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Hormonal Regulation of Plant Growth and Development

Chapter 39

Growth and Development

Photosynthesis, respiration, and transpiration are the three major functions that drive plant growth and development

How well a plant is able to regulate these functions greatly affects its ability to grow, compete, and reproduce

Control of Growth and Development

Internal mechanisms govern plant growth and development

Changes in environment turn such mechanisms on or off at different times


To sense environmental cues and to respond appropriately plants rely on signaling molecules like hormones

Summary of Embryo Development

After differentiation is complete, the embryo consists of four major parts:


Cotyledon(s) is the source of nutrients during embryo development Epicotyl is the embryonic shoot Hypocotyl is below the cotyledons and contributes to stem development Radicle is below the hypocotyl and contributes to root development

Hypophysis is the uppermost cell of suspensor, it contributes to root formation The development of each part of the embryo depends on gradients of plant hormone, auxin Cotyledons
Epicotyl Hypocotil Radicle Hypophysis

Structure of Plant Seed

Monocots versus Eudicots

Monocot embryo has only one cotyledon, a dicot embryo has two In monocots, the cotyledon rarely stores food In dicots, cotyledons usually strore the nutrients In monocots, the cotyledon passes the food from endosperm to the embryo In dicots, as food reserves are used up the cotyledons wither and fall off

Seed Germination

As seed matures, it dehydrates and enters a phase referred to as dormancy

Seed dormancy increases the chances that germination will occur at a time or place most favorable to the seedling Germination is a process by which the plant embryo resumes growth after seed dispersal

Depends on internal factors

Abscisic acid (ABA) - a germination inhibitor within a seed

Depends upon environmental factors

Temperature

Soil moisture
Oxygen levels

Splitting the Seed Coat

Germination of seeds is initiated by the process called imbibition, which is the uptake of water

Attracted by hydrophilic groups of storage proteins water molecules move into a seed

As water moves in, the seed swells and the coat ruptures

More oxygen reaches the embryo Aerobic respiration provides energy for growth

Growth of a Bean Plant


(Eudicot)
radicle

The first organ to emerge is radicle

true leaf

Then, the hypocotyl breaks through the soil surface

It is bent in a hairpin shape

epicotyl hypocotyl withered cotyledon

Stimulated by light, the hypocotyl straightens and brings up the cotyledons and epicotyl As the true leaves begin making food by photosynthesis

primary root two cotyledons

the cotyledons wither and fall off

branch root

Growth of a Corn Plant


(Monocot)

Monocots use a different method for breaking ground when they germinate Primary shoot (epicotyl) pushes upward through the soil and into the air
seed coat coleoptile

foliage leaves

coleoptile

radicle
radicle radicle

Growth of a Corn Plant


(Monocot)

The primary shoot is protected by the coleoptile a hollow tube When coleoptile reaches the surface, it stops growing and the true leaf breaks out through the coleoptile tip

seed coat coleoptile

foliage leaves

coleoptile

radicle
radicle radicle

Coleoptile of Monocots

first foliage leaf

coleoptile

Coleoptile of Monocots

The coleoptiles of monocots have been a favorite experimental object for studying phototropism Tropism is a growth response that results in a movement of plant organs either toward or away from the stimulus

Phototropism is a directional growth toward light

Phototropism and Darwin

One example of phototropism is

the bending of a grass seedling toward light

In late 19th century, Charles Darwin and his son Francis

conducted some of the earliest experiments on phototropism

Phototropism and Darwin


EXPERIMENT In 1880, Charles Darwin and his son Francis designed an experiment to determine what part
of the coleoptile senses light.

RESULTS
Control Shaded side of coleoptile Light Darwin and Darwin (1880)

Light

Illuminated side of coleoptile

Tip removed

Tip covered by opaque cap

Tip covered by transparent cap

Base covered by opaque shield

CONCLUSION In the Darwins experiment, a phototropic response occurred only when light could
reach the tip of coleoptile. Therefore, they concluded that only the tip senses light.

Phototropism and Boysen-Jensen


EXPERIMENT In 1913, Peter Boysen-Jensen conducted an experiment to determine how the signal for
phototropism is transmitted.

