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Introduction to Animal Behavior Why do they do that?

Introduction to Animal Behavior Why do they do that?

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Page 1: Introduction to Animal Behavior Why do they do that?

Introduction to Animal Behavior

Why do they do that

Warm Up

Personification Putting human characteristics onto animals to explain behavior

Youtube httpwwwyoutubecomwatchv=f-Kt_kuYVtU

Unit Map Follow Along in your packet

WHAT ARE YOU LEARNING Explain factors that serve to stimulate or

discourage various types of animal behaviorRecognize the normality curve of animal

behavior

Know Understand Do

KnowTypes of

BehaviorHistory of

studying animal behavior

Understandbull Stimuli

influence on behavior

bull How ethology has evolved

Dobull Observe and

interpret animal behaviors

bull Evaluate animal studies

Key Learning Animal Behavior

Unit EQ How does animal behavior influence animal husbandry

Concept Influence

Lesson EQWhat can alter behavior

VocabAgnostic

Concept History

Lesson EQHow are animals studied for behavior

VocabFAP Skinner Box

Concept TypesLesson EQ

What are categories of behavior

VocabStimuli Behavior Ethology

What is Animal Behavior

The study of how and why animals interact with each other (both within and among species) and their environment

Proximate questions - howmechanisms responsible for interactions

Ultimate questions - whyhow these interactions influence an

individuals survival and reproduction

Some examples

Intraspecific interactions

mate choice

male competition

alarm calls

parental care

Some examples

Interspecific interactions

predation

parasitism

mutualism

competition

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 2: Introduction to Animal Behavior Why do they do that?

Warm Up

Personification Putting human characteristics onto animals to explain behavior

Youtube httpwwwyoutubecomwatchv=f-Kt_kuYVtU

Unit Map Follow Along in your packet

WHAT ARE YOU LEARNING Explain factors that serve to stimulate or

discourage various types of animal behaviorRecognize the normality curve of animal

behavior

Know Understand Do

KnowTypes of

BehaviorHistory of

studying animal behavior

Understandbull Stimuli

influence on behavior

bull How ethology has evolved

Dobull Observe and

interpret animal behaviors

bull Evaluate animal studies

Key Learning Animal Behavior

Unit EQ How does animal behavior influence animal husbandry

Concept Influence

Lesson EQWhat can alter behavior

VocabAgnostic

Concept History

Lesson EQHow are animals studied for behavior

VocabFAP Skinner Box

Concept TypesLesson EQ

What are categories of behavior

VocabStimuli Behavior Ethology

What is Animal Behavior

The study of how and why animals interact with each other (both within and among species) and their environment

Proximate questions - howmechanisms responsible for interactions

Ultimate questions - whyhow these interactions influence an

individuals survival and reproduction

Some examples

Intraspecific interactions

mate choice

male competition

alarm calls

parental care

Some examples

Interspecific interactions

predation

parasitism

mutualism

competition

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 3: Introduction to Animal Behavior Why do they do that?

Unit Map Follow Along in your packet

WHAT ARE YOU LEARNING Explain factors that serve to stimulate or

discourage various types of animal behaviorRecognize the normality curve of animal

behavior

Know Understand Do

KnowTypes of

BehaviorHistory of

studying animal behavior

Understandbull Stimuli

influence on behavior

bull How ethology has evolved

Dobull Observe and

interpret animal behaviors

bull Evaluate animal studies

Key Learning Animal Behavior

Unit EQ How does animal behavior influence animal husbandry

Concept Influence

Lesson EQWhat can alter behavior

VocabAgnostic

Concept History

Lesson EQHow are animals studied for behavior

VocabFAP Skinner Box

Concept TypesLesson EQ

What are categories of behavior

VocabStimuli Behavior Ethology

What is Animal Behavior

The study of how and why animals interact with each other (both within and among species) and their environment

Proximate questions - howmechanisms responsible for interactions

Ultimate questions - whyhow these interactions influence an

individuals survival and reproduction

Some examples

Intraspecific interactions

mate choice

male competition

alarm calls

parental care

Some examples

Interspecific interactions

predation

parasitism

mutualism

competition

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 4: Introduction to Animal Behavior Why do they do that?

Know Understand Do

KnowTypes of

BehaviorHistory of

studying animal behavior

Understandbull Stimuli

influence on behavior

bull How ethology has evolved

Dobull Observe and

interpret animal behaviors

bull Evaluate animal studies

Key Learning Animal Behavior

Unit EQ How does animal behavior influence animal husbandry

Concept Influence

Lesson EQWhat can alter behavior

VocabAgnostic

Concept History

Lesson EQHow are animals studied for behavior

VocabFAP Skinner Box

Concept TypesLesson EQ

What are categories of behavior

VocabStimuli Behavior Ethology

What is Animal Behavior

The study of how and why animals interact with each other (both within and among species) and their environment

Proximate questions - howmechanisms responsible for interactions

Ultimate questions - whyhow these interactions influence an

individuals survival and reproduction

Some examples

Intraspecific interactions

mate choice

male competition

alarm calls

parental care

Some examples

Interspecific interactions

predation

parasitism

mutualism

competition

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 5: Introduction to Animal Behavior Why do they do that?

