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Mutualism (biology)From Wikipedia, the free encyclopedia

Hummingbird Hawkmoth drinking fromDianthus. Pollination is a classic example of mutualism.

Arbuscular mycorrhiza

Mutualism is the way two organismsbiologically interact where each individual derives afitnessbenefit (i.e.

increased reproductive output). Similar interactions within a species are known asco-operation. It can be

contrasted withinterspecific competition, in which each species experiencesreducedfitness, and exploitation,

orparasitism, in which one species benefits at the expense of the other. Mutualism and symbiosisare

sometimes used as if they are synonymous, but this is strictly incorrect: symbiosis is a broad category, defined

to include relationships which are mutualistic, parasiticorcommensal. Mutualism is only one type.

http://en.wikipedia.org/wiki/Hummingbird_Hawkmothhttp://en.wikipedia.org/wiki/Dianthushttp://en.wikipedia.org/wiki/Pollinationhttp://en.wikipedia.org/wiki/Biological_interactionhttp://en.wikipedia.org/wiki/Biological_interactionhttp://en.wikipedia.org/wiki/Fitness_(biology)http://en.wikipedia.org/wiki/Fitness_(biology)http://en.wikipedia.org/wiki/Co-operation_(evolution)http://en.wikipedia.org/wiki/Co-operation_(evolution)http://en.wikipedia.org/wiki/Co-operation_(evolution)http://en.wikipedia.org/wiki/Interspecific_competitionhttp://en.wikipedia.org/wiki/Interspecific_competitionhttp://en.wikipedia.org/wiki/Cheating_(biology)http://en.wikipedia.org/wiki/Parasitismhttp://en.wikipedia.org/wiki/Parasitismhttp://en.wikipedia.org/wiki/Symbiosishttp://en.wikipedia.org/wiki/Symbiosishttp://en.wikipedia.org/wiki/Parasitichttp://en.wikipedia.org/wiki/Parasitichttp://en.wikipedia.org/wiki/Commensalhttp://en.wikipedia.org/wiki/Commensalhttp://en.wikipedia.org/wiki/File:Arbuscular_mycorrhiza_cross-section.pnghttp://en.wikipedia.org/wiki/File:Arbuscular_mycorrhiza_cross-section.pnghttp://en.wikipedia.org/wiki/File:Hummingbird_hawkmoth_a.jpghttp://en.wikipedia.org/wiki/File:Hummingbird_hawkmoth_a.jpghttp://en.wikipedia.org/wiki/Dianthushttp://en.wikipedia.org/wiki/Pollinationhttp://en.wikipedia.org/wiki/Biological_interactionhttp://en.wikipedia.org/wiki/Fitness_(biology)http://en.wikipedia.org/wiki/Co-operation_(evolution)http://en.wikipedia.org/wiki/Interspecific_competitionhttp://en.wikipedia.org/wiki/Cheating_(biology)http://en.wikipedia.org/wiki/Parasitismhttp://en.wikipedia.org/wiki/Symbiosishttp://en.wikipedia.org/wiki/Parasitichttp://en.wikipedia.org/wiki/Commensalhttp://en.wikipedia.org/wiki/Hummingbird_Hawkmoth

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A well known example of mutualism is the relationship betweenungulates(such as cows) andbacteria within

theirintestines. The ungulates benefit from the cellulase produced by the bacteria, which facilitates digestion;

the bacteria benefit from having a stable supply ofnutrients in thehostenvironment.

Mutualism plays a key part in ecology. For example, mutualistic interactions are vital for

terrestrialecosystem function as more than 48% of land plants rely on mycorrhizalrelationships withfungi to

provide them with inorganic compounds and trace elements.

In addition, mutualism is thought to have driven the evolution of much of the biological diversity we see, such

as flowerforms (important forpollination mutualisms) and co-evolutionbetween groups of species.[1] However

mutualism has historically received less attention than other interactions such as predation and parasitism.[2][3]

Measuring the exactfitness benefit to the individuals is not always straightforward, particularly when the

individuals can receive benefits from a range of species, for example most plant-pollinatormutualisms. It is

therefore common to categorise mutualisms according to the closeness of the association, using terms suchas obligate versusfacultative. Defining "closeness," however, is also problematic. It can refer to mutual

dependency (the species cannot live without one another) or the biological intimacy of the relationship in

relation to physical closeness (e.g. one species living within the tissues of the other species).[4]

Contents

[hide]

