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Chapter 40

Plant Reproduction
BIOLOGY
Thirteenth Edition
Raven, Johnson, Mason, Losos,
Duncan

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No reproduction or further distribution permitted without the prior written consent of McGraw Hill, LLC.
Lecture Outline

40.1 Reproductive Development


40.2 Making Flowers
40.3 Structure and Evolution of
Flowers
40.4 Pollination and Fertilization
40.5 Embryo Development
40.6 Germination
40.7 Asexual Reproduction
40.8 Plant Life Spans
©Heather Angel/Natural Visions

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Reproductive Development

• Angiosperms represent an evolutionary innovation with


their production of flowers and fruits
• Plants go through developmental changes leading to
reproductive maturity by adding structures to existing ones
with meristems

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Life Cycle of an Angiosperm

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Initiation of Flowering

• Once plants are competent to reproduce, a combination of factors –


including light, temperature, and both promotive and inhibitory internal
signals – determines when a flower is produced
• Undergo phase change – subtle or obvious

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Phase Change

• Internal developmental changes allow plants to obtain


competence to respond to external or internal signals that
trigger flower formation
• May be morphologically obvious or very subtle

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Examples of Phase Changes

(a) mauritius images GmbH/Alamy Stock Photo; (b) Pat Breen, Oregon State University

The lower branches in the oak tree retain their leaves in the winter,
because these lower branches were initiated by juvenile meristems and
have not made a phase change. Juvenile ivy makes adventitious roots
that can cling to walls, but after a phase change mature ivy lacks the
ability to produce adventitious roots.

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Delay of Flowering

In Arabidopsis, the gene


embryonic flower (emf)
prevents early flowering
• emf mutants flower
immediately.
Flowering is the default state
Many mechanisms have
evolved to delay flowering

Courtesy of Lingjing Chen & Renee Sung

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Inducing Flowering

• The juvenile-to-adult transition can be induced by


overexpressing a flowering gene called LEAFY
• LEAFY (LFY) was cloned in Arabidopsis and replaced with
a viral promoter that results in constant, high levels of LFY
transcription
• Overexpression of LFY in aspen causes flowering to occur
in weeks instead of years

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Accelerated Phase Change

(a) Roadrunner1866/Shutterstock; (b Top) Blickwinkel/Teigler/Alamy Stock Photo; (b Bottom) ©Ove Nilsson/Umeå Plant Science Centre and Detlef Weigel/Max Planck Institute for
Developmental Biology

Normally aspen trees grow for several years before producing flowers.
Overexpression of the Arabidopsis flowering gene LFY causes rapid
flowering in transgeneic aspen.
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Flower Production

Four genetically regulated pathways to flowering have been


identified
1. The light-dependent pathway
2. The temperature-dependent pathway
3. The gibberellin-dependent pathway
4. The autonomous pathway
Plants can rely primarily on one pathway, but all four
pathways can be present

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Light-Dependent Pathway

• Also termed the photoperiodic pathway


• Keyed to changes in the proportion of light to dark in the
daily 24-hr cycle (day length)
• Short-day plants flower when daylight becomes shorter
than a critical length
• Long-day plants flower when daylight becomes longer
than a critical length
• Day-neutral plants flower when mature regardless of day
length

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Day Length Affects Flowering

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Obligate and Facultative Plants

In obligate long- or short-day plants there is a sharp


distinction between short and long nights, respectively
In facultative long- or short-day plants, the photoperiodic
requirement is not absolute
• Flowering occurs more rapidly or slowly depending on the
length of day.