Key Learning Animal Behavior

Unit EQ How does animal behavior influence animal husbandry

Concept Influence

Lesson EQWhat can alter behavior

VocabAgnostic

Concept History

Lesson EQHow are animals studied for behavior

VocabFAP Skinner Box

Concept TypesLesson EQ

What are categories of behavior

VocabStimuli Behavior Ethology

What is Animal Behavior

The study of how and why animals interact with each other (both within and among species) and their environment

Proximate questions - howmechanisms responsible for interactions

Ultimate questions - whyhow these interactions influence an

individuals survival and reproduction

Some examples

Intraspecific interactions

mate choice

male competition

alarm calls

parental care

Some examples

Interspecific interactions

predation

parasitism

mutualism

competition

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 6: Introduction to Animal Behavior Why do they do that?

What is Animal Behavior

The study of how and why animals interact with each other (both within and among species) and their environment

Proximate questions - howmechanisms responsible for interactions

Ultimate questions - whyhow these interactions influence an

individuals survival and reproduction

Some examples

Intraspecific interactions

mate choice

male competition

alarm calls

parental care

Some examples

Interspecific interactions

predation

parasitism

mutualism

competition

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 7: Introduction to Animal Behavior Why do they do that?

Some examples

Intraspecific interactions

mate choice

male competition

alarm calls

parental care

Some examples

Interspecific interactions

predation

parasitism

mutualism

competition

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 8: Introduction to Animal Behavior Why do they do that?

Some examples

Interspecific interactions

predation

parasitism

mutualism

competition

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 9: Introduction to Animal Behavior Why do they do that?

Some examples

Interactions with the environment

foraging

nest site selection

signal modification

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 10: Introduction to Animal Behavior Why do they do that?

Why study behavior

Possible first science Our survival dependent on knowledge of other animals (preycompetitorspredators)

Controlmanagement of species Food and game species agricultural pests invasive species endangered species

Understandingmodification of our own behavior Studies of how birds learn and develop songs provide unique insights into the development and neural control of speech in humans

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 11: Introduction to Animal Behavior Why do they do that?

Curiosity

Science for sciencersquos sake

Achieve a better understanding of the species we share the Earth with

Almost any behavior performed by any animal may be interesting to study

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 12: Introduction to Animal Behavior Why do they do that?

Paleolithic art from 40000+ years ago provide indirect evidence that primitive humans observed the behavior of animals

Cave paintings portray herding animals in groups animal migration certain predators hunting in packs and solitary animals alone

History of the study of animal behavior

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 13: Introduction to Animal Behavior Why do they do that?

- Discriminated data from theory

- Developed hypotheses

- Used reasoned skepticism

Blurton-Jones (1976) documented Kalahari bushmenrsquos (Kung) knowledge of animal behavior

Hunter-gatherer society similar to most of humanrsquos history

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 14: Introduction to Animal Behavior Why do they do that?

Introduction

Why do animals do what they do Why do birds sing How do sea turtles navigate

the ocean to lay their eggs on the same beach where they were hatched

How do honeybees know when the hive needs more food

Image from httpwwwnpsgov

Image from httpwwwscottcamazinecom

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 15: Introduction to Animal Behavior Why do they do that?

Introduction

Animal behavior asks what why and how

Animal behavior is also referred to as ethology Scientists who study

animal behavior are called ethologists

Image from httpwwwarcamaxcom

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 16: Introduction to Animal Behavior Why do they do that?

Introduction

Animal behavior is centered around the ability to moveAnimals seek

food water shelter

Animals play with each other

Animals seek matesImage from httpwwwe-magineeducationtasgovau

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 17: Introduction to Animal Behavior Why do they do that?

IntroductionIn order for an animal to move it uses muscles So in a way we can think of animal behavior as

being dependent on muscle movement

Image from httpwwwwildaboutcatsorg

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 18: Introduction to Animal Behavior Why do they do that?

Introduction

Behavior results as a reaction to a stimulusA stimulus is a detectable

change in the animalrsquos internal or external environment

HungerSoundPainVisual cuesHormonal changes

Image from httpwww3nauedubiology

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 19: Introduction to Animal Behavior Why do they do that?

What is really happeninghellip

Lets watch

As we watch when the video pauses explain WHAT the animal is doing and WHY you think the animal is doing this

youtube httpwwwyoutubecomwatchv=LU8DDYz68kM

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 20: Introduction to Animal Behavior Why do they do that?

