Chapter 39: Plant Responses to Internal and

External Signals

19 April 2008

Contents
1 Signal Transduction and Plant Responses 2
1.1 Signal transduction pathways link internal and environmental sig-
nals to chemical responses . . . . . . . . . . . . . . . . . . . . . . 2
1.1.1 Reception . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.1.2 Transduction . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.1.3 Response . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.1.4 The Greening Proteins. . . . . . . . . . . . . . . . . . . . 3

2 Plant Responses to Hormones 3

2.1 Research on how plants grow toward light led to the discovery of
plant hormones . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2.2 Plant hormones help coordinate growth, development, and re-
sponses to environmental stimuli . . . . . . . . . . . . . . . . . . 3
2.2.1 Auxin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2.2.2 Cytokinins . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2.2.3 Gibberellins . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2.2.4 Abscisic Acid . . . . . . . . . . . . . . . . . . . . . . . . . 5
2.2.5 Ethylene . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
2.2.6 Brassinosteroids . . . . . . . . . . . . . . . . . . . . . . . 7

3 Plant Responses to Light 7

3.1 Blue-light photoreceptors are a heterogenous group of pigments . 7
3.2 Phytochromes function as photoreceptors in many plant responses
to light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
3.2.1 The Phytochrome Switch and Seed Germination . . . . . 7
3.2.2 The Phytochroem Switch and Shade Avoidance . . . . . . 8
3.3 Biological clocks control circadian rhythmns in plants and other
eukaryotes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
3.4 Light entrains the biological clock . . . . . . . . . . . . . . . . . . 8
3.5 Photoperiodism synchronizes many plant responses to changes of
season . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

1
 3.5.1 Photoperiodism and the Control of Flowering . . . . . . . 8
3.5.2 Is There a Flowering Hormone? . . . . . . . . . . . . . . . 9
3.5.3 Meristem Transition from Vegetative Growth to Flowering 9

4 Plant Responses to Environmental Stimuli Other Than Light 9

4.1 Plants respond to environmental stimuli through a combination
of developmental and physiological mechanisms . . . . . . . . . . 9
4.1.1 Responses to Gravity . . . . . . . . . . . . . . . . . . . . 9
4.1.2 Responses to Mechanical Stimuli . . . . . . . . . . . . . . 9
4.1.3 Responses to Stress . . . . . . . . . . . . . . . . . . . . . 10

5 Plant Defense: Responses to Herbivores and Pathogens 11

5.1 Plants deter herbivores with both physical and chemical defenses 11
5.2 Plants use multiple lines of defense against pathogens . . . . . . 11
5.2.1 Gene-for-Gene Recognition . . . . . . . . . . . . . . . . . 11
5.2.2 Hypersensitive Response . . . . . . . . . . . . . . . . . . . 11
5.2.3 Systemic Acquired Resistance . . . . . . . . . . . . . . . . 11

1 Signal Transduction and Plant Responses

1.1 Signal transduction pathways link internal and envi-
ronmental signals to chemical responses

1. Plant germinate in dark elongate stems.

2. Greening elongation rate slow, leaves expand, roots elongate, chlorophyll.

1.1.1 Reception
1. Phytochrome. Light-absorbing pigment attached to protein. Cytoplasmic.

2. Discovered from aurea (L. gold-colored ) mutants.

1.1.2 Transduction
1. Phytochrome very sensitive.

2. Second messengers activate cascade. Activate transcription factor:

(a) cGMP (cyclic guanoside monophosphate).

2+
(b) Ca -calmodulin complex.

1.1.3 Response
Transcriptional Regulation. Transcription factors bind to DNA. Repress-
able or inducible.

2
Post-Translational Modication of Proteins.
1. Protein kinases catalyze phosphorylation.

2. Protein phosphatases dephosphorylateturn o  signals.

1.1.4 The Greening Proteins.

1. Enzymes that participate in photosynthesis.

2. Enzymes to supply chemical stock for chlorophyll.

3. Hormones.

2 Plant Responses to Hormones

1. Produced one place; receive elsewhere.

2. Minute quantities sucient.

2.1 Research on how plants grow toward light led to the

discovery of plant hormones

1. Tropism is growth towards or away from stimulus.

(a) Phototropism.
2. Coleoptilecasing of grass shoot. Shaded cells grow faster, bending plants
towards light. Experiments:

(a) Darwin & Darwin: tip of coleoptile sense light.

(b) Boysen-Jensen: growth signal chemical; travels down.

(c) Went: curve because ↑ [auxin] on darker side of coleoptile.

3. Others (e.g. dicots) dierent: ↑ [growth inhibitors] on light side.

2.2 Plant hormones help coordinate growth, development,

and responses to environmental stimuli

1. Small; pass across cell walls.

2. Aect division, elongation, dierentiation.

3. Relative concentration, not absolute amount more signicant.

3
2.2.1 Auxin
1. IAA (indoleacetic acid).

2. Polar transport:

(a) Shoot tip to base.