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Manipulation of Photoperiod

• Using light as a cue allows


plants to flower when abiotic
conditions are optimal
• Manipulation of photoperiod
in greenhouses ensures that
short-day poinsettias flower
in time for the winter holidays

Don Hammond/Design Pics

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Phytochrome and Cryptochrome

Conformational change in a phytochrome (red-light sensitive)


or cryptochrome (blue-light sensitive) light-receptor molecule
triggers a cascade of events that leads to the production of a
flower
In Arabidopsis, regulate via the gene CONSTANS (CO)
which encodes a transcription factor that turns on genes that
are needed for flowering
• This signaling cascade leads to expression of LFY
Phytochrome regulates the transcription of CO

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CONSTANS

CO protein is produced day and night


The levels of CO are maintained in accordance with the
circadian clock
• Levels of CO mRNA are lower at night because of
targeted protein degradation by ubiquitin
• Phytochrome causes an increase in transcription at
daybreak
• Cryptochrome prevents degradation by the ubiquitin-
dependent pathway during the day

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Phytochrome increases CO level mRNA at daybreak
Cryptochrome (blue light receptor) prevents degradation of
CO protein

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Temperature-Dependent Pathway

Some plants require a period of chilling before flowering –


vernalization
• Described in the 1930s by Ukrainian scientist T.D.
Lysenko.
• Winter wheat would not flower without a period of chilling.
• Seeds could be chilled and planted in the spring.

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Gibberellin-Dependent Pathway

Decreased levels of gibberellins have been shown to delay


flowering in some species.
Gibberellin has been shown to bind to the promoter of the
LFY gene, which supports a model where gibberellin
induces an increase in LFY gene expression.
• This would directly affect flowering.

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Autonomous Pathway

• Does not depend on external cues except for basic


nutrition
• Presumably delays flowering
• A balance between floral promoting and inhibiting signals
may regulate when flowering occurs
• Can test determination for flowering by changing the
environment and ascertaining whether developmental fate
has changed

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Flowering Pathways

The four flowering pathways lead to an adult meristem


becoming a floral meristem
• Activate or repress the inhibition of floral meristem identity
genes.
The floral meristem identity genes: LFY and AP1
• Turn on floral organ identity genes.
• Define the four concentric whorls.
• Sepal, petal, stamen, and carpel.

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Model for Flowering

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The Floral Meristem and the Flower

A floral meristem is a modified shoot apical meristem that produces


flowers
A flower contains four kinds of organs: sepals, petals, stamens, and
carpels
• Each of these organs is arranged in a characteristic pattern of whorls
within whorls
How does the floral meristem produce these four organs in their
characteristic arrangement?

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Figure 38.21

(a) Whorls of cells in (b) Whorls of organs in flower


floral meristem

3 4
2 4
1 2 3
1

Sepal Petal Stamen Carpel

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The Genetic Control of Flower Structures

Several types of mutant flowering plants are homeotic mutants in which


one kind of floral organ is replaced by another
Homeotic gene is responsible for the development of a certain body
structure
Elliot Meyerowitz and colleagues found that Arabidopsis floral homeotic
mutants can be divided into three general classes
• Some had only carpels and stamens; others had only sepals and
carpels; others had only petals and sepals
• Each type of mutant lacks the elements normally found in two of the
four whorls

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Figure 38.22

Wild type Mutant class 1 Mutant class 2 Mutant class 3


Normal arrangement of organs Only carpels and stamens Only sepals and carpels Only petals and sepals

Whorl 4: Carpel
Whorl 3: Stamen
Whorl 2: Petal
Whorl 1: Sepal

34 34 44 21
12 43 11 12

Sepal Petal Stamen Carpel Carpel Stamen Stamen Carpel Sepal Sepal Carpel Carpel Sepal Petal Petal Sepal

© 2017 Pearson Education, Ltd.


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The ABC Model

The biologists hypothesized that each class of homeotic mutant was


caused by a defect in a single gene
• Their hypothesis for genetic control of flower development is called
the ABC model

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The ABC Model

Three basic ideas underlie the ABC model:


1. Each of the three genes involved is expressed in two adjacent
whorls
2. A total of four different combinations of gene products can occur
3. Each of the four combinations of gene products triggers the
development of a different floral organ

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The ABC Model

The Meyerowitz group proposed that


• The A protein alone causes cells to form sepals
• A combination of A and B proteins sets up the formation of petals
• B and C combined specify stamens
• The C protein causes cells to form carpels
• The A protein inhibits production of the C protein
• The C protein inhibits production of the A protein