1 Types of relationships

1.1 Resource-resource

relationships

1.2 Service-resource

relationships

1.3 Service-service

relationships

2 Humans and mutualism

3 Mathematical theory

3.1 Type II functional

response

4 See also

4.1 Notes

5 References

http://en.wikipedia.org/wiki/Ungulateshttp://en.wikipedia.org/wiki/Ungulateshttp://en.wikipedia.org/wiki/Ungulateshttp://en.wikipedia.org/wiki/Cowshttp://en.wikipedia.org/wiki/Cowshttp://en.wikipedia.org/wiki/Bacteriahttp://en.wikipedia.org/wiki/Intestineshttp://en.wikipedia.org/wiki/Intestineshttp://en.wikipedia.org/wiki/Cellulasehttp://en.wikipedia.org/wiki/Digestionhttp://en.wikipedia.org/wiki/Nutrientshttp://en.wikipedia.org/wiki/Nutrientshttp://en.wikipedia.org/wiki/Host_(biology)http://en.wikipedia.org/wiki/Host_(biology)http://en.wikipedia.org/wiki/Ecologyhttp://en.wikipedia.org/wiki/Ecosystemhttp://en.wikipedia.org/wiki/Ecosystemhttp://en.wikipedia.org/wiki/Mycorrhizalhttp://en.wikipedia.org/wiki/Mycorrhizalhttp://en.wikipedia.org/wiki/Fungihttp://en.wikipedia.org/wiki/Fungihttp://en.wikipedia.org/wiki/Flowerhttp://en.wikipedia.org/wiki/Pollinationhttp://en.wikipedia.org/wiki/Pollinationhttp://en.wikipedia.org/wiki/Co-evolutionhttp://en.wikipedia.org/wiki/Co-evolutionhttp://en.wikipedia.org/wiki/Predationhttp://en.wikipedia.org/wiki/Parasitismhttp://en.wikipedia.org/wiki/Parasitismhttp://en.wikipedia.org/wiki/Fitness_(biology)http://en.wikipedia.org/wiki/Fitness_(biology)http://en.wikipedia.org/wiki/Pollinatorhttp://en.wikipedia.org/wiki/Obligatehttp://en.wikipedia.org/wiki/Facultativehttp://en.wikipedia.org/wiki/Facultativehttp://en.wikipedia.org/wiki/Mutualism_(biology)http://en.wikipedia.org/wiki/Mutualism_(biology)http://en.wikipedia.org/wiki/Ungulateshttp://en.wikipedia.org/wiki/Cowshttp://en.wikipedia.org/wiki/Bacteriahttp://en.wikipedia.org/wiki/Intestineshttp://en.wikipedia.org/wiki/Cellulasehttp://en.wikipedia.org/wiki/Digestionhttp://en.wikipedia.org/wiki/Nutrientshttp://en.wikipedia.org/wiki/Host_(biology)http://en.wikipedia.org/wiki/Ecologyhttp://en.wikipedia.org/wiki/Ecosystemhttp://en.wikipedia.org/wiki/Mycorrhizalhttp://en.wikipedia.org/wiki/Fungihttp://en.wikipedia.org/wiki/Flowerhttp://en.wikipedia.org/wiki/Pollinationhttp://en.wikipedia.org/wiki/Co-evolutionhttp://en.wikipedia.org/wiki/Predationhttp://en.wikipedia.org/wiki/Parasitismhttp://en.wikipedia.org/wiki/Fitness_(biology)http://en.wikipedia.org/wiki/Pollinatorhttp://en.wikipedia.org/wiki/Obligatehttp://en.wikipedia.org/wiki/Facultativehttp://en.wikipedia.org/wiki/Mutualism_(biology)

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6 Further reading

[edit]Types of relationships

Mutualistic transversals can be thought of as a form of "biological barter"

[4]

in which species trade resources (forexamplecarbohydrates or inorganic compounds) or services such asgamete, offspring dispersal, or protection

from predators.

[edit]Resource-resource relationships

Resource-resource interactions, in which one type of resource is traded for a different resource, are probably

the most common form of mutualism; for examplemycorrhizal associations between plantrootsandfungi, with

the plant providingcarbohydratesto thefungus in return for primarilyphosphate but

also nitrogenous compounds. Other examples includerhizobiabacteria which fix nitrogen

forleguminousplants (family Fabaceae) in return for energy-containing carbohydrates.[5]

[edit]Service-resource relationships

The Red-billed Oxpeckereats ticks on theimpala's coat

Service-resource relationships are also common.

Pollination in whichnectarorpollen (food resources) are traded for pollen dispersal (a service) orant protection

ofaphids, where the aphids tradesugar-rich honeydew(a by-product of their mode of feeding on plantsap) in

return for defense against predators such as ladybirdbeetles.

Phagophilesfeed (resource) onectoparasites, thereby providing anti-pest service.

Zoochory is an example where animals disperse the seeds of plants. This is similar to pollination in that the

plant produces food resources (for example, fleshy fruit, overabundance of seeds) for animals that disperse the

seeds (service).