Introduction

Ethologists do not attempt to describe WHY an animal does a behavior before describing WHAT the animal is doingThis removes as much bias as possible ndash good

scientists donrsquot want to just ldquosee what they want to seerdquo

Need to make objective observations of animal behaviors analyze the data statistically then come to conclusions about WHY an animal behaves a certain way

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 21: Introduction to Animal Behavior Why do they do that?

Introduction

For example you see two gophers interacting with each other rolling and hopping around running to and from each other As a behavioral ecologist you would first state

the behavior you are observing Once you made the observations about WHAT

was happening you could begin to determine WHY they are behaving that way

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 22: Introduction to Animal Behavior Why do they do that?

Introduction

The behavior you observed could have been many different things Play Mating rituals Aggression

defending territory

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 23: Introduction to Animal Behavior Why do they do that?

Movie BreakPlease get a piece of paper and copy the

followingANSWER FOR EACH ANIMAL PRESENTED

1 What animal2 Where is it commonly found3 What is the ldquosmartrdquo behavior4 Is this behavior (in your opinion) instinctual

learned or other

AT THE END OF THE VIDEOWhat animal do YOU believe is the smartest and

why (10 sentences)

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 24: Introduction to Animal Behavior Why do they do that?

Video Worldrsquos Smartest Animals

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 25: Introduction to Animal Behavior Why do they do that?

Types of behaviors defined

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 26: Introduction to Animal Behavior Why do they do that?

Pet Activity Warm Up On a separate sheet of paper1 Write the name of one of your pets Is

it a bird dog or cat Other2 What behaviors do you think about

when you think of your pet Give a list of behaviors Indicate if the behavior was genetic ldquoinnaterdquo or learned

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 27: Introduction to Animal Behavior Why do they do that?

Behavioral Ecology

Behavioral ecology emphasizes evolutionary hypothesis science as a process

Based on expectation that animals behave in ways that will increase their Darwinian fitness (reproductive success)

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 28: Introduction to Animal Behavior Why do they do that?

Stimuli Review

Certain stimuli trigger innate behaviors called fixed action patterns

A fixed action pattern (FAP) is a highly stereotypical innate behavior that continues to completion after initiation by an external stimulus

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 29: Introduction to Animal Behavior Why do they do that?

Learning

Learning is experience based modification of behavior

Some learning is due mostly to inherent maturation

Habituation is learning involving loss of sensitivity to unimportant stimuli

Associative learning involves linking one stimulus with another

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 30: Introduction to Animal Behavior Why do they do that?

Classical conditioning (Pavlov)

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 31: Introduction to Animal Behavior Why do they do that?

Operant conditioning (Skinner)

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 32: Introduction to Animal Behavior Why do they do that?

Operant Conditioning1048708 Animal learns to behave in a certain waythrough repeated practice1048708 Trial amp error learning ndash animal testsconditions for desired responseeg Skinner box1048708 Animal learns that a behavior gets a certainresponse1048708 eg rat presses lever gets food

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 33: Introduction to Animal Behavior Why do they do that?

Rhythmic Behaviors

Rhythmic behaviors synchronize an animalrsquos activities with daily and seasonal changes in the environment

Governed by endogenous clocks which in turn require exogenous cues to keep the behavior properly timed with the external environment

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 34: Introduction to Animal Behavior Why do they do that?

Foraging Behavior

Ecologists are using costbenefit analysis to study foraging behavior

Species may be generalists or specialists as foragers

Animals modify behavior to favor a high ratio of energy intake to expenditure

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 35: Introduction to Animal Behavior Why do they do that?

Social Behavior

Sociobiology places social behavior in evolutionary context

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 36: Introduction to Animal Behavior Why do they do that?

Competitive Social Behaviors

Agonistic behavior competitor gains advantage by getting a limited resource like food or a mate

Natural selection survival of the fittestldquoPecking orderrdquo dominance hierarchies

with differently ranked individuals permitted options according to their status

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 37: Introduction to Animal Behavior Why do they do that?

How natural selections leads to behavioral traitsVariation exists fraction of the species T

elegans (garter snakes) had ability to recognize slugs by chemoreception

Increased fitness That variation has higher chance to survive and reproduce (genes passed on)

Led to changes in the population over time

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 38: Introduction to Animal Behavior Why do they do that?

Mating Behavior

Promiscuity ndash having many random mates

Monogamy ndash having only one matePolygamy ndash having a few selected mates

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 39: Introduction to Animal Behavior Why do they do that?

Mating behavior

Promiscuous Strong bonds

Monogamous(sex morphology similar)

Polygamous

Polyandry(dimorphic Larger Showy males)

Polygyny(dimorphicLarger Showy females)

Factors influencing evolution of mating systems-Need of young-Paternity certainty

- certainty increases with external fertilization

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 40: Introduction to Animal Behavior Why do they do that?