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Testing the ABC Model

Researchers tested the ABC model, determining the pattern of


expression of the A, B, and C genes
• They found that A genes are expressed in the outer two whorls, B
genes are expressed in the middle two whorls, and C genes are
expressed in the inner two whorls
Thus A, B, and C genes were expressed in the predicted regions,
supporting the ABC model

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Figure 38.23

(a) The ABC model Whorl 1 2 3 4

A A
Idea 1: The products of three genes
Active genes B B pattern the flower; each gene is expressed
in two adjacent whorls.
C C

Assume: Resulting proteins A A+B B+C C Idea 2: Four different combinations


Protein A inhibits gene C (no A + C) of proteins occur.
Protein C inhibits gene A
Sepal Petal Stamen Carpel Idea 3: Each protein combination triggers
Floral organ
(Se) (Pe) (St) (Ca) development of a different floral organ.

(b) Predictions of the ABC model


Wild type: ABC intact Gene A missing Gene B missing Gene C missing
Whorls 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4
A A A A A A A A
Active B B B B B B
genes
C C C C C C C C

Identity of
floral organ
in whorl
Se Pe St Ca Ca St St Ca Se Se Ca Ca Se Pe Pe Se

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Flower Structure

Floral organs are thought to have evolved from leaves


A complete flower has four whorls
• Calyx, corolla, androecium, and gynoecium.

An incomplete flower lacks one or more of the whorls

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Flower Morphology

Calyx = Consists of flattened sepals


Corolla = Consists of petals
Androecium = Collective term for all the stamens (male
structures) of a flower
• Stamen consists of a filament and an anther.
Gynoecium = Collective term for all carpels (female
structures) of a flower
• Carpel consists of ovary, style, and stigma.
• Ovules produced in ovary.

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Complete Flower

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Trends in Floral Specialization

2 major trends
1. Floral parts have grouped
together
2. Floral parts lost or
reduced
Modifications often relate to
pollination mechanisms

Dr. Thomas Barnes/University of Kentucky/USFWS

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Trends in Floral Symmetry

• Primitive flowers are radially symmetrical


• Advanced flowers are bilaterally symmetrical

©Traci Tatman

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Genetic Regulation of Asymmetry

Courtesy of Enrico Coen

Snapdragon flowers normally have bilateral symmetry, like the one


shown here on the left. The CYCLOIDEA gene regulates floral symmetry,
and cycloidea mutant snapdragons, like the one on the right, have
radially symmetrical flowers.
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Gamete Production

Alternation of generations
Diploid sporophyte → haploid gametophyte
In angiosperms, the gametophyte generation is very small
and is completely enclosed within the tissues of the parent
sporophyte
• Male gametophyte – pollen grains.
• Female gametophyte – embryo sac.

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Reproductive Organs of Angiosperms

Gametes are produced in separate, specialized structures of


the flower
Reproductive organs of angiosperms differ from those of
animals in two ways
1. Both male and female structures usually occur together in
the same individual
2. Reproductive structures are not permanent parts of the
adult individual

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Gametophyte Formation

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Pollen Formation

• Anthers contain four microsporangia which produce


microspore mother cells (2n)
• Microspore mother cells produce microspores (n) through
meiosis
• Microspore develops by mitosis into pollen
• Generative cell in the pollen grain will later divide to form
two sperm cells

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Pollen Grains

©L. DeVos/Free University of Brussels

In the Easter lily, Lilium candidum, the pollen tube emerges from the
pollen grain through the groove or furrow that occurs on one side of the
grain. In a plant of the sunflower family, Hyoseris longiloba, three pores
are hidden among the ornamentation of the pollen grain. The pollen tube
may grow out through any one of them.
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Embryo Sac Formation

• Within each ovule, a diploid megaspore mother cell


undergoes meiosis to produce four haploid megaspores
• Usually only one megaspore survives
• Enlarges and undergoes repeated mitotic divisions to
produce eight haploid nuclei
• Enclosed within a seven-celled embryo sac

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Mature Embryo Sac

(right) K. Stern’s

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Pollination

Process by which pollen is placed on the stigma


Self-pollination
• Pollen from a flower’s anther pollinates stigma of the same
flower, or the stigma of another flower on the same plant
Cross-pollination
• Pollen from anther of one flower pollinates another
flower’s stigma.
• Also termed outcrossing.