[edit]Service-service relationships

http://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=1http://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/wiki/Gametehttp://en.wikipedia.org/wiki/Gametehttp://en.wikipedia.org/wiki/Gametehttp://en.wikipedia.org/wiki/Biological_dispersalhttp://en.wikipedia.org/wiki/Predatorhttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=2http://en.wikipedia.org/wiki/Mycorrhizalhttp://en.wikipedia.org/wiki/Mycorrhizalhttp://en.wikipedia.org/wiki/Roothttp://en.wikipedia.org/wiki/Roothttp://en.wikipedia.org/wiki/Roothttp://en.wikipedia.org/wiki/Fungihttp://en.wikipedia.org/wiki/Fungihttp://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/wiki/Fungushttp://en.wikipedia.org/wiki/Fungushttp://en.wikipedia.org/wiki/Phosphatehttp://en.wikipedia.org/wiki/Phosphatehttp://en.wikipedia.org/wiki/Nitrogenoushttp://en.wikipedia.org/wiki/Rhizobiahttp://en.wikipedia.org/wiki/Rhizobiahttp://en.wikipedia.org/wiki/Rhizobiahttp://en.wikipedia.org/wiki/Leguminoushttp://en.wikipedia.org/wiki/Leguminoushttp://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=3http://en.wikipedia.org/wiki/Red-billed_Oxpeckerhttp://en.wikipedia.org/wiki/Impalahttp://en.wikipedia.org/wiki/Impalahttp://en.wikipedia.org/wiki/Pollinationhttp://en.wikipedia.org/wiki/Nectarhttp://en.wikipedia.org/wiki/Nectarhttp://en.wikipedia.org/wiki/Pollenhttp://en.wikipedia.org/wiki/Pollenhttp://en.wikipedia.org/wiki/Anthttp://en.wikipedia.org/wiki/Aphidshttp://en.wikipedia.org/wiki/Aphidshttp://en.wikipedia.org/wiki/Sugarhttp://en.wikipedia.org/wiki/Sugarhttp://en.wikipedia.org/wiki/Honeydew_(melon)http://en.wikipedia.org/wiki/Honeydew_(melon)http://en.wikipedia.org/wiki/Saphttp://en.wikipedia.org/wiki/Saphttp://en.wikipedia.org/wiki/Predatorhttp://en.wikipedia.org/wiki/Ladybirdhttp://en.wikipedia.org/wiki/Phagophilehttp://en.wikipedia.org/wiki/Phagophilehttp://en.wikipedia.org/wiki/Ectoparasitehttp://en.wikipedia.org/wiki/Ectoparasitehttp://en.wikipedia.org/wiki/Zoochoryhttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=4http://en.wikipedia.org/wiki/File:Impala_mutualim_with_birds_wide.jpghttp://en.wikipedia.org/wiki/File:Impala_mutualim_with_birds_wide.jpghttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=1http://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/wiki/Gametehttp://en.wikipedia.org/wiki/Biological_dispersalhttp://en.wikipedia.org/wiki/Predatorhttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=2http://en.wikipedia.org/wiki/Mycorrhizalhttp://en.wikipedia.org/wiki/Roothttp://en.wikipedia.org/wiki/Fungihttp://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/wiki/Fungushttp://en.wikipedia.org/wiki/Phosphatehttp://en.wikipedia.org/wiki/Nitrogenoushttp://en.wikipedia.org/wiki/Rhizobiahttp://en.wikipedia.org/wiki/Leguminoushttp://en.wikipedia.org/wiki/Carbohydrateshttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=3http://en.wikipedia.org/wiki/Red-billed_Oxpeckerhttp://en.wikipedia.org/wiki/Impalahttp://en.wikipedia.org/wiki/Pollinationhttp://en.wikipedia.org/wiki/Nectarhttp://en.wikipedia.org/wiki/Pollenhttp://en.wikipedia.org/wiki/Anthttp://en.wikipedia.org/wiki/Aphidshttp://en.wikipedia.org/wiki/Sugarhttp://en.wikipedia.org/wiki/Honeydew_(melon)http://en.wikipedia.org/wiki/Saphttp://en.wikipedia.org/wiki/Predatorhttp://en.wikipedia.org/wiki/Ladybirdhttp://en.wikipedia.org/wiki/Phagophilehttp://en.wikipedia.org/wiki/Ectoparasitehttp://en.wikipedia.org/wiki/Zoochoryhttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=4

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An example of mutual symbiosis is the relationship betweenOcellaris clownfishthat dwell among thetentaclesofRitteri sea

anemones.

Strict service-service interactions are very rare, for reasons that are far from clear.[4]One example is the

relationship between sea anemones andanemonefish in the family Pomacentridae: the anemones provide the

fish with protection from predators (which cannot tolerate the stings of the anemone's tentacles) and the fish

defend the anemones against butterflyfish(familyChaetodontidae) which eat anemones. However, in common

with many mutualisms, there is more than one aspect to it: in the anemonefish-anemone mutualism,

wasteammonia from the fish feed the symbioticalgae that are found in the anemone's tentacles.[6][7] Therefore

what appears to be a service-service mutualism in fact has a service-resource component. A second example

is that of the relationship between some ants in the genus Pseudomyrmexand trees in the genusAcacia, such

as theWhistling Thorn and Bullhorn Acacia. The ants nest inside the plant's thorns. In exchange for shelter, the

ants protect acacias from attack byherbivores (which they frequently eat, introducing a resource component to

this service-service relationship) and competition from other plants by trimming back vegetation that would

shade the acacia. In addition, another service-resource component is present, as the ants regularly feed on

lipid-rich food-bodies called Beltian bodiesthat are on theAcacia plant.