Sexual selectionSexual selection (selective pressure)

evolution of male behavior and anatomyStalked-eyed flies

Females more likely to mate with males with longer eyestalks

Why Correlation between genetic disorders and inability to develop long eyestalks

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 41: Introduction to Animal Behavior Why do they do that?

Social Interactions

Social interactions depend on diverse modes of communication

Some animals communicate with smells

Honeybees communicate through ldquodancingrdquo

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 42: Introduction to Animal Behavior Why do they do that?

Social learning

Experience involves observing othersCulture information transfer through

social learningVervet monkey alarm callsMemes (Richard Dawkins)

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 43: Introduction to Animal Behavior Why do they do that?

Altruistic Behavior

Inclusive fitness accounts for most altruistic behavior

Best explained by a ldquokinrdquo theory animals try to maintain the survival of others who share their genes

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 44: Introduction to Animal Behavior Why do they do that?

AltruismCostbenefit of selfish vs unselfish

behaviorAltruism reduces individual fitness but

increases fitness of others

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 45: Introduction to Animal Behavior Why do they do that?

Reciprocal altruism

Some animals behave altruistically toward others who are not relatives A wolf may offer food to another wolf even though they share no kinship

Such behavior can be adaptive if the aided individual returns the favor in the future

This sort of exchange of aid is called reciprocal altruism

Commonly used to explain altruism in humans

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 46: Introduction to Animal Behavior Why do they do that?

Agonistic behavior

RitualizedWinner gains access to resourcesPhysical and behavioral characteristics

involvedUsually harm is not done

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 47: Introduction to Animal Behavior Why do they do that?

Reasoning

Analyze problem amp devise solution using past experiences

Most DogsEg No canrsquot unwind leash from tree

Most HorsesNo

PrimatesYES

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 48: Introduction to Animal Behavior Why do they do that?

Sociobiology

Human sociobiology connects biology to the humanities and social sciences

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 49: Introduction to Animal Behavior Why do they do that?

Sociobiology (EO Wilson)

Connects human culture to evolutionary theory

Social behaviors exist because they are perpetuated by natural selection

Does not mean all social behaviors are hardwired (nature vs nurture)

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 50: Introduction to Animal Behavior Why do they do that?

Self-quiz

Bees can see colors we cannot see and detect minute amounts of chemicals we cannot smell But unlike many insects bees cannot hear very well Which of the following statements best fits into the perspective of behavioral ecology

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 51: Introduction to Animal Behavior Why do they do that?

Possible answers

A Bees are too small to have functional ears

B Hearing must not contribute much to a beersquos fitness

C If a bee could hear its tiny brain would be swamped with information

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 52: Introduction to Animal Behavior Why do they do that?

Possible answers

D This is an example of a fixed action pattern

E If bees could hear the noise of the hive would distract the bees from their work

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 53: Introduction to Animal Behavior Why do they do that?

Challenge question

Starting with the very first time a bee leaves the hive it always flies in a circle around the hive before heading out on a foraging trip

If it is prevented from seeing the hive when it leaves or if the hive is moved while the bee is gone the bee is not able to locate the hive when it returns

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 54: Introduction to Animal Behavior Why do they do that?

Challenge Question

For this reason beekeepers know that a hive should only be moved helliphellipwhen Why

What part of the bees ldquoorientation flightrdquo behavior appears to be innate

What component shows learning

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 55: Introduction to Animal Behavior Why do they do that?

Temple Grandin Who is she What is her ldquodisabilityrdquo What was her ldquomajor break throughrdquo at her Auntrsquos cattle farm How does this break through help Temple through college What problems arise at college with her break through How does she over come these problems What does Temple realize when she first visits the feed lot How does she go about her research Why is her job at the newspaper critical for Temple What does Temple design for the gentleman who comes to the

paper What happens the 1st time they use her design Why does this happen How does Temple plan on ldquofixing the industryrdquo What ldquobreak throughrdquo does Temple have at the funeral What break through does Temple have at the grocery store How

does this play a role in her slaughter house design What does Temple do at the conference How does her disability help her ldquoseerdquo

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 56: Introduction to Animal Behavior Why do they do that?

Review Types of

BehaviorSocialCompetitive

Agnostic Natural Selection ldquoPecking Orderrdquo

ForagingAltruistic Rhythmic

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 57: Introduction to Animal Behavior Why do they do that?

Behavioral biology is the study of what animals do when interacting with their environment

Behavior can be interpreted in terms of proximate causes (immediate interaction with the environment) or ultimate causes (evolutionary differences)

Behavioral biologists study the actions of animals in their natural environments

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 58: Introduction to Animal Behavior Why do they do that?

