Chapter 39.

Plant signals and Behavior

Stimuli and a Stationary Life

▪ Plants receive signals from the environment

and respond by altering growth and
development

▪ For example, the bending of a

dodder seedling toward a host plant
occurs in response to chemicals
released by the host

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Concept 39.1: Signal transduction pathways link signal reception to
response

▪ A potato left growing in darkness produces pale stems, unexpanded leaves, and
short roots

▪ These are morphological adaptations for growing in darkness, collectively called

etiolation

▪ After exposure to light, a potato undergoes changes called de-etiolation, in which

shoots and roots grow normally

▪ A potato’s response to light is an example of cell signal processing

▪ The stages are reception, transduction, and response

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1. Reception

▪ Signals are detected by receptors, proteins that change in shape in response to

specific stimuli

▪ In de-etiolation, the receptor is a phytochrome capable of detecting light

2. Transduction

▪ Second messengers transfer and amplify signals from receptors to proteins that
cause responses

▪ Two types of second messengers play an important role in de-etiolation: calcium

ions (Ca2+) and cyclic GMP (cGMP)

▪ The phytochrome receptor responds to light by

- Opening Ca2+ channels, which increases Ca2+ levels in the cytosol

- Activating an enzyme that produces cGMP

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3. Response

▪ A signal transduction pathway leads to regulation of one or more cellular activities

▪ In most cases, these responses to stimulation involve increased activity of enzymes

▪ This can occur by transcriptional regulation or post-translational modification

1) Post-translational Modification of Preexisting Proteins

▪ Post-translational modification involves modification of existing proteins in the

signal response

▪ Modification often involves the phosphorylation of specific amino acids

▪ The second messengers cGMP and Ca2+ activate protein kinases directly

▪ Protein kinases often work in a cascade linking initial stimuli to gene expression
through phosphorylation of transcription factors

▪ Protein phosphatases “switch off” the signal transduction pathways by

dephosphorylating proteins

2) Transcriptional Regulation

▪ Specific transcription factors bind directly to specific regions of DNA and control
transcription of specific genes

▪ Some transcription factors are activators that increase the transcription of specific
genes

▪ Other transcription factors are repressors that decrease the transcription of specific
genes

3) De-etiolation (“Greening”) Proteins

▪ De-etiolation activates enzymes that

▪ Function in photosynthesis directly

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 ▪ Supply the chemical precursors for chlorophyll production

▪ Affect the levels of plant hormones that regulate growth

Concept 39.2: Plants use chemicals to communicate

▪ Plant hormones are chemical signals that modify or control one or more specific
physiological processes within a plant

▪ Plant hormones are also called plant growth regulators

▪ Plant hormones are produced in very low concentrations, but can have profound
effects on growth and development

▪ Each hormone has multiple effects, but multiple hormones can influence a single
process

▪ Plant responses depend on amount and concentration of specific hormones and

often on the combination of hormones present

1. A Survey of Plant Hormones

▪ The major plant hormones include

Auxin, Cytokinins, Gibberellins, Abscisic acid, Ethylene,

Brassinosteroids, Jasmonates, Strigolactones

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1) Auxin

▪ Any response resulting in curvature of organs toward or away from a stimulus is

called a tropism

▪ In the late 1800s, Charles Darwin and his son Francis conducted experiments on
phototropism, a plant’s response to light

▪ They observed that a grass seedling could bend toward light only if the tip of the
coleoptile was present and exposed to light

▪ They postulated that a signal was transmitted from the tip to the elongating region

▪ In 1913, Peter Boysen-Jensen demonstrated that the signal was a mobile chemical
substance

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▪ The term auxin refers to any chemical that promotes elongation of coleoptiles

▪ Indoleacetic acid (IAA) is a common auxin in plants; in this lecture the term auxin
refers specifically to IAA

▪ Transport of auxin is polar; it is produced in shoot tips and is transported down the
stem

▪ Auxin transporter proteins move the hormone from the basal end of one cell into
the apical end of the neighboring cell

▪ The direction of auxin does not change in response to gravity

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① The Role of Auxin in Cell Elongation

▪ According to the acid growth hypothesis, auxin stimulates proton pumps in the
plasma membrane

▪ Proton pumps move H+ into the cell, lowering the pH in the cell wall and increasing
the membrane potential

▪ Reduced pH activates expansins, enzymes that loosen the fabric of the cell wall

▪ Osmotic uptake of water into the cell increases turgor pressure

▪ Increased cell wall plasticity combined with increased turgor pressure enable the
cell to elongate