RESULTS
Control Shaded side of coleoptile Light

Boysen-Jensen (1913)

Light

Illuminated side of coleoptile

Tip separated by gelatin block

Tip separated by mica

CONCLUSION

Boysen-Jensen observed that a phototropic response occurred if the tip was separated by a permeable barrier (gelatin), but not if separated by an impermeable solid barrier (a mineral called mica). These results suggested that the signal is a light-activated mobile chemical.

Genes Govern Development

All cells in a plant inherit the same set of genes and, thus, retain the ability to produce a whole individual
(e.g. tissue culture propagation)
Undifferentiated callus

Single parenchyma cell

Genes Govern Development

During development, some plant cells differentiate, others continue to divide


a) positional

differences and b) unequal cytoplasmic divisions lead to


differences in metabolic output selective gene expression

As a result, daughter cells begin to vary and specialized tissues start to form

Hormones and Development

Hormones are signal molecules that coordinate changes in different parts of an organism

Being produced at specific locations, they alter processes in target cells at other locations
Though produced at very low concentrations, even a minute amount of hormonal signal can have a profound effect on growth and development of a plant organ

Hormones and Development

In their action hormones interact

with each other, with gene products and with the environmental cues flowering, aging, root and stem growth, color of fruits and many other conditions

Hormones can affect

Differential expression of genes regulated by hormones

starts cell lineages down different developmental pathways


(e.g. meristematic vs permanent)

Plant Hormones
The six major classes of plant hormones are:

Auxins

Gibberellins

Cytokinins

Brassinosteroids Abscisic Acid

Ethylene

Hormonal Response in Plants

Plants have cellular receptors

that they use to detect hormonal signals or important changes in the environment certain cells must have an appropriate receptor

For a stimulus to elicit a response

Signal transduction is a pathway that links signals, receptors, and plant cell responses

Signal Transduction

Hormonal signals are detected by receptors that change in response to specific stimuli
CELL WALL 1 Reception CYTOPLASM

Relay molecules Receptor

Activation of cellular responses

Hormone or environmental stimulus

Plasma membrane

Signal Transduction

Activated receptor activates other molecules

Second messengers transform and amplify signal from a receptor and bring it to proteins that cause specific responses
CELL WALL 1 Reception CYTOPLASM

2 Transduction

3 Response

Relay molecules Receptor

Activation of cellular responses

Hormone or environmental stimulus

Plasma membrane

Signal Transduction

Ultimately, signal transduction leads to regulation

of certain cellular activities by stimulating specific proteins (enzymes, transporters, transcription factors)
CELL WALL 1 Reception CYTOPLASM

2 Transduction

3 Response

Relay molecules Receptor

Activation of cellular responses

Hormone or environmental stimulus

Plasma membrane

Plant Hormones

Gibberellins

More than 120 forms have been isolated from plants

Although a much smaller number occur in each plant species

Chemically all gibberellins are diterpenoid acids

Gibberellins

Gibberellins are synthesized in seeds and young shoot tissues Their synthesis is increased in the dark, when photosynthesis is arrested, and decreased in the light

Gibberellins

Natural gibberellins have a variety of effects

Help seeds and buds break dormancy Influence stems elongation Affect flowering and fruit growth

Gibberellins

Gibberellins are used commercially

Applied by growers to delay ripening and increase fruit size

control

GA sprayed during fruit development

Gibberellins

Gibberellins stimulate growth of both leaves and stems

In stems gibberellins stimulate cell elongation and division

Gibberellins

Gibberellin was discovered as a substance secreted by a parasitic fungi (Gibberella fujikuroi) that causes rice bakanae disease (from Japanese foolish seedling)

Plants grow fast, look spindly and pale and break off easily Compare foolish cabbages that were treated with GA

GA Action in Germination

After water is imbibed,

the release of gibberellins from the embryo signals the seed to break dormancy and germinate

Barley Seed Germination

endosperm

seed coat aleurone

Aleurone is a specialized layer of cells in the endosperm During barley seed germination, Gibberellin performs its act in cells of the aleurone

large cotyledon

embryo sporophyte

GA Action in Germination

After a seed imbibes water, the embryo releases GA as a signal to the aleurone

Aleurone

Cotyledon

GA

GA

Water

Endosperm

Radicle

GA Action in Germination

Aleurone responds by synthesizing and secreting digestive enzymes (e.g. a-amylase) that hydrolyze nutrients stored in the endosperm