In theNeotropics, the ant,Myrmelachista schumannimakes its nest in special cavities in Duroia hirsute. Plants

in the vicinity which belong to other species are killed with formic acid. This selective gardening can be so

aggressive that small areas of the rainforest are dominated byDuroia hirsute. These peculiar patches are

known by local people as "devil's gardens".[8]

In some of these relationships, the cost of the ants protection can be quite expensive.Cordiasp. trees in

the Amazonian rainforest have a kind of partnership withAllomerus sp. ants, which make their nests in

modified leaves. To increase the amount of living space available, the ants will destroy the trees flower buds.

The flowers die and leaves develop instead, provisioning the ants with more dwellings. Another type

ofAllomerus sp. ant lives with the Hirtella sp. tree in the same forests, but in this relationship the tree has turned

the tables on the greedy ants. When the tree is ready to produce flowers, the ant abodes on certain branches

http://en.wikipedia.org/wiki/Ocellaris_clownfishhttp://en.wikipedia.org/wiki/Ocellaris_clownfishhttp://en.wikipedia.org/wiki/Ocellaris_clownfishhttp://en.wikipedia.org/wiki/Tentaclehttp://en.wikipedia.org/wiki/Tentaclehttp://en.wikipedia.org/wiki/Tentaclehttp://en.wikipedia.org/wiki/Heteractis_magnificahttp://en.wikipedia.org/wiki/Heteractis_magnificahttp://en.wikipedia.org/wiki/Heteractis_magnificahttp://en.wikipedia.org/wiki/Heteractis_magnificahttp://en.wikipedia.org/wiki/Sea_anemonehttp://en.wikipedia.org/wiki/Anemonefishhttp://en.wikipedia.org/wiki/Anemonefishhttp://en.wikipedia.org/wiki/Pomacentridaehttp://en.wikipedia.org/wiki/Predatorhttp://en.wikipedia.org/wiki/Butterflyfishhttp://en.wikipedia.org/wiki/Butterflyfishhttp://en.wikipedia.org/wiki/Chaetodontidaehttp://en.wikipedia.org/wiki/Chaetodontidaehttp://en.wikipedia.org/wiki/Ammoniahttp://en.wikipedia.org/wiki/Ammoniahttp://en.wikipedia.org/wiki/Symbiotichttp://en.wikipedia.org/wiki/Algaehttp://en.wikipedia.org/wiki/Algaehttp://en.wikipedia.org/wiki/Antshttp://en.wikipedia.org/wiki/Genushttp://en.wikipedia.org/wiki/Acaciahttp://en.wikipedia.org/wiki/Acaciahttp://en.wikipedia.org/wiki/Whistling_Thornhttp://en.wikipedia.org/wiki/Whistling_Thornhttp://en.wikipedia.org/wiki/Bullhorn_Acaciahttp://en.wikipedia.org/wiki/Antshttp://en.wikipedia.org/wiki/Herbivoreshttp://en.wikipedia.org/wiki/Herbivoreshttp://en.wikipedia.org/wiki/Beltian_bodieshttp://en.wikipedia.org/wiki/Beltian_bodieshttp://en.wikipedia.org/wiki/Neotropicshttp://en.wikipedia.org/wiki/Neotropicshttp://en.wikipedia.org/wiki/Myrmelachista_schumannihttp://en.wikipedia.org/wiki/Myrmelachista_schumannihttp://en.wikipedia.org/wiki/Myrmelachista_schumannihttp://en.wikipedia.org/wiki/Cordiahttp://en.wikipedia.org/wiki/Cordiahttp://en.wikipedia.org/wiki/Cordiahttp://en.wikipedia.org/wiki/Amazon_Rainforesthttp://en.wikipedia.org/w/index.php?title=Allomerus&action=edit&redlink=1http://en.wikipedia.org/wiki/Hirtellahttp://en.wikipedia.org/wiki/File:Common_clownfish_curves_dnsmpl.jpghttp://en.wikipedia.org/wiki/File:Common_clownfish_curves_dnsmpl.jpghttp://en.wikipedia.org/wiki/Ocellaris_clownfishhttp://en.wikipedia.org/wiki/Tentaclehttp://en.wikipedia.org/wiki/Heteractis_magnificahttp://en.wikipedia.org/wiki/Heteractis_magnificahttp://en.wikipedia.org/wiki/Sea_anemonehttp://en.wikipedia.org/wiki/Anemonefishhttp://en.wikipedia.org/wiki/Pomacentridaehttp://en.wikipedia.org/wiki/Predatorhttp://en.wikipedia.org/wiki/Butterflyfishhttp://en.wikipedia.org/wiki/Chaetodontidaehttp://en.wikipedia.org/wiki/Ammoniahttp://en.wikipedia.org/wiki/Symbiotichttp://en.wikipedia.org/wiki/Algaehttp://en.wikipedia.org/wiki/Antshttp://en.wikipedia.org/wiki/Genushttp://en.wikipedia.org/wiki/Acaciahttp://en.wikipedia.org/wiki/Whistling_Thornhttp://en.wikipedia.org/wiki/Bullhorn_Acaciahttp://en.wikipedia.org/wiki/Antshttp://en.wikipedia.org/wiki/Herbivoreshttp://en.wikipedia.org/wiki/Beltian_bodieshttp://en.wikipedia.org/wiki/Neotropicshttp://en.wikipedia.org/wiki/Myrmelachista_schumannihttp://en.wikipedia.org/wiki/Cordiahttp://en.wikipedia.org/wiki/Amazon_Rainforesthttp://en.wikipedia.org/w/index.php?title=Allomerus&action=edit&redlink=1http://en.wikipedia.org/wiki/Hirtella

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begin to wither and shrink, forcing the occupants to flee, leaving the trees flowers to develop free from ant

attack.[8]

[edit]Humans and mutualism

Dogs and sheep were among the first animals to bedomesticated.