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▪ Auxin also alters gene expression and stimulates a sustained growth response

② Auxin’s Role in Plant Development

▪ Polar transport of auxin plays a role in pattern formation of the developing plant

▪ Reduced auxin flow from the shoot of a branch stimulates growth in lower branches

▪ Auxin transport plays a role in phyllotaxy, the arrangement of leaves on the stem

▪ Polar transport of auxin from leaf margins directs leaf venation pattern

▪ The activity of the vascular cambium is under control of auxin transport

▪ Organization of female angiosperm gametophytes is likely regulated by an auxin

gradient

③ Practical Uses for Auxins

▪ The auxin indolebutyric acid (IBA) stimulates adventitious roots and is used in
vegetative propagation of plants by cuttings

▪ Synthetic auxins used in herbicides such as 2,4-D kill eudicots by causing a

hormonal overdose; monocots are able to inactivate these hormones

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▪ Developing seeds produce auxin, which promotes fruit development

▪ Greenhouse tomatoes produce few seeds

▪ Spraying synthetic auxins on greenhouse tomatoes improves fruit development

2) Cytokinins

▪ Cytokinins are so named because they stimulate cytokinesis (cell division)

▪ The most common natural cytokinin is zeatin because it was first discovered in
maize (Zea mays)

▪ Cytokinins influence cell division, cell differentiation, and apical dominance

① Control of Cell Division and Differentiation

▪ Cytokinins are produced in actively growing tissues such as roots, embryos, and
fruits

▪ Cytokinins work together with auxin to control cell division and differentiation

-If auxin and cytokinins are in equal concentration, a mass of

undifferentiated cells grows

-If cytokinin levels increase, shoot buds develop; if auxin levels increase,
roots form

② Control of Apical Dominance

▪ Apical dominance is a terminal bud’s ability to suppress development of axillary

buds

▪ It is under the control of sugar, cytokinins, auxin, and strigolactone

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▪ Removal of the apical bud increases sugar availability and decreases auxin and
strigolactone levels, initiating axillary bud growth

③ Anti-aging Effects

▪ Cytokinins slow the aging of some plant organs by inhibiting protein breakdown,
stimulating RNA and protein synthesis, and mobilizing nutrients from surrounding
tissues

3) Gibberellins

▪ Gibberellins have a variety of effects, such as stem elongation, fruit growth, and
seed germination

① Stem Elongation

▪ Gibberellins are produced in young roots and leaves

▪ Gibberellins stimulate growth of leaves and stems by enhancing cell elongation and
cell division

▪ Bolting, rapid growth of the floral stalk, is induced by gibberellins

② Fruit Growth

▪ In many plants, both auxin and gibberellins must be present for fruit to develop
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▪ Gibberellins are used in spraying of Thompson seedless grapes

③ Germination

▪ After water is imbibed, release of gibberellins from the embryo signals seeds to
germinate

4) Abscisic Acid

▪ Abscisic acid (ABA) slows growth, often by antagonizing the actions of growth
hormones

▪ ABA has many other effects on plants including seed dormancy and drought
tolerance

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① Seed Dormancy

▪ Seed dormancy increases the likelihood that

the seed will germinate only in optimal
conditions

▪ Many dormant seeds germinate when ABA is

removed or inactivated

▪ The ratio of ABA to gibberellins often affects

whether seeds will break dormancy

▪ Precocious (early) germination can be caused

by inactive or low levels of ABA

② Drought Tolerance

▪ ABA is the primary internal signal that enables plants to withstand drought

▪ ABA accumulation in wilting leaves causes stomata to close rapidly

▪ Transport of ABA from water-stressed root systems to leaves can act as an “early
warning system”

5) Ethylene

▪ Plants produce ethylene in response to stresses such as drought, flooding,

mechanical pressure, injury, and infection

▪ The effects of ethylene include response to mechanical stress, senescence, leaf

abscission, and fruit ripening

① The Triple Response to Mechanical Stress

▪ Ethylene is produced when a seedling tip pushes against an obstacle

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▪ The production of ethylene induces a triple response in which stem elongation is
slowed, the stem thickens, and the stem begins to grow horizontally

▪ Vertical growth resumes when the effects of the ethylene wear off

▪ Some Arabidopsis mutants have abnormal triple responses

▪ Ethylene-insensitive (ein) mutants fail to undergo the triple response after

exposure to ethylene

▪ Ethylene-overproducing (eto) mutants undergo the triple response even in

the absence of obstacles

▪ Constitutive triple-response (ctr) mutants undergo a triple response even if

ethylene is not present

② Senescence

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▪ Senescence is the programmed death of certain cells or organs or entire plants