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Self vs cross pollination

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Successful Pollination

• Successful pollination in many angiosperms depends on


regular attraction of pollinators
• Floral morphology has coevolved with pollinators
• Early seed plants wind pollinated
• Among insect-pollinated angiosperms, the most numerous
groups are those pollinated by bees

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Bee Pollination

• Bees typically visit yellow or blue flowers


• Many have stripes or lines of dots that indicate the location
of the nectaries

Holly Hildreth/McGraw Hill

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How a Bee Sees a Flower

©Thomas Eisner, Cornell University

The yellow flower of Ludwigia peruviana (Peruvian primrose)


photographed in normal light (on the left) appears yellow, but under
ultraviolet light it has a conspicuous central bull’s-eye. This is because
the outer sections of the petals reflect both yellow and ultraviolet, while
the inner portions reflect yellow only and therefore appear dark in the
photograph that emphasizes ultraviolet reflection.
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Other Insect Pollinators

• Flowers that are visited regularly by butterflies often have


flat “landing platforms”
• Flowers that are visited regularly by moths are often white
or pale in color
• Also tend to be heavily scented, making them easy to
locate at night

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Bird Pollination

Flowers that are visited regularly by birds must produce large


amounts of nectar
Often have a red color
• Conspicuous to birds, but usually inconspicuous to insects.

Adam Jones/Getty Images

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Other Animal Pollinators

• Signals are species-specific


• Small rodents may pollinate plants
• Bats, bird, and insects pollinate saguaro cacti

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Wind Pollination

Some angiosperms are wind-pollinated


• Characteristic of early seed plants.
Flowers are small, green, and odorless, with reduced or
absent corollas
Often grouped and hanging down in tassels
Stamen- and carpel-containing flowers are usually separated
between individuals
• Strategy that greatly promotes outcrossing.

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Staminate and Pistillate Flowers

Guenter Fischer/Imagebroker/Alamy Stock Photo

Birches (Betula sp) are monoecious; their staminate flowers hang down
in long, yellowish tassels, and their pistillate flowers mature into clusters
of small, brownish, conelike structures.
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Wind-pollinated Flowers

Noppharat4969/Shutterstock

The large yellow anthers, dangling on very slender filaments, are hanging out,
about to shed their pollen to the wind. Later, these flowers will become pistillate,
with long, feathery stigmas – well-suited for trapping windblown pollen – sticking
far out of them. Many grasses, like this one, are therefore dichogamous.
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Self-Pollination

Outcrossing is generally advantageous for plants and for


eukaryotic organisms
Nevertheless, self-pollination also occurs in some
angiosperms
2 basic reasons for frequency of self-pollination
1. Self-pollination is favored in environments where
pollinators are scarce
2. Offspring are more uniform and probably better adapted
to their environment, which is favored in stable
environments

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Promotion of Outcrossing

Several evolutionary strategies promote outcrossing


• Separation of stamens and pistils in space.
• Dioecious plants produce only ovules or only pollen.
• Monoecious plants produce male and female flowers on the same
plant, but they may mature at different times (dichogamy).

• Self-incompatibility that prevents self-fertilization.

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Dichogamy
the ripening of the stamens and pistils of a flower at different
times, so that self-fertilization is prevented.

(a) Heiti Paves/Alamy Stock Photo; (b) Rafael Campillo/agefotostock

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Self-incompatibility

Self-incompatibility increases outcrossing


Pollen and stigma recognize each other as being genetically
related and pollen tube growth is blocked
Controlled by alleles at the S locus
2 types of self-incompatibility
1. Gametophytic self-incompatibility
• Depends on the haploid S locus of the pollen and the diploid S
locus of the stigma.
2. Sporophytic self-incompatibility
• If the alleles in the stigma match either of the pollen parent’s S
alleles, the haploid pollen will not germinate.