Humans also engage in mutualisms with other species, including theirgut flora (without which they would not

be able to digest food efficiently) anddomesticated animals such as horses, which provide transportation in

return for food and shelter. In traditional agriculture, many plants will function mutualistically as companion

plants, providing each other with shelter, soil fertility and the repelling of pests. For example,beansmay grow

upcornstalks as a trellis, while fixing nitrogen in the soil for the corn, as exploited in the ThreeSisters gardening technique.

[edit]Mathematical theory

In 1989, David Hamilton Wright developed a mathematical explanation for mutualism using the LotkaVolterra

equation. Wright modified theLotka-Volterra equations by adding a new term, M/K, to represent a mutualistic

relationship.[9]

The mutalistic relationship is quantified by:

dN/dt = r1 N( 1 N/K1 + 12 M/K1 )

dM/dt = r2 M ( 1 M/K2 + 21 N/K2 )

where,

N and M = the population density

r = intrinsic growth rate of the population

http://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=5http://en.wikipedia.org/wiki/Dogshttp://en.wikipedia.org/wiki/Sheephttp://en.wikipedia.org/wiki/Domesticationhttp://en.wikipedia.org/wiki/Domesticationhttp://en.wikipedia.org/wiki/Gut_florahttp://en.wikipedia.org/wiki/Domesticationhttp://en.wikipedia.org/wiki/Domesticationhttp://en.wikipedia.org/wiki/Companion_planthttp://en.wikipedia.org/wiki/Companion_planthttp://en.wikipedia.org/wiki/Companion_planthttp://en.wikipedia.org/wiki/Beanhttp://en.wikipedia.org/wiki/Beanhttp://en.wikipedia.org/wiki/Beanhttp://en.wikipedia.org/wiki/Maizehttp://en.wikipedia.org/wiki/Maizehttp://en.wikipedia.org/wiki/Three_Sisters_(agriculture)http://en.wikipedia.org/wiki/Three_Sisters_(agriculture)http://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=6http://en.wikipedia.org/wiki/Lotka%E2%80%93Volterra_equationhttp://en.wikipedia.org/wiki/Lotka%E2%80%93Volterra_equationhttp://en.wikipedia.org/wiki/Lotka%E2%80%93Volterra_equationhttp://en.wikipedia.org/wiki/Lotka%E2%80%93Volterra_equationhttp://en.wikipedia.org/wiki/Lotka%E2%80%93Volterra_equationhttp://en.wikipedia.org/wiki/File:Backing_sheep_at_sheepdog_competition.jpghttp://en.wikipedia.org/wiki/File:Backing_sheep_at_sheepdog_competition.jpghttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=5http://en.wikipedia.org/wiki/Dogshttp://en.wikipedia.org/wiki/Sheephttp://en.wikipedia.org/wiki/Domesticationhttp://en.wikipedia.org/wiki/Gut_florahttp://en.wikipedia.org/wiki/Domesticationhttp://en.wikipedia.org/wiki/Companion_planthttp://en.wikipedia.org/wiki/Companion_planthttp://en.wikipedia.org/wiki/Beanhttp://en.wikipedia.org/wiki/Maizehttp://en.wikipedia.org/wiki/Three_Sisters_(agriculture)http://en.wikipedia.org/wiki/Three_Sisters_(agriculture)http://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=6http://en.wikipedia.org/wiki/Lotka%E2%80%93Volterra_equationhttp://en.wikipedia.org/wiki/Lotka%E2%80%93Volterra_equationhttp://en.wikipedia.org/wiki/Lotka%E2%80%93Volterra_equation

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K =carrying capacityof its local environmental setting.

= coefficient converting encounters with one species to new units of the

other

Mutualism is essentially thelogistic growth equation + mutualistic interaction. The mutualistic interaction termrepresents the increase in population growth of species one as a result of the presence of greater numbers of

species two, and vice versa. Wright also considered the concept of saturation, which means that with higher

densities, there are decreasing benefits of further increases of the mutualist population. Without saturation,

species' densities would increase indefinitely. Because that isn't possible due to environmental constraints and

carrying capacity, a model that includes saturation would be more accurate. Wright's mathematical theory is

based on the premise of a simple two-species mutualism model in which the benefits of mutualism become

saturated due to limits posed by handling time. Wright defines handling time as the time needed to process a

food item, from the initial interaction to the start of a search for new food items and assumes that processing of

food and searching for food are mutually exclusive. Mutualists that display foraging behavior are exposed to the

restrictions on handling time. Mutualism can be associated with symbiosis

[edit]Type II functional response

In 1959, C. S. Holling performed his classic disc experiment that assumed the following: that (1), the number of

food items captured is proportional to the allotted searching time; and (2), that there is a variable ofhandling

time that exists separately from the notion of search time. He then developed an equation for the Type

