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Plant Development
March 28
Developmental Biology, Spring 2016
Migration from water to land
• Life evolved in aquatic environments for 3 billion years
before migrating to land
• Evolutionary pilgrimage of complex organisms onto land
began only 500 million years ago
• Terrestrial colonies founded by plants completely
transformed the biosphere
Adaptation to land
• Advantages
– Access to sunlight
– CO2 as gas
– (initially) few predators/herbivores
• Challenges
– Dessiccate environment
– Mobility
– Transport nutrients
– Supportive structures
– Reproduction
Adaptation to land
• Advantages
– Access to sunlight
– CO2 as gas
– (initially) few predators/herbivores
• Challenges
– Dessiccate environment (waxy leaves, seed coats)
– Mobility (leaf structure, cell shape)
– Transport nutrients (vasculature)
– Supportive structures (roots, shoots)
– Reproduction (embryos, seeds, flowers, dispersal)
Ancestral green algae
(450 million years ago)
Land plants
Vascular plants
Seed plants
Last common ancestor of
animals and plants was a
single-celled organism
The plant body
• Plants have three basic
organs
– Roots
– Stems
– Leaves
• Plant organs are
composed of three tissue
systems
– Dermal
– Vascular
– Ground
Angiosperms
• Derived from Greek: ‘enclosed seed’
• Largest phylum of land plants; >300,000 species
• Vascular seed plants
• Ovule is fertilized
• Ovule develops into a seed inside an enclosed ovary
• Ovary is enclosed inside a flower
• Flower contains the male or female organs, or both
• Flower develops into a fruit
• Gymnosperms (the other phylum of seed-bearing plants)
do not develop their seeds inside an ovary (seeds are on
the surface, e.g. cones)
Life cycle
Pollen
• Male gamete
• Mature pollen contains two cells
• Tube cell: guides pollen
germination and growth of the
pollen tube
• Generative cell: divides to
produce 2 sperm
• One sperm will fertilize the egg
• One sperm will participate in the
formation of the endosperm (a
structure that provides
nourishment for the embryo)
Carpel
• Contains the stigma, style, and
an ovary containing one or more
ovules
• Ovules are attached by a
placenta to the ovary wall
• Fully developed ovules are called
seeds
• Integuments are layers of cells
that enclose the spores
• Pollen tube will grow through the
micropyle
• There are four female
gametophytes inside the
megaspore – one of these is the
egg
Self incompatibility
• Pollination does not
guarantee fertilization
• Interspecific incompatibility
(between species)
• Intraspecific incompatibility
(within species)
• Recognition of self depends
on the self-incompatibility
locus (S locus)
Self incompatibility
Multiple alleles of the S locus dictate incompatibility
Double Fertilization
• The pollen tube enters the embryo sac and two sperm cells are
released
• The ovule has to undergo meiosis and produces 4 haploid
megaspores
• 3 megaspores degenerate, and one haploid megaspore remains
• This megaspore undergoes mitosis to form 8 nuclei (embryo sac)
• Cell walls start to form
• Egg cell and polar nuclei are fertilized
Embryogenesis
• Radial patterning: produces three tissue systems – Ground
– Dermal
– Vascular
• Axial patterning: produces apical-basal axis (root-shoot) – Auxin hormone
– Activates transcription by degrading a repressor of transcription
• Assymetric cell division
• Set aside meristems for post-embryonic development
• Establish accessible food reserve for germinating embryo – Development of cotyledons from the endosperm
Embryogenesis
Dormancy
• At the end of embryogenesis there is a shift from constructing
the body plan to creating a food reserve
• Genes coding for seed storage proteins start to get activated
• Metabolism slows, and the seed desiccates
• A seed coat forms from the integuments
• Gene called viviparous triggers dormancy
• Viviparous mutants act like ferns – goes directly to
postembryonic development
• Abscisic acid (hormone) maintains dormancy
• Gibberelins (hormone) block dormancy
• Seeds can stay dormant for years
Plant development
Embryonic vs. Post-embryonic development
Most animal development is embryonic
Plant development
Embryonic vs. Post-embryonic development
In plants, the mature embryo and young plant have only two leaves
Plant development
Embryonic vs. Post-embryonic development
New organs are made in response to environmental cues
Regeneration and extreme flexibility
The plant body
Where does growth occur?
Meristems: Stem cell population and the source of new plant tissues
Pluripotent
Shoot apical
meristem (SAM)
Leaves,
branches,
flowers
Root apical
meristem (RAM)
Root organs
Shoot apical meristem
Root apical meristem
• Meristem
grows in two
directions
• The meristem
itself doesn’t
give rise to
appendages
(roots)
• Lateral roots
grow out of
mature tissue
Stem cells
Stem cell niche
Specifying fate
Flowering or Inflorescence
• Inflorescence signals a reproductive transition in plants
• Signals from leaves trigger flowering
• In some species this is dependent on photoperiodicity (day length)
• Phytochrome pigments transduce signals from the environment
• Vernalization (period of cold) enhances the flowering signal
• Vegetative to reproductive state
A derivative of a stem cell (yellow) is
displaced to the periphery of the shoot
meristem, becomes part of a floral
meristem and is incorporated into a flower
Transition of Shoot Apical Meristem
from vegetative to floral meristem
Flower
• Specialized shoot with four circles of modified leaves
• Sepals
• Petals
• Stamens (male)
• Carpels (female)
Structure of a flower
Floral identity
• Specification of floral meristems
• Genes like LEAFY, APETALA, CAULIFLOWER are floral
meristem identity genes
Wild type Mutant
Floral identity
Wild-rose
Simple flower with
one row of 5 petals
Hybrid tea rose
A genetic mutant with
(35 or 40 petals)
selected by breeders
• What kind of mutation is it?