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Comparing Types of Self-Incompatibility

Determined by the Determined by the genotype


haploid pollen genotype of the diploid pollen parent

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Double Fertilization

Only in angiosperms
Requires two sperm cells
Double fertilization results in two key developments
• Fertilization of the egg.
• Formation of endosperm that nourishes the embryo.

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Process of Double Fertilization 1

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Process of Double Fertilization 2

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Embryo Development

Begins once the egg cell is fertilized


The growing pollen tube enters angiosperm embryo sac and
releases two sperm cells
• One sperm fertilizes central cell with its polar nuclei and
initiates endosperm development.
• Other sperm fertilizes the egg to produce a zygote.
• Cell division soon follows, creating the embryo.

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First Zygotic Division

First zygote division is asymmetrical, resulting in cells with 2


different fates
• Small cell divides repeatedly forming a ball of cells, which
will form the embryo.
• Large cell divides repeatedly forming an elongated
structure called a suspensor.
• Transports nutrients to embryo.

The root–shoot axis also forms at this time


• Cells near suspensor become root.
• Cells at the other end become shoot.
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Early Cell Divisions 1

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Early Cell Divisions 2

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The Suspensor (sus) Mutant

Edward Yeung, University of Calgary and David Meinke, Oklahoma State University

This suspensor (sus) mutant of Arabidopsis has a defect in embryo development.


Inhibition of embryo development in the suspensor is blocked, resulting in
embryo-like development of the suspensor. SUS, an allele normally present in the
embryo, is required to suppress embryo development in suspensor cells

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Body Plan

In plants, three-dimensional shape and form arise by


regulating amount and pattern of cell divisions
• Vertical axis (root–shoot axis) becomes established at a
very early stage.
• Same is true for establishment of a radial axis (inner–outer
axis).
First cell division gives rise to a single row of cells, cells soon
begin dividing in different directions, producing a solid ball of
cells

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Apical Meristems

Apical meristems establish the root–shoot axis in the globular


stage, from which the three basic tissue systems arise
• Dermal.
• Ground.
• Vascular tissue.

These tissues are organized radially around the root–shoot


axis

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SHOOTMERISTEMLESS

• SHOOTMERISTEMLESS (STM) needed for shoot formation


• STM gene codes for a transcription factor with a homeobox region
• The stm mutant of Arabidopsis has a normal root meristem but fails to
produced a shoot meristem between its two cotyledons.

©Jan Lohmann, Max Planck Institute for Developmental Biology

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Auxin

One way that auxin induces gene expression is by activating


a transcription factor
• MONOPTEROS (MP) is a gene that codes for an auxin-
induced transcription factor.
• Necessary for root formation, but not shoot.

• Once activated, MP protein binds to the promoter of


another gene, leading to transcription of a gene or genes
needed for root meristem formation.

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HOBBIT Gene Action

HOBBIT (HOB) mutation affects root development

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MONOPTEROS Gene Action

MONOPTEROS mutation affects root development

Hobbit
represses the
production of
the repressor of
auxin-induced
genes

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HOBBIT and MONOPTEROS

(c-e) Ben Scheres, University of Utrecht

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Formation of Tissue Systems

Primary meristems differentiate while the plant embryo is still


at the globular stage
• No cell movements are involved.

Outer protoderm develops into dermal tissue that protects


the plant
Ground meristem develops into ground tissue that stores
food and water
Inner procambium develops into vascular tissue that
transports water and nutrients

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Morphogenesis

The globular stage gives rise to heart-shaped embryo with


bulges called cotyledons
• Two in eudicots and one in monocots.

These bulges are produced by embryonic cells, and not by


the shoot apical meristem
• This process is called morphogenesis.
• Results from changes in planes and rates of cell division.

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Plant Body Form

Form of a plant body is largely determined by the plane in


which its cells divide
• Also controlled by changes in cell shape as cells expand
due to turgor pressure after they form.
Based on the position of the cell plate
• Determined by microtubules and actin.
• Microtubules also guide cellulose deposition as the cell
wall forms around the new cell.
• Cells expand in the directions of the two sides with the least
cellulose reinforcement.