II functional response, which showed that the feeding rate is equivalent to

ax/(1 + axTH).

where,

a = the instantaneous discovery rate

x = food item density

TH = handling time

The equation that incorporates Type II functional response and mutualism is:

dN/dt = N [r(1-cN) + M(X+M)]

where,

N and M = density of the two mutualists

r = intrinsic rate of increase of N

c = coefficient measuring negative intraspecific interaction

X = 1/a TH

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= b/ TH

a = instantaneous discovery rate

b = coefficient converting encounters with M to new units of N

Rearranged: dN/dt = N[r(1-cN) + baM/(1+aTHM)]

These two graphs show the isoclinesof mutualistic relationships. Presence of the mutualist has a positive

effect on the recipient population at low densities of mutualists but at high densities, saturation takes place, and

further increases in mutualist populations have little effect. Arrows indicate the stability or instability of

equilibrium points.

Graph A illustrates a facultative mutualism in which both species derive benefits from each other, but do not

depend on the existence of the other for survival.

Graph B demonstrates anobligate mutualism in which both species are interdependent on each other in a way

that one species cannot survive without the other. Once the population of one species drops below the first

equilibrium point, both populations will go extinct.

The model presented above is most effectively applied to free-living species that encounter a number of

individuals of the mutualist part in the course of their existences. Of note, as Wright points out, is that models of

biological mutualism tend to be similar qualitatively, in that the featured isoclines generally have a positive

decreasing slope, and by and large similar isocline diagrams. Mutualistic interactions are best visualized as

positively sloped isoclines, which can be explained by the fact that the saturation of benefits accorded to

mutualism or restrictions posed by outside factors contribute to a decreasing slope.

[edit]See also

Co-adaptation

Co-evolution

Ecological facilitation

Frugivory

Greater Honeyguide - has an interesting mutualism with humans

Interspecific communication

List of symbiotic relationships

Mllerian mimicry

Mutualisms and Conservation

Mutual Aid: A Factor of Evolution

http://en.wikipedia.org/wiki/Isoclinehttp://en.wikipedia.org/wiki/Isoclinehttp://en.wikipedia.org/wiki/Facultativehttp://en.wikipedia.org/wiki/Obligatehttp://en.wikipedia.org/wiki/Obligatehttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=8http://en.wikipedia.org/wiki/Co-adaptationhttp://en.wikipedia.org/wiki/Co-evolutionhttp://en.wikipedia.org/wiki/Ecological_facilitationhttp://en.wikipedia.org/wiki/Frugivoryhttp://en.wikipedia.org/wiki/Greater_Honeyguidehttp://en.wikipedia.org/wiki/Interspecific_communicationhttp://en.wikipedia.org/wiki/List_of_symbiotic_relationshipshttp://en.wikipedia.org/wiki/M%C3%BCllerian_mimicryhttp://en.wikipedia.org/wiki/Mutualisms_and_Conservationhttp://en.wikipedia.org/wiki/Mutual_Aid:_A_Factor_of_Evolutionhttp://en.wikipedia.org/wiki/Isoclinehttp://en.wikipedia.org/wiki/Facultativehttp://en.wikipedia.org/wiki/Obligatehttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edit&section=8http://en.wikipedia.org/wiki/Co-adaptationhttp://en.wikipedia.org/wiki/Co-evolutionhttp://en.wikipedia.org/wiki/Ecological_facilitationhttp://en.wikipedia.org/wiki/Frugivoryhttp://en.wikipedia.org/wiki/Greater_Honeyguidehttp://en.wikipedia.org/wiki/Interspecific_communicationhttp://en.wikipedia.org/wiki/List_of_symbiotic_relationshipshttp://en.wikipedia.org/wiki/M%C3%BCllerian_mimicryhttp://en.wikipedia.org/wiki/Mutualisms_and_Conservationhttp://en.wikipedia.org/wiki/Mutual_Aid:_A_Factor_of_Evolution

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Symbiogenesis

[edit]Notes

1. ^ Thompson, J. N. 2005 The geographic mosaic of coevolution. Chicago, IL:

University of Chicago Press.

2. ^ Bronstein, JL. 1994. Our current understand of mutualism.Quarterly Review

of Biology 69 (1): 31-51 March 1994

3. ^ Begon, M., J.L. Harper, and C.R. Townsend. 1996.Ecology: individuals,

populations, and communities, Third Edition. Blackwell Science Ltd.,

Cambridge, Massachusetts, USA.

4. ^ abcOllerton, J. 2006. "Biological Barter": Patterns of Specialization

Compared across Different Mutualisms. pp. 411-435 in: Waser, N.M. &

Ollerton, J. (Eds) Plant-Pollinator Interactions: From Specialization toGeneralization. University of Chicago Press.