Aleurone

Cotyledon
a-amylase GA

GA

Water

Endosperm

Radicle

GA Action in Germination

Sugars and other nutrients absorbed by cotyledon are transferred to the embryo

Sugar monomers fuel aerobic respiration ATP from the aerobic respiration provides energy for growth of the primary root and shoot
Aleurone

Cotyledon
a-amylase GA

Coleoptile
Sugar

GA

Water

Endosperm

Radicle

Auxins

The term auxin is used for any chemical that promotes cell elongation in different tissues

Indole Acetic Acid (IAA) is the most important natural auxin

It is produced in cells of apical meristem both in shoots and roots Auxins play an essential role in coordination of many growth and behavioral processes Auxins typically act in concert with (or in opposition to) other plant hormones

Auxins

Promote stem lengthening Play a role in responses to gravity and light Apical dominance

The main stem grows more strongly than the side stems Auxin from the apical bud inhibits the development of lateral bud growth

Root growth and development Fruit growth

Wounding response
An overdose of auxins can kill a plant

Certain synthetic auxins are used as herbicides

Auxins

Auxin is involved in the formation and branching of roots

When planted into medium pretreated with auxin, plants develop more roots
control plant auxin treated

Auxin and Embryo Patterning


(Friml et al, Nature 2003)

Axis formation and patterning

occur in plants, as in animals, during early embryogenesis

The first mitotic division of zygote is transverse


Creates the apical-basal axis Splits the fertilized egg into a larger basal cell and a smaller terminal cell Basal cell develops into suspensor Terminal cell becomes a proembryo

Auxin and Embryo Patterning


(Friml et al, Nature 2003)

The pattern formation becomes apparent at the globular stage, when

The outer layer of cells differentiates into the protoderm

Two layers of inner cells have distinct developmental fates

Upper layer produces apical meristem and cotyledons Lower layer produces the hypocotyl the founder of root meristem

Hypophysis is

Auxin and Embryo Patterning


(Friml et al, Nature 2003)

Major players are PIN proteins, auxin transporters: PIN1, PIN4, PIN7

PIN trans-membrane proteins control the auxin flow Localization of PIN proteins on a cell membrane determines the direction of auxin transport apical cells accumulate maternally derived auxin through PIN7 transport from basal cells Apical pole specification and Proembryo and suspensor differentiation

At the two-cell stage,

This results in

Auxin and Embryo Patterning


(Friml et al, Nature 2003)

At the globular stage

auxin production starts in the apical region of proembryo The PIN7-dependent transport of auxin is reversed and now directed toward the suspensor In addition, PIN1 and PIN4 also mediate auxin transport out of the embryo

As a result, auxin gradient reverses

displaying new maximum in the uppermost suspensor cell, hypophysis, thus, triggering root pole specification (root meristem)

Auxin Action in Shoot Growth

In monocots, primary leaf is protected by coleoptile In 1880, Charles Darwin showed that the tip of coleoptile drives shoot growth In 1913, Boysen-Jensen showed that a light-activated mobile chemical is first produced in the tip of coleoptile and then transmitted to the base of the shoot

coleoptile first foliage leaf

Auxin Action in Shoot Growth

In 1926, Frits Went showed that the growth-promoting chemical (which he named auxin) produced by cells in the tip causes elongation of cells below

A seedling with its tip cut off will not lengthen Application of auxin extracted from the tip on the top of damaged coleoptile restores the growth

A modified experiment of Frits Went addressing phototropism


Excised tip on agar cube

Growth-promoting chemical diffuses into agar cube

Control

Control (agar cube lacking chemical)

Offset cubes

Auxin Action in Shoot Growth


Auxin activates shoot growth by promoting cell elongation According to a model called the Acid Growth Hypothesis,

auxin activates enzymes that break cross-links between cellulose microfibrils in the cell wall

Now cells can expand, as turgor pressure builds up inside


Cross-linked cellulose microfibrils in primary cell wall

expansion under turgor pressure

Acid Growth Hypothesis


Parenchyma cells transport auxin from apical meristem downward to the region of cell elongation The AUX1 protein transports auxin in, and PIN protein transports auxin out Proton pumps (H+ pump) play a major role in the model
cell wall (pH 5)

AUX1 protein

cytoplasm of parenchyma cell (pH 7)