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Cell Division

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Cell Expansion

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Early Embryonic Development

Early in embryonic development, most cells can give rise to a


wide range of cell and organ types, including leaves
• As development proceeds, the cells with multiple
potentials are restricted to the meristem regions.
• Many meristems have been established by the time
embryogenesis ends and the seed becomes dormant.
After germination, apical meristems continue adding cells to
the growing root and shoot tips

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Critical Developmental Events

During embryogenesis, angiosperms undergo three other


critical events
1. Development of a food supply
2. Development of seed coat
3. Development of fruit surrounding seed

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Endosperm Variation

The sporophyte transfers nutrients via the suspensor to the


endosperm in angiosperms
Endosperm varies between plants
• In coconuts it includes the liquid “milk”.
• In corn it is solid.
• it expands with heat to form the white edible part of popcorn.

• In peas and beans it is used up during embryogenesis.


• Nutrients are stored in thick, fleshy cotyledons.

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Endosperm

(Top right) Somchai Som/ Shutterstock; (Bottom right) Metta image/Alamy Stock Photo

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Seeds

• In many angiosperms, development of the embryo is


arrested soon after meristems and cotyledons differentiate
• Integuments develop into a relatively impermeable seed
coat
• Encloses the seed with its dormant embryo and stored
food

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Initiation of Germination

• Germination: the emergence of the radicle (first root)


through the seed coat
• Germination cannot take place until water and oxygen
reach the embryo
• Stratification: some seeds require periods of time at low
temperatures before germination

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Food Storage in the Seed

• Germination and early seeding growth require the


utilization of metabolic reserves stored as starch in
amyloplasts and protein bodies
• Depending on the kind of plant, these reserves may be
stored in the embryo or in the endosperm
• Scutellum: in kernels of cereal grains, the single cotyledon
is modified into this structure, which transfers nutrients
from the endosperm to the embryo

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Hormonal Effects

In response to the absorption of water by a seed, the embryo


produces gibberellic acid
This signals the aleurone (the outer layer of the endosperm)
to produce α-amylase
• This enzyme is responsible for breaking down starch.
Levels of abscisic acid, which inhibits starch breakdown, may
be reduced when a seed absorbs water

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Hormonal Regulation of Germination

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Germination

(a1) Nigel Cattlin/Alamy Stock Photo; (b1) Martin Shields/Alamy Stock Photo

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Asexual Reproduction

Produces genetically identical individuals because only


mitosis occurs
More common in harsh, unchanging environments
• All clones are adapted.
• Variations may not be adapted.

Should conditions change dramatically, there will be less


variation in the population for natural selection to act on, and
the species may be less likely to survive.
Used in agriculture to propagate a particularly desirable plant
with traits that would be altered by sexual reproduction

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Apomixis

• Apomixis – asexual development of a diploid embryo in


the ovule
• Offspring is genetically identical to the sporophyte that
produced it
• Gain advantage of seed dispersal usually associated with
sexual reproduction.

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Vegetative Reproduction

New plant individuals are


cloned from parts of adults
Comes in many and varied
forms
• Runners or stolons.
• Rhizomes.
• Suckers.
• Adventitious plantlets.
Jerome Wexler/Science Source

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Plant Life Spans

Once established, plants live for variable periods of time,


depending on the species
Woody plants, which have extensive secondary growth,
typically live longer than herbaceous plants, which don’t
• Bristlecone pine, for example, can live upward of 4,000
years.
Depending on the length of their life cycles, herbaceous
plants may be annual, biennial, or perennial

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Annual, Biennial, or Perennial

Annual plants grow, flower, and form fruits and seeds within
one growing season
• They then die when the process is complete.

Biennial plants have life cycles that take two years to


complete
• Store energy in year one, flower in year two.

Perennial plants continue to grow year after year


• They may be herbaceous or woody.

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Annual and Perennial Plants

(a) Anthony Arendt/Alamy Stock Photo; (b) SuperStock/Alamy Stock Photo

Desert annuals complete their entire life span in a few weeks, flowering just once.
Some trees, such as the giant redwood, Sequoiadendron giganteum, which
occurs in scattered groves along the western slopes of the Sierra Nevada in
California, live 2000 years or more, and reproduce year after year.
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