Early insights into the nature of behavior came from studies by Nobel laureates Karl von Frisch Konrad Lorenz and Niko Tinbergen

They were among the first experimentalists in behavioral biology

Tinbergen and Lorenz performed experimental studies of innate behavior and simple forms of learning

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 59: Introduction to Animal Behavior Why do they do that?

Behavioral ecologists are especially interested in the ultimate causes of behavior which are evolutionary Natural selection preserves behaviors that enhance fitness

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 60: Introduction to Animal Behavior Why do they do that?

Niko Tinbergen

Niko Tibergen was a pioneer in the field of animal behaviorHe observed animals in

their natural conditions then manipulated or varied the conditions to see how the animals responded

Image from httpnobelprizeorg

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 61: Introduction to Animal Behavior Why do they do that?

A classic Tinbergen experiment deals with the nesting behavior of the digger wasp

The female wasp often excavates and cares for four or five separate nests

Tinbergen used this experiment to test his prediction that digger wasps use landmarks to keep track of the location of their nests

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 62: Introduction to Animal Behavior Why do they do that?

Niko Tinbergen

Tinbergen observed how a wasp called the beewolf finds its nest among other beewolf nestsHe observed that the beewolf

would circle its nest in an ever-widening circle before flying away to hunt

This behavior was an action pattern ndash it was performed exactly the same way each time

Image from httpwwwsciencenewsorg

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 63: Introduction to Animal Behavior Why do they do that?

Niko Tinbergen

After the beewolf flew off Tinbergen would move certain landmarks around the nests

When the beewolf returned it was disorientedSo by manipulating the

beewolfrsquos environment Tinbergen came to the conclusion that the beewolf commits landmarks to memory to be able to find its nest when it comes back from hunting

Image from httpwwwearthlifeorg

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 64: Introduction to Animal Behavior Why do they do that?

In the experiment Tinbergen placed a circle of pinecones around a nest opening

Figure 371 Part 11

Nest

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 65: Introduction to Animal Behavior Why do they do that?

Nest

After the female flew away Tinbergen moved the pinecones a few feet to one side of the nest opening

When the female wasp returned she flew to the middle of the circle of pinecones rather than to the actual nest opening

Figure 371 Part 22

No Nest

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 66: Introduction to Animal Behavior Why do they do that?

Tinbergen next arranged the pinecones in a triangle around the nest and made a circle of small stones off to one side of the nest opening

This time the wasp flew to the stones

Nest

Figure 371 Part 33

No Nest

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 67: Introduction to Animal Behavior Why do they do that?

The wasp cued in on the arrangement of the landmarks rather than the landmarks themselves

This experiment demonstrated that the wasp did use landmarks and that she could learn new ones to keep track of her nest

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 68: Introduction to Animal Behavior Why do they do that?

Niko Tinbergen

Tinbergen had to describe and investigate WHAT the organism was doing before attempting to explain WHY

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 69: Introduction to Animal Behavior Why do they do that?

Spatial learning and cognitive maps

Spatial learning (Tinbergen) experience consists of spatial structures of the environmentUse of landmarks

ReliableCognitive maps Internal

representation of spatial relationships

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 70: Introduction to Animal Behavior Why do they do that?

Recap

WhatWhat was the experiment

WhereWhere did it take place

WhenWhen was the behavior altered

WhyWhy was this a break through in the study of animal

behavior How

How would this discovery impact (for example) a bee keeper

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 71: Introduction to Animal Behavior Why do they do that?

Guidelines For Studying Animal Behavior

Ask clear specific questionsKeep the question simple Are you sure it is a

question that can be easily answeredPut the question into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesisDecide on the type of data you need to collect and

how you will gather the dataLeave time to run statistical analyses on the data

and form conclusions based on your results

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 72: Introduction to Animal Behavior Why do they do that?

Check your understandinghellip

Draw a cartoon of the Niko Wasp Experiment

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 73: Introduction to Animal Behavior Why do they do that?

Animal Behavior

Influences and Actions

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 74: Introduction to Animal Behavior Why do they do that?

Animal behavior often involves a combination of genetic programming (innate behavior) and environmental experiences (learning)both genes and the environment influence the

development of behavioral phenotypes- just like any other traits

372 Behavior results from both genes and environmental factors

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 75: Introduction to Animal Behavior Why do they do that?

What influences behavior

Environmental pressuresInternal influences such as hormonesLearningGenetic predisposition (sometimes

referred to as ldquoinstinctrdquo)

There isnrsquot a definite combination of these influences that affects all behavior

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 76: Introduction to Animal Behavior Why do they do that?

Proximate vs Ultimate Causes

The question of ldquoWHYrdquo can have different answersProximate causes are related to internal changes in the

animalHormonesMessages from the nervous system

ldquoProximaterdquo means closeUltimate causes are related to the survival and

reproductive success of the animal ldquoUltimaterdquo means furthest or utmost

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 77: Introduction to Animal Behavior Why do they do that?