H+ pump

PIN protein

IAA H+

Auxin Transport
H+

IAAH H+ IAA + H+ H+

As auxin (IAA: Indol Acetic Acid) diffuses through the acidic cell wall it binds proton H+ AUX1 transports non-ionized form of auxin into cytoplasm Within the cell, at pH7 auxin looses proton H+ and PIN transports auxin out

H+

IAA
H+ H+

H+
IAAH H+ IAA + H+ H+ H+

IAA H+

Acid Growth Hypothesis


H+

IAAH H+ IAA + H+ H+

As it moves down, in each cell auxin increases the activity of proton pumps

H+

H+ cations pumped out of cell lower the pH in the cell wall


Acidification of the wall activates expansins that separate cellulose microfibrils, thus, allowing enzymes to break the cross-links between them

IAA
H+ H+

H+
IAAH H+ IAA + H+ H+ H+

Acid Growth Hypothesis


H2O
Plasma membrane Cell wall

When the cross-links are broken, the microfibrils become loose and able to slide Thus, the flexibility of the cell wall is increased Inside turgor pressure and wall plasticity allow cells to elongate


Nucleus Cytoplasm Vacuole

Cytokinins

Promote cell division

Called cytokinins because they stimulate cell division i.e. cytokinesis

Most abundant in root and shoot meristems and in maturing fruits

In mature plants, produced in roots and transported to shoots through xylem


Used to artificially extend the shelf life of cut flowers; delays leaf death Opposes effects of auxin by promoting growth of lateral buds

Ethylene

Ethylene is a gaseous hormone

It is synthesized in nodes of stems


The biosynthesis of ethylene can be induced by ethylen itself, auxin and cytokinin; it is inhibited by the abscisic acid, however The environmental cues can also induce ethylene formation, e.g. plants produce ethylene in response to stresses

Drought Flooding Mechanical pressure Injury or infection

The triple response to mechanical stress

Under mechanical stress such as seedling growing into an obstacle, ethylene induces triple response

Slowing of stem elongation


Thickening of the stem Horizontal growth

The triple response allows a growing shoot to avoid obstacles


In response to ethylene, seedling undergo triple response slowing of stem elongation, stem thickening, and horizontal stem growth. The response is greater with higher ethylene concentration.

0.00

0.10

0.20

0.40

0.80

Ethylene concentration (parts per million)

Arabidopsis triple response mutants

Studies of triple response mutants illustrate the identification of signal transduction pathway

(a) Ethylene insensitive

(b) Constitutive triple-response

Arabidopsis triple response mutants

Some mutants are ethylene insensitive (ein) and fail undergo the triple response after exposure to ethylene Why?

(a) ein mutant

Arabidopsis triple response mutants

Some mutants are ethylene insensitive (ein) and fail undergo the triple response after exposure to ethylene They lack a functional ethylene receptor
(a) ein mutant

Arabidopsis triple response mutants

Some mutants (ctr) undergo the triple response even when there are no physical obstacles Why?

(b) ctr mutant

Arabidopsis triple response mutants

Some mutants (ctr) undergo the triple response even when there are no physical obstacles In constitutive triple-response (ctr) mutants, ethylene signal transduction is permanently turned on
(b) ctr mutant

Ethylene

Ethylene is involved in abscission, the dropping of leaves, fruits, or flowers It ripens fruits by increasing activity of enzymes that soften fruit

It was an early practice to prepare citrus for market by storage in a room with kerosene stove. It turned out that incomplete combustion of kerosene produced ethylene which ripens fruits A barrel of ripening apples can induce ripening of a bunch of bananas some distance away

Abscisic Acid (ABA)

Produced in root cells and transferred to leaves by the transpiration stream (xylem)

but could be synthesized by all cells

ABA is called stress hormone it causes the suspension of growth; promotes dormancy of buds and seeds Used commercially to induce dormancy in plants to be shipped

Abscisic Acid (ABA)

ABA plays a role in drought response (causes closure of stomata) ABA binds receptor on the guard cell membrane Chloride and potassium ions flow out Water follows, turgor pressure drops and stomata close

Plant Hormone Applications

Plant Tropisms
Adjustment of

plant growth toward or away

from an environmental stimulus


Phototropism -

stimulus is light Gravitropism - stimulus is gravity Thigmotropism - stimulus is contact with an object