(b) Parenchyma to parenchyma.

(c) Gravity independent.

(d) Coupled to proton pump.

The Role of Auxin in Cell Elongation.

1. Synthesized at apical meristem.

2. Stimulate growth at ↓ [ ].

(a) ↑[ ] inhibit elongation by synthesizing ethylene.

3. Acid growth hypothesis: auxin stimulates proton pump. Acidies wall.

(a) Activate expansins that break cross links between cellulose.

(b) Increase membrane potential →↑ ions →↑ water.

4. Also stimulates protein synthesis for sustained growth.

Lateral and Adventitious Root Formation.

Auxins as Herbicides.
1. 2,4-dinitrophenol (2,4-D) kills dicot weedsoverdose.

2. Monocots (maize, turfgrass) ne.

Other Eects of Auxin.

1. Induce division in vascular cambium.

2. Fruit growth (secreted by developing seeds).

2.2.2 Cytokinins
1. Stimulate cytokinesis. Modied adenine.

2. Zeatin most common.

2+
3. Surface receptor. Some plants open Ca channels.

4. Cytoplasmic: stimulate transcription in isolated nuclei.

4
Control of Cell Division and Dierentiation.
1. Produced in growing tissue: root, embryo, fruit.

2. Concert with auxin. Ratio determine rate of dierentiation.

(a) Cytokinins + auxin: Divide. Balanced, undierentiated callus.

(b) ↑ cytokinin → shoot buds.

Control of Apical Dominance.

1. Direct inhibition hypothesis: auxin, cytokinin antagonistically aect api-
cal bud growth.

2. But, cutting o terminal bud increases auxin in axillary buds.

Anti-Aging Eects.
1. Inhibit protein breakdown, stimulate synthesis, mobilize nutrients.

2. Leaves in cytokinin solution last much longer.

3. Florists spray on cut owers.

2.2.3 Gibberellins
Produced by Gibberella fungus; also by plants, but lower concentrations.

Stem Elongation.
1. Produced at roots, young leaves.

2. Stems: cell elongation and division.

3. Cell wall loosening: faciliate penetration of expansins.

4. Treating normal plants often does nothing: already at optimal levels.

Fruit Growth.
1. Auxin + gibberellins.

2. Thompson seedless grapes: bigger, more space.

Germination. Released after water is imbibed. Synthesis of α-amylase.

2.2.4 Abscisic Acid

1. ABA originally thought to be important for leaf abscission.

2. Slows growth. Antagonize other hormones.

5
Seed Dormancy.
1. ↑ in developing seeds → inhibit germination.

2. ABA must be removed, inactivated somehow. Rain, cold, light.

Drought Stress.
1. ABA accumulates when wilting; close stomata.

2. May stress root rst. ABA transported for early warning.

2.2.5 Ethylene
1. Produce in response to stress: drought, ooding, injury, infection.

2. Produced in fruit ripening.

3. Released in bursts.

4. Articial auxin induces this.

The Triple Response to Mechanical Stress: Using Mutants to Dissect

a Signal-Transduction Pathway.
1. Slow elongation.

2. Thicken stem.

3. Grow sideways.

Apoptosis: Programmed Cell Death.

1. Shedding leaves, death of annual, xylem maturation.

2. New enzymes break stu down. Salvage.

Leaf Abscission.
1. Prevent dessication when roots cannot absorb water.

2. Essential elements salvaged in stem parenchyma; restored later.

3. Abscission layer weakend by hydrolysis of cell walls of petiole.

4. Cork forms protective scar to prevent infection.

5. Gravity + wind breaks it o.

6. Aging leaf less auxin →↑ sensitive to ethylene.

6
Fruit Ripening.
1. Break down cell wall, convert acid → sugar.

2. Scents and colors.

3. Positive feedback: ethylene → ripening → ethylene.

(a) May aect adjacent fruit.

(b) Can manipulate: add ethylene or CO2 to inhibit.

2.2.6 Brassinosteroids
1. Similar to auxin (eects), cholesterol/sex hormones (structure).

(a) Elongation, division.

(b) Retard abscission, promote xylem dierentiation.

3 Plant Responses to Light

1. Photomorphogenesis is eect of light on plant morphology.
2. Action spectrum is a graph relating physiological response to wave-
length.

3. Red, blue light most important in photomorphogenesis.

3.1 Blue-light photoreceptors are a heterogenous group of

pigments

Hypocotyl elongation ( cryptochromes), stomata opening (zeaxanthin), coleop-

tile curvature ( phototropin).

3.2 Phytochromes function as photoreceptors in many plant

responses to light

3.2.1 The Phytochrome Switch and Seed Germination

1. Red light stimulate.

2. Far-red (near edge of visibility, 730nm) inhibits.

3. Last ash of light determines. Reversible.

red light
(a) Pf −−−−−→ Pf r .
(b) Presence of Pf r activates pathway.

7
3.2.2 The Phytochroem Switch and Shade Avoidance
1. Also conveys quality of light.