The gathering of nest materials by lovebirds has genetic and environmental components

Figure 372

Single long strip carried in beak(Fischerrsquos lovebird)

Several short strips tucked under feathers(peach-faced lovebird)

Hybrid behavior

Tuckingfailure

Strip inbeak

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 78: Introduction to Animal Behavior Why do they do that?

Proximate vs Ultimate Causes

An example incorporating both proximate and ultimate causes Beldingrsquos ground squirrelsWhen males reach about two

months old they leave the burrow where they were bornIt is an increase in testosterone or

a hormonal change that triggers this behavior

So the proximate cause of the nest-leaving behavior involves the increase in testosterone levels in the squirrel

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 79: Introduction to Animal Behavior Why do they do that?

Proximate vs Ultimate Causes

There is more to the story than just hormones When males leave the nest they avoid

inbreeding with sisters or cousins etc Their offspring are therefore healthier The male offspring inherit the same genetic

information that induces them to leave their nests at a young age

So this behavior is passed on genetically and it makes for a healthier population of squirrels

Avoiding inbreeding is therefore the ultimate cause of this ldquoearly nest-leavingrdquo behavior

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 80: Introduction to Animal Behavior Why do they do that?

How to determine WHY ndash action patterns

Action patterns are complex behaviors that are always repeated the same way by a species of animal We say that action patterns are stereotyped

since they occur the same way each time and through to completion

After repeatedly observing action patterns an ethologist can analyze the data statistically

Only then do we attempt to determine WHY a behavior is being done

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 81: Introduction to Animal Behavior Why do they do that?

Sign stimuli (often a simple cue in an animalrsquos environment) trigger innate essentially unchangeable fixed action patterns (FAPs)

The genetic programming underlying FAPs ensures that such activities are performed correctly without practice

373 Innate behavior often appears as fixed action patterns

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 82: Introduction to Animal Behavior Why do they do that?

Fixed action pattern (FAP)

Sequence of unlearned behaviors

Nearly unchangeableCarried out to completionSign stimulus (releaser)

behaviorExample of an innate

behavior

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 83: Introduction to Animal Behavior Why do they do that?

More on Action Patterns

The egg-rolling behavior of the greylag goose is a good example of an action pattern

Niko Tibergen and another pioneer in ethology Konrad Lorentz originally observed this behavior

Image from httpwwwgrayimagescouk

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 84: Introduction to Animal Behavior Why do they do that?

More on Action Patterns

The goose will roll an egg that is outside the nest back into the nest in the same manner every timeInterestingly the goose will

do this with any round object placed outside the nest

Every time this action pattern is initiated it is carried through to completion

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 85: Introduction to Animal Behavior Why do they do that?

The graylag goose always retrieves an egg that has been bumped out of her nest in the same manner

This is a fixed action pattern

She carries this sequence to completion even if the egg slips away during the process

Figure 373A

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 86: Introduction to Animal Behavior Why do they do that?

Several key events in the life cycle of the European cuckoo are determined by fixed action patterns

Egg-laying behavior

Figure 373B

1 2 3

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 87: Introduction to Animal Behavior Why do they do that?

The behavior of the cuckoo hatchling ejecting the host eggs from the nest

The feeding behavior of a foster mother to the cuckoo chick

Figure 373B

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 88: Introduction to Animal Behavior Why do they do that?

Learning is a change in behavior resulting from experience

Habituation is one of the simplest forms of learningAn animal learns not to respond to a repeated

stimulus that conveys little or no informationFor example birds eventually become

habituated to scarecrows and no longer avoid nearby fruit trees

Learning ranges from simple behavioral changes to complex problem solving

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 89: Introduction to Animal Behavior Why do they do that?

Table 374

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 90: Introduction to Animal Behavior Why do they do that?

Imprinting is learning that involves both innate behavior and experience

Imprinting is irreversible learning limited to a sensitive period in an animals life it enhances fitness by enabling rapid learning

Example Lorenz used the graylag goose to demonstrate imprinting He took over the maternal role for a group of goslings

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 91: Introduction to Animal Behavior Why do they do that?

Not all examples of imprinting involve parent-offspring bonding

Although newly hatched salmon do not receive any parental care they imprint on the complex mixture of odors unique to the freshwater stream where they hatch

This allows salmon to find their way back to the stream to spawn after spending a year or more at sea

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 92: Introduction to Animal Behavior Why do they do that?

Imprinting plays an important role in song development for many kinds of birds

Figure 375B

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 93: Introduction to Animal Behavior Why do they do that?

Associative learning is learning that a particular stimulus or response is linked to a reward or punishmentThese ducks have

learned to associate humans with food handouts

They congregate rapidly whenever a person approaches the shoreline

Many animals learn by association and imitation

Figure 376A

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 94: Introduction to Animal Behavior Why do they do that?