Gravitropism

Roots tend to grow toward pull of gravity; shoots grow against it Gravitational field is sensed via position of statoliths (cell organelles that accumulate starch)

Statoliths

20 m

due to gravity statoliths settle at the lowest part of cell


(a) (b)

Thigmotropism

Growth in response to contact with a solid object

Allows vines and tendrils to wrap around supports


Touched cells produce auxin and transport it to untouched cells

Cells on untouched side elongate faster, causing stem to curl

Phototropism

Change in growth in response to light Light cues many critical events in plant growth and development

Phototropism

Plants not only detect the presence of light

But also its direction, intensity, and wavelength (color)

A graph called an action spectrum

Depicts the relative response of a process to different wavelengths of light


Are useful in the study of any process that depends on light

Action spectra

Action Spectrum
1.0 0.8 Phototropic effectiveness

0.6
0.4 0.2

The graph illustrates that only light wavelength below 500 nm induce response

400

450

500

550

600

650

700

Wavelength (nm) (a) Phototropism action spectrum Light Time 0 min

The color of the light source is indicated on coleoptiles

Time 90 min (b) Coleoptiles before and after light exposures

Phototropism

Research on action spectra and absorption spectra of pigments

Led to the identification of two major classes of light receptors: blue-light photoreceptors and phytochromes

Phototropism

Various blue-light photoreceptors

Control hypocotyl elongation, stomatal opening, and phototropism Regulate many of a plants responses to light throughout its life

Phytochromes

Phytochromes

A phytochrome is the photoreceptor responsible for the opposing effects of red and far-red light
A phytochrome consists of two identical proteins joined to form one functional molecule. Each of these proteins has two domains.

Chromophore

Photoreceptor activity. One domain, which functions as the photoreceptor, is covalently bonded to a nonprotein pigment, or chromophore.

Kinase activity. The other domain has protein kinase activity. The photoreceptor domains interact with the kinase domains to link light reception to cellular responses triggered by the kinase.

Phytochromes

Phytochromes exist in two photoreversible states

the conversion of Pr to Pfr triggers many developmental responses

Pr

Pfr

Red light Synthesis


Responses: seed germination, control of flowering, etc. Far-red light

Slow conversion in darkness (some plants)

Enzymatic destruction

Signal Transduction Activated by Light


1 Reception
CYTOPLASM Plasma membrane

2 Transduction

3 Response
Transcription factor 1 NUCLEUS

cGMP
Second messenger produced

Specific protein kinase 1 activated Transcription factor 2 P Specific protein kinase 2 activated

Phytochrome activated by light Cell wall

Transcription

Light Translation 1 The light signal is detected by the phytochrome receptor, which then activates at least two signal transduction pathways. Ca2+

Ca2+

channel opened

De-etiolation (greening) response proteins

Signal Transduction Activated by Light


1 Reception
CYTOPLASM Plasma membrane

2 Transduction

3 Response
Transcription factor 1 NUCLEUS

cGMP
Second messenger produced

Specific protein kinase 1 activated Transcription factor 2 P Specific protein kinase 2 activated

Phytochrome activated by light Cell wall

2 One pathway uses cGMP as a second messenger that activates a specific protein kinase.The other pathway involves an increase in cytoplasmic Ca2+ that activates another specific protein kinase.

Transcription

Light Translation 1 The light signal is detected by the phytochrome receptor, which then activates at least two signal transduction pathways. Ca2+

Ca2+

channel opened

De-etiolation (greening) response proteins

Signal Transduction Activated by Light


1 Reception
CYTOPLASM Plasma membrane

2 Transduction

3 Response
Transcription factor 1 NUCLEUS

cGMP
Second messenger produced

Specific protein kinase 1 activated Transcription factor 2 P Specific protein kinase 2 activated

Phytochrome activated by light Cell wall

2 One pathway uses cGMP as a second messenger that activates a specific protein kinase.The other pathway involves an increase in cytoplasmic Ca2+ that activates another specific protein kinase.

Transcription

Light Translation 1 The light signal is detected by the phytochrome receptor, which then activates at least two signal transduction pathways. Ca2+ 3 Both pathways lead to expression of genes for proteins that function in a particular response.

Ca2+

channel opened

De-etiolation (greening) response proteins