2. Day, equilibrium.

3. But shade → far red > red → Pr accumulates. Inhibit.

4. Also tracks day, season passage.

3.3 Biological clocks control circadian rhythmns in plants

and other eukaryotes

1. Processes oscillate daily.

2. Circadian rhythmns. Endogenous. Calibrated by environmental cues.

3. Perhaps negative feedback.

(a) 1st half: ↑ transcription factor.

(b) 2nd half: ↑ due to own inhibition.

3.4 Light entrains the biological clock

Sudden burst of Pf r trigger.

3.5 Photoperiodism synchronizes many plant responses to

changes of season

3.5.1 Photoperiodism and the Control of Flowering

1. Short-day plants (long-night) need period shorter than a critical length
to ower.

(a) Chrysanthemum, poinsettia, soybean.

2. Long-day plants (short-night) need period longer than critical.

(a) Spring, early summer.

(b) Spinach, Radish, lettuce.

3. Day-neutral plants ower when they reach maturity.

(a) Tomato, rice, dandelion.

8
Critical Night Length.
1. Continuous darkness length, not daytime length matters.

2. Interrupt night → eect.

3. Very accuratesome within minute of critical length.

(a) Some on same date each year.

(b) Some after single exposure; others need multiple.

(c) Vernalizationcold exposure required too.

3.5.2 Is There a Flowering Hormone?
1. Exposure of one leaf sucient to cause all owers to bloom.

2. Don't what what hormone, or ratio of two.

3.5.3 Meristem Transition from Vegetative Growth to Flowering

Meristem-identity genes for bud switched on.

4 Plant Responses to Environmental Stimuli Other

Than Light
4.1 Plants respond to environmental stimuli through a
combination of developmental and physiological mech-
anisms

4.1.1 Responses to Gravity

1. Gravitropism: positive → towards ground; negative → away.

(a) Ensures shoots grow upwards.

(b) Auxin.

2. Statoliths signal gravity. Dense plastids with starch. Bottom of cell.

(a) Not strictly necessary; perhaps enhance a system.

4.1.2 Responses to Mechanical Stimuli

1. Thigmomorphogenesis is change resulting from mechanical disturbance.

2. Plants very sensitive: measure leaf with ruler → grow dierently.

3. Thigmotropism is directional growth in response to touch.

9
 (a) Mimosa sensitive plant. Pulvini (motor neurons) lose K → water
leaves cells. Flaccid → leaves drop.

(b) Transmission: other leaets follow suit. Action potential resembles

neurons.

4.1.3 Responses to Stress

Drought.
1. ABA → keep stomata closed.

2. Inhibit growth of young leaves.

3. Wilting rolles up shape to minimize wind.

4. Deep roots grow; shallow don't.

Flooding.
1. Mangroves: specialized to breath.

2. Ethylene → root cortex apoptosis → creates air tubes snorkels.

Salt Stress.
1. Lowers ψ→ water decit despite water.

2. Ions toxic at high [ ].

+
3. Secrete organic ions that substitute for Na to maintain membrane po-
tential.

4. Halophytes have salt glands; pump salt across leaf epidermii.

Heat Stress.
1. Evaporative cooling (transpiration).

2. Heat-shock proteins hold proteins in place. Prevent denaturation.

Cold Stress.
1. Problem: freezing lipid membrane.

2. ↑ unsaturated fatty acids → stay uid.

3. Takes time.

4. Freezing: ice in cell walls, intercellular space. Water osmoses out. Toxic
concentrations of ions in cytoplasm. Cytoplasm usually does not freeze
because of solutes.

5. Frost-tolerant: increase sugar, other solutes that are better tolerated at

high concentration.

10
5 Plant Defense: Responses to Herbivores and
Pathogens
5.1 Plants deter herbivores with both physical and chem-
ical defenses

1. Canavanine resembles arginine, but dierent enough to adversely aect

proteins. Eater dies.

2. Some plants recruit predators to eat their predators.

(a) Parasitoid wasp recruited when caterpillars start eating leaf. Wasp
lays eggs within caterpillars.

(b) Early warning system → neighbors activate defense genes.

5.2 Plants use multiple lines of defense against pathogens

First line is skin. Not very impermeable.

5.2.1 Gene-for-Gene Recognition

1. Avirulent is the evolution of compromise. Pathogen gains access to
host to reproduce, but does not cause severe damage.

2. Gene-for-gene recognition is precise matchup of plant and pathogen

allele.

(a) R gene recognizes complementary Avr genes.

(b) R codes for receptor.

3. Signal transduction pathway → spread warning.

5.2.2 Hypersensitive Response

1. Elicitors released by cell-wall damage induce phytoalexins.

(a) PR proteins. Antimicrobial, signal.

(b) Cross-linking, lignin deposition.

2. Cells at infection site mount defense, they apoptose.

5.2.3 Systemic Acquired Resistance

1. Salicylic acid released by local cells before they die.

2. Antimicrobial molecules produced to protect cell for several days.

11