Trial-and-error learning is a common form of associative learning

An animal learns to associate one of its own behavioral acts with a positive or negative effect

Figure 376B

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 95: Introduction to Animal Behavior Why do they do that?

Imitation is learning by observing and mimicking the behavior of othersThis form of learning is not limited to a sensitive

periodMany predators including cats and coyotes seem

to learn some of their basic hunting tactics by observing and imitating their mother

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 96: Introduction to Animal Behavior Why do they do that?

Some animals exhibit problem-solving behaviorExamples chimpanzees and

ravens

Animal cognition includes problem-solving behavior

Figure 377A B

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 97: Introduction to Animal Behavior Why do they do that?

Animal Behavior

ECOLOGICAL ROLES OF BEHAVIOR

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 98: Introduction to Animal Behavior Why do they do that?

Behavior is an evolutionary adaptation that enhances survival and reproductive success

Behavior evolves as natural selection fine-tunes an animal to its environmentThe hunting and reproduction behaviors of

jaguarsNest location by digger waspsImprinting of goslings

378 An animals behavior reflects its evolution

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 99: Introduction to Animal Behavior Why do they do that?

Animals exhibit a great variety of rhythmic behavior patterns

Circadian rhythms are patterns that are repeated dailySleepwake cycles in animals and plants

Circadian rhythms appear to be timed by an internal biological clock

379 Biological rhythms synchronize behavior with the environment

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 100: Introduction to Animal Behavior Why do they do that?

In the absence of environmental cues these rhythms continueBut they become out of phase with the

environment

Figure 379A

1212 (natural)Constant darkness

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 101: Introduction to Animal Behavior Why do they do that?

Movement in a directed way enables animals to avoid predatorsmigrate to a more favorable environmentobtain foodfind mates and nest sites

3710 Animal movement may be oriented to stimuli or landmarks

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 102: Introduction to Animal Behavior Why do they do that?

1 kinesis- simplest type of animal movement - random movement in response to a stimulus

2 taxis- another simple type A more or less automatic movement directed toward

or away from some stimulus Examples include rheotaxis (current) chemotaxis

and phototaxis

3 Some animals use landmarks to find their way within an area

TYPES of ANIMAL MOVEMENT

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 103: Introduction to Animal Behavior Why do they do that?

Directed movements

Strong genetic influenceKinesis versus taxisMigration

Migrating blackcaps kept in captivity exhibited behaviors of ldquomigratory restlessnessrdquo at night

Migratory and nonmigratory blackcaps mated and subjected to both environments

40 of offspring exhibited ldquomigratory restlessnessrdquo

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 104: Introduction to Animal Behavior Why do they do that?

Many animals formulate cognitive maps Internal representations of spatial relationships among

objects in their surroundings (wasp example)

Some animals undertake long-range migrationsExamples whales sea turtles birds monarch butterflies

Animals navigate using the sun stars temperature gradients landmarks or Earths magnetism

3711 Movement from place to place often depends on internal maps

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 105: Introduction to Animal Behavior Why do they do that?

Migrating gray whales use coastal landmarks to stay on course

Figure 3711A

FEEDINGGROUNDS

Siberia

Arctic Ocean

Alaska

NORTHAMERICA

PacificOcean

Baja California

BREEDING GROUNDS

AtlanticOcean

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 106: Introduction to Animal Behavior Why do they do that?

The indigo bunting learns a star map and navigates by fixing on the North Star

Figure 3711B

Paper

Ink padFunnel-shapedcage

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 107: Introduction to Animal Behavior Why do they do that?

Letrsquos Read More About ItRead the article on Indigo BuntingsAnswer the questions that follow

Put the experiment into the ldquoWhat is the effect of _______ on ________rdquo model

Formulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)What are the resultsHow can this influence studies of other migratory birdsHow can human behavior alter this natural behavior in

birds

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 108: Introduction to Animal Behavior Why do they do that?

Bird Migration Video

For each bird What is the speciesHow far does it travelTry to get an idea of where 2 birds mentioned in

the video are from Afterwards we will try to locate possible modes

of remembrance during migration Landmarks Stars Earthrsquos Magnetism Other

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 109: Introduction to Animal Behavior Why do they do that?

Animals are generally selective and efficient in their food choicesSome animals such

as gulls are feeding ldquogeneralistsrdquo

Other animals such as koalas are feeding ldquospecialistsrdquo

3712 Behavioral ecologists use costbenefit analysis in studying feeding behavior

Figure 3712A B

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 110: Introduction to Animal Behavior Why do they do that?

The mechanism that enables an animal to find particular foods efficiently is called a search image

Natural selection seems to have shaped feeding behavior to maximize energy gain and minimize the expenditure of time and energyThis is the theory of optimal foraging

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 111: Introduction to Animal Behavior Why do they do that?