• Homeotic mutations
• In a developing flower, the order of each primordium’s emergence determines its fate: sepal, petal, stamen, or carpel
• Plant biologists have identified several organ identity genes (plant homeotic genes) that regulate the development of floral pattern
• These are MADS-box transcription factor genes
• Not HOX genes
• A mutation in a plant organ identity gene can cause abnormal floral development
Floral identity
In a developing flower, the order of each
primordium’s emergence (outer to inner) determines
its fate: sepal, petal, stamen, or carpel
© 2011 Pearson Education, Inc.
Developing
leaves
Shoot Apical Meristem (SAM) Inflorescence
Transition from shoot apical meristem to infloresence
Rule 1. Each gene acts in two whorls
(e.g. A is expressed in whorls 1 & 2)
A A
B B
C C
Three rules of the ABC model
Rule 1. Each gene acts in two whorls
(e.g. A is expressed in whorls 1 & 2)
Rule 2. Combinations of gene products determine organ identity
(e.g. activity A + B specify a petal)
A A
B B
C C
Three rules of the ABC model
Rule 1. Each gene acts in two whorls
(e.g. A is expressed in whorls 1 & 2)
Rule 2. Combinations of gene products determine organ identity
(e.g. activity A + B specify a petal)
Rule 3. A and C activities are mutually exclusive
(e.g. when A is mutated, C is expressed in that whorl)
A A
B B
C C
Three rules of the ABC model
For Rule 3, if the products of gene A and gene C gene have
antagonistic action, will A and C ever be expressed in the same
whorl?
Whorls
Organs
For Rule 3, if the products of gene A and gene C gene have
antagonistic action, will A and C ever be expressed in the same
whorl?
Answer: No
Whorls
Organs
For Rule 3, if the products of gene A and gene C gene have
antagonistic action, will A and C ever be expressed in the same
whorl?
Answer: No
Follow-up question:
If gene A is normally expressed in whorls 1 and 2, which gene
will be expressed in those whorls 1 and 2 in a mutant of gene A?
Whorls
Organs
For Rule 3, if the products of gene A and gene C gene have
antagonistic action, will A and C ever be expressed in the same
whorl?
Answer: No
Follow-up question:
If gene A is normally expressed in whorls 1 and 2, which gene
will be expressed in those whorls 1 and 2 in a mutant of gene A?
Answer: Gene C
Whorls
Organs
What are the organs in a gene A mutant?
Why is it called Apetela?
Whorls
Organs
Gene A is missing
When A is missing, then C is expressed in the whorl where A used to be…
Remember Rule 3 of ABC model
A and C activities are mutually exclusive
(e.g. when A is mutated, C is expressed in that whorl)
C C C C B B
Apetala 2 (no petals) = mutation in Gene A
Wild-type flower
A A B B
C C
What types of floral organs are made in a mutant in gene B?
Whorls
Organs
Apetala 3 and Pistillata (carpel is also called pistil, thus Pistillata)
= Gene B mutant
Wild-type flower
A A B B
C C
A A
C C
What types of floral organs are made in a mutant in gene C?
Why is it called Agamous?
Whorls
Organs
Agamous (no gametes) = Gene C mutant
Wild-type flower
A A B B
C C
A A A A B B
Normal flower
Gene A Apetala 2 mutant (no petals) Only carpels and stamen)
Gene B Apetala 3 and Pistalata mutant (no petals or stamen) Only sepals and carpels
Gene C Agamous mutant (no gametes) Only sepals and petals
What types of floral organs are made in a double mutant of gene B
and gene C?
Whorls
Organs
What types of organs are made in a triple mutant A, B and C?
Whorls
Organs
The triple ABC mutant makes leaves in the 4 whorls
What does this tell you about the relationship between
leaves and flowers?
This result supports the theory that flowers evolved from leaves,
This theory dates to 1790, and was proposed by Goethe the existentialist
Goethe proposed that the different parts of a plant result from the
“metamorphosis (meaning transformation) of a basic organ, the “ideal leaf”
into other more specialized organs.
The forth whorl does not contain carpels
(female organ), but instead contains extra
stamen (male organs) hence the name
superman.
The superman mutant
Follow up question:
What is the normal function of
wild-type Superman gene?
Whorls
Organs
In the superman
mutant
the B gene is
expressed
in 4th whorl, so whorl 4
becomes a stamen
instead of a carpel
Organs
B
stamen
Wild-Type superman
Suppresses B gene in whorl 4
Answer: Wild-type superman gene normally suppresses
gene B activity in 4th whorl
Superman Gene
Fruiting
• Mature ovary
• Helps seed dispersal for reproduction
• Wall of the ovary becomes the pericarp which is the thick
wall of the fruit
Type of fruit Floral origin Example
Simple Single ovary of one
flower
Aggregate Many ovaries of one
flower
Multiple Many ovaries of many
clustered flowers
Midterm exam
Class average: 89
Class average without extra credit: 83
For next class
In-class quiz (10 questions, 15 minutes)
• Plant Development
Paper discussion
Kareem et al. Plethora genes control regeneration
by a two-step mechanism.
There are two supplemental readings to help with
the material
Email two questions about the paper