5. ^ Denison RF, Kiers ET 2004. Why are most rhizobia beneficial to their plant

hosts, rather than parasitic.Microbes and Infection 6 (13): 1235-1239

6. ^ Porat, D. & Chadwick-Furman, N. E. 2004 Effects of anemonefish on giant

sea anemones: expansion behavior,growth, and survival.Hydrobiologia 530,

513520. (doi:10.1007/s10750-004-2688-y)

7. ^ Porat, D. & Chadwick-Furman, N. E. 2005 Effects of anemonefish on giant

sea anemones: ammonium uptake,zooxanthella content and tissue

regeneration. Mar. Freshw.Behav. Phys. 38, 4351.

(doi:10.1080/102362405000 57929)

8. ^ abPiper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious

and Unusual Animals,Greenwood Press.

9. ^ Wright, David Hamilton. 1989. A Simple, Stable Model of Mutualism

Incorporating Handling Time.The American Naturalist, Vol. 134, No. 4, pp.

664-667.

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Porat, D. & Chadwick-Furman, N. E. 2005. Effects of anemonefish on giant sea

anemones: ammonium uptake,zooxanthella content and tissue regeneration. Mar.

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Thompson, J. N. 2005. The Geographic Mosaic of Coevolution. University of

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Wright, David Hamilton. 1989. A Simple, Stable Model of Mutualism Incorporating

Handling Time. The American Naturalist, Vol. 134, No. 4, pp. 664667.

[edit]Further reading

Wikimedia Commons has

media related to:Mutualism

Look up mutualism

(biology)inWiktionary, the

free dictionary.

Boucher, D. G., James, S. & Kresler, K. (1984) The ecology of

mutualism.Annual Review of Ecology and Systematics, 13: 315-347.

Boucher, D. H. (editor) (1985) The Biology of Mutualism : Ecology and

Evolution London : Croom Helm 388 p.ISBN 0-7099-3238-3

[show]vdeInter-species biological interactions in ecology

[show]vdeEvolutionary ecology

[hide]vdeSwarming

Swarm algorithms

Agent-based modelsAnt colony optimizationAnt roboticsArtificial AntsBees algorithmBee colony

optimizationBoidsCrowd simulationFirefly algorithmGlowworm swarm optimizationParticle swarm

optimizationSelf-propelled particlesSwarm intelligenceSwarm (simulation)

Biological swarming

Agent-based model in biologyBait ballCollective animal behaviorFeeding

frenzyFlockFlockingHerdHerd behaviorMixed-species foraging flockMobbing behaviorPack

hunterPatterns of self-organization in antsSardine runShoaling and schoolingSort solSwarming

behaviourSwarming (honey bee)Swarming motility

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Animal migrationAnimal migrationBird migrationBird migration flywaysDiel vertical migrationFish migrationHoming

(biology)Insect migrationLessepsian migrationLepidoptera migrationReverse migration

Swarm roboticsI-SwarmMicroboticsSwarm roboticsSymbrion

Related topicsAllee effectBiomimicryCollective intelligenceEusocialityGroup size measuresMutualismPredator

satiationSpatial organizationStigmergySwarming (military)Task allocation and partitioning of social insects