Foraging behavior

Optimal foraging theory behaviors exist as a compromise between benefits of nutrition and cost of obtaining food

Predation must be a factor

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 112: Introduction to Animal Behavior Why do they do that?

Whenever an animal has food choices there are a number of tradeoffs

A bass can get more usable energy from minnows but crayfish are easier to catch

However it may take more time to eat a crayfish because of its tough exoskeleton

Figure 3712C

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 113: Introduction to Animal Behavior Why do they do that?

The kangaroo rat selects high-energy foods (seeds) in a manner that reduces time spent above the ground where it is exposed to predators

Figure 3712E

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 114: Introduction to Animal Behavior Why do they do that?

Read more about it

Read the Indigo Bunting and answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow can this influence the feed industry for cattle

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 115: Introduction to Animal Behavior Why do they do that?

Animal Behavior

SOCIAL BEHAVIOR AND SOCIOBIOLOGY

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 116: Introduction to Animal Behavior Why do they do that?

Social behavior is defined as the interaction among members of a population

The discipline of sociobiology studies social behavior in the context of evolution

3713 Sociobiology places social behavior in an evolutionary context

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 117: Introduction to Animal Behavior Why do they do that?

Agonistic behavior is social behavior consisting of threats and combat that settles disputes between individuals in a population

Agonistic behavior can directly affect an individuals evolutionary fitness The victor often gains

first or exclusive access to mates

3714 Rituals involving agonistic behavior often resolve confrontations between competitors

Figure 3714

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 118: Introduction to Animal Behavior Why do they do that?

Many animals live in social groups maintained by agonistic behaviors

Dominance hierarchy is the ranking of individuals based on social interactions

3715 Dominance hierarchies are maintained by agonistic behavior

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 119: Introduction to Animal Behavior Why do they do that?

Chickens establish a ldquopeck orderrdquo

Resources are often partitioned based upon the dominance hierarchy

Figure 3715

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 120: Introduction to Animal Behavior Why do they do that?

Humans tend to space themselves out when they are close to othersThey establish

what we might call personal territories

3717 Territorial behavior parcels space and resources

Figure 3717A

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 121: Introduction to Animal Behavior Why do they do that?

Many animals exhibit territorial behavior

It is a form of social behavior that partitions resources

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 122: Introduction to Animal Behavior Why do they do that?

A territory is an area that individuals defend and from which other members of the same species are usually excluded

The size of the territory varies with species the function and the available resources

Territories are typically used for feeding mating andor rearing young

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 123: Introduction to Animal Behavior Why do they do that?

Territoriality is often maintained by agonistic behavior

These New Zealand gannets maintain their individual nesting territories by calling and pecking at each other

Figure 3717B

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 124: Introduction to Animal Behavior Why do they do that?

Territoriality can enhance fitness if the benefits of possessing a territory outweigh the energy costs of defending one

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 125: Introduction to Animal Behavior Why do they do that?

Territorial rights are proclaimed continually in a variety of ways

Bird songsNoises such as the bellowing of sea lions and

the chattering of squirrelsDefecation in open

areasScent markers

such as urine

Figure 3717C

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 126: Introduction to Animal Behavior Why do they do that?

Signals and communication

Signal causes change in another organismrsquos behavior

Difference between communication and language

Pheromones (reproductive and nonreproductive behaviors)

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 127: Introduction to Animal Behavior Why do they do that?

Auditory communication

Songs of birds are partly learnedCritical period

Some insects such as male Drosophila produce a song even when reared in isolationVery little variation why

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 128: Introduction to Animal Behavior Why do they do that?

Zebra Danio Experiment

Read the Article and Answer the followingPut the experiment into the ldquoWhat is the effect of

_______ on ________rdquo modelFormulate a complete hypothesis in the IFTHAN formatWhat is the type of data collectedHow is the data gatheredWhat is the timing for the experiment (How long did

they run)How many trials were neededWhat are the resultsHow could this influence the sale of multiple Danio fish

to someone in a pet store

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment

Page 129: Introduction to Animal Behavior Why do they do that?

Animal Behavior REVIEWDefine ethology animal behavior stimulus action pattern

proximate cause (give example) ultimate cause (give example) FAP (give example) agnostic behavior rhythmic behavior Circadian Rhythm sociobiology

Answer in complete sentences What influences behavior Know the types of behavior and be able to explaingive examples of each What are the 5 types of learning Explain Nikorsquos experiment (you can use drawings for support) What did this tell us about

animal behavior Explain the indigo bunting experiment What did this tell us about animal behavior What was the Skinner box What type of behavior does this illustrate What was Pavlovrsquos experiment and what type of behavior does this illustrate Why was

this an important gain in animal behavior Who was Lorenz and what did he study What did this tell us about animal behavior How was the winged Migration movie successful What research did they use as a base

for their experiment