[show]Modelling ecosystems- trophiccomponents

[show]Modelling ecosystems- other components

Categories:Mutualism (biology) |Biological interactions| Symbiosis

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http://en.wikipedia.org/wiki/Animal_migrationhttp://en.wikipedia.org/wiki/Animal_migrationhttp://en.wikipedia.org/wiki/Bird_migrationhttp://en.wikipedia.org/wiki/Bird_migrationhttp://en.wikipedia.org/wiki/Category:Bird_migration_flywayshttp://en.wikipedia.org/wiki/Category:Bird_migration_flywayshttp://en.wikipedia.org/wiki/Category:Bird_migration_flywayshttp://en.wikipedia.org/wiki/Diel_vertical_migrationhttp://en.wikipedia.org/wiki/Diel_vertical_migrationhttp://en.wikipedia.org/wiki/Diel_vertical_migrationhttp://en.wikipedia.org/wiki/Fish_migrationhttp://en.wikipedia.org/wiki/Fish_migrationhttp://en.wikipedia.org/wiki/Homing_(biology)http://en.wikipedia.org/wiki/Homing_(biology)http://en.wikipedia.org/wiki/Homing_(biology)http://en.wikipedia.org/wiki/Insect_migrationhttp://en.wikipedia.org/wiki/Insect_migrationhttp://en.wikipedia.org/wiki/Lessepsian_migrationhttp://en.wikipedia.org/wiki/Lessepsian_migrationhttp://en.wikipedia.org/wiki/Lepidoptera_migrationhttp://en.wikipedia.org/wiki/Lepidoptera_migrationhttp://en.wikipedia.org/wiki/Reverse_migrationhttp://en.wikipedia.org/wiki/Reverse_migrationhttp://en.wikipedia.org/wiki/Swarm_roboticshttp://en.wikipedia.org/wiki/Intelligent_Small_World_Autonomous_Robots_for_Micro-manipulationhttp://en.wikipedia.org/wiki/Intelligent_Small_World_Autonomous_Robots_for_Micro-manipulationhttp://en.wikipedia.org/wiki/Microboticshttp://en.wikipedia.org/wiki/Microboticshttp://en.wikipedia.org/wiki/Swarm_roboticshttp://en.wikipedia.org/wiki/Swarm_roboticshttp://en.wikipedia.org/wiki/Symbrionhttp://en.wikipedia.org/wiki/Symbrionhttp://en.wikipedia.org/wiki/Allee_effecthttp://en.wikipedia.org/wiki/Allee_effecthttp://en.wikipedia.org/wiki/Biomimicryhttp://en.wikipedia.org/wiki/Collective_intelligencehttp://en.wikipedia.org/wiki/Collective_intelligencehttp://en.wikipedia.org/wiki/Eusocialityhttp://en.wikipedia.org/wiki/Eusocialityhttp://en.wikipedia.org/wiki/Eusocialityhttp://en.wikipedia.org/wiki/Group_size_measureshttp://en.wikipedia.org/wiki/Group_size_measureshttp://en.wikipedia.org/wiki/Predator_satiationhttp://en.wikipedia.org/wiki/Predator_satiationhttp://en.wikipedia.org/wiki/Predator_satiationhttp://en.wikipedia.org/wiki/Predator_satiationhttp://en.wikipedia.org/wiki/Spatial_organizationhttp://en.wikipedia.org/wiki/Stigmergyhttp://en.wikipedia.org/wiki/Stigmergyhttp://en.wikipedia.org/wiki/Stigmergyhttp://en.wikipedia.org/wiki/Swarming_(military)http://en.wikipedia.org/wiki/Swarming_(military)http://en.wikipedia.org/wiki/Task_allocation_and_partitioning_of_social_insectshttp://en.wikipedia.org/wiki/Task_allocation_and_partitioning_of_social_insectshttp://en.wikipedia.org/wiki/Mutualism_(biology)http://en.wikipedia.org/wiki/Ecosystem_modelhttp://en.wikipedia.org/wiki/Ecosystem_modelhttp://en.wikipedia.org/wiki/Ecosystem_modelhttp://en.wikipedia.org/wiki/Trophic_levelhttp://en.wikipedia.org/wiki/Trophic_levelhttp://en.wikipedia.org/wiki/Mutualism_(biology)http://en.wikipedia.org/wiki/Ecosystem_modelhttp://en.wikipedia.org/wiki/Ecosystem_modelhttp://en.wikipedia.org/wiki/Ecosystem_modelhttp://en.wikipedia.org/wiki/Special:Categorieshttp://en.wikipedia.org/wiki/Category:Mutualism_(biology)http://en.wikipedia.org/wiki/Category:Mutualism_(biology)http://en.wikipedia.org/wiki/Category:Biological_interactionshttp://en.wikipedia.org/wiki/Category:Biological_interactionshttp://en.wikipedia.org/wiki/Category:Biological_interactionshttp://en.wikipedia.org/wiki/Category:Symbiosishttp://en.wikipedia.org/w/index.php?title=Special:UserLogin&returnto=Mutualism_(biology)&campaign=ACP2http://en.wikipedia.org/w/index.php?title=Special:UserLogin&returnto=Mutualism_(biology)&campaign=ACP2http://en.wikipedia.org/wiki/Mutualism_(biology)http://en.wikipedia.org/wiki/Talk:Mutualism_(biology)http://en.wikipedia.org/wiki/Mutualism_(biology)http://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edithttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=edithttp://en.wikipedia.org/w/index.php?title=Mutualism_(biology)&action=historyhttp://en.wikipedia.org/wiki/Main_Pagehttp://en.wikipedia.org/wiki/Portal:Contentshttp://en.wikipedia.org/wiki/Portal:Featured_contenthttp://en.wikipedia.org/wiki/Portal:Featured_contenthttp://en.wikipedia.org/wiki/Portal:Current_eventshttp://en.wikipedia.org/wiki/Special:Randomhttp://wikimediafoundation.org/wiki/Special:Landingcheck?landing_page=WMFJA085&language=en&utm_source=donate&utm_medium=sidebar&utm_campaign=20101204SB002http://en.wikipedia.org/wiki/Help:Contentshttp://en.wikipedia.org/wiki/Wikipedia:Abouthttp://en.wikipedia.org/wiki/Wikipedia:Community_portalhttp://en.wikipedia.org/wiki/Wikipedia:Community_portalhttp://en.wikipedia.org/wiki/Special:RecentChangeshttp://en.wikipedia.org/wiki/Wikipedia:Contact_ushttp://ar.wikipedia.org/wiki/%D8%AA%D9%86%D8%A7%D9%81%D8%B9http://ar.wikipedia.org/wiki/%D8%AA%D9%86%D8%A7%D9%81%D8%B9http://be.wikipedia.org/wiki/%D0%9C%D1%83%D1%82%D1%83%D0%B0%D0%BB%D1%96%D0%B7%D0%BChttp://bg.wikipedia.org/wiki/%D0%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