▪ A burst of ethylene is associated with the onset of apoptosis, programmed cell

death

③ Leaf Abscission

▪ A change in the balance of auxin

and ethylene controls leaf
abscission, the process that occurs
in autumn when a leaf falls

④ Fruit Ripening

▪ In many cases, a burst of ethylene production in a fruit triggers the ripening process

▪ Ethylene triggers ripening, and ripening triggers release of more ethylene

▪ Fruit producers can control ripening by picking green fruit and controlling ethylene
levels

6) More Recently Discovered Plant Hormones

▪ Brassinosteroids are chemically similar to cholesterol and the sex hormones of

animals

▪ They induce cell elongation and division in stem segments and seedlings at low
concentration

▪ They slow leaf abscission and promote xylem differentiation

▪ Jasmonates, including jasmonate (JA) and methyl jasmonate (MeJA) play important
roles in plant defense and development

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▪ They are produced in response to wounding and are involved in controlling plant
defenses

▪ Jasmonates also regulate many other physiological processes, including

① Nectar secretion ② Fruit ripening ③ Pollen production ④ Flowering time

⑤ Seed germination ⑥ Root growth ⑦ Tuber formation ⑧ Mycorrhizal symbiosis

⑨ Tendril coiling

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Concept 39.3: Responses to light are critical for plant success

▪ Light cues many key events in plant growth and development

▪ Effects of light on plant morphology are called photomorphogenesis

▪ Plants detect not only 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

▪ Action spectra are useful in studying any process that depends on light

▪ Action spectra can be used to determine which responses are mediated by a specific
photoreceptor

▪ There are two major classes of light receptors:

blue-light photoreceptors and phytochromes

1. Blue-Light Photoreceptors

▪ Blue light initiates a variety of plant responses such as hypocotyl elongation,

stomatal opening, and phototropism

▪ Phototropin is a protein kinase involved in mediating plant responses to blue light

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2. Phytochrome Photoreceptors

▪ Phytochromes are pigments that absorb mostly red and far-red light

▪ They regulate many of a plant’s responses to light including de-etiolation, seed

germination, and shade avoidance

1) Phytochromes and Seed Germination

▪ Many seeds remain dormant until light and other conditions are near optimal

▪ In the 1930s, scientists at the U.S. Department of Agriculture determined the action
spectrum for light-induced germination of lettuce seeds

▪ Red light increased germination, while far-red light inhibited germination

▪ The effects of red and far-red light are reversible; the final light exposure determines
the response

▪ The photoreceptors responsible for the opposing effects of red and far-red light
are phytochromes

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▪ Phytochromes exist in two photoreversible states, with conversion of P r to Pfr
triggering many developmental responses

▪ Red light triggers the conversion of Pr to Pfr

▪ Far-red light triggers the conversion of Pfr to Pr

▪ The conversion of Pr to Pfr is faster than the reverse process

▪ Sunlight, containing both red and far-red light, increases the ratio of Pfr to Pr and
triggers germination

2) Phytochromes and Shade Avoidance

▪ The phytochrome system also provides the plant with information about the quality
of light

▪ Leaves in the canopy absorb red light and allow far-red light to pass through to
the shaded plants below

▪ When a tree is shaded, the phytochrome ratio shifts in favor of Pr, inducing vertical
growth

▪ This is called the “shade avoidance” response

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3. Biological Clocks and Circadian Rhythms

▪ Many plant processes oscillate during the day in response to light and temperature
changes

▪ Many other processes oscillate with a frequency of 24 hours, even under constant
environmental conditions

▪ For example, many legumes lower their leaves in the evening and raise them
in the morning, even when kept in constant light or darkness

▪ Circadian rhythms are cycles that are about 24 hours long and are governed by
an internal “clock”

▪ These cycles can be free-running, varying from 21 to 27 hours, when organisms are
kept in a constant environment

▪ The 24-hour period arises from the transcription of “clock genes” regulated through
negative-feedback loops

4. The Effect of Light on the Biological Clock

▪ Light is the factor that entrains the biological clock to precisely 24 hours every day

▪ Both phytochromes and blue-light photoreceptors can entrain circadian rhythms in

plants

▪ Phytochrome conversion marks sunrise and sunset, providing the biological clock
with environmental cues
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5. Photoperiodism and Responses to Seasons

▪ Many critical events in plant life cycles, such as seed germination and flowering,
occur seasonally

▪ Plants detect the time of year based on changes in photoperiod, the relative lengths
of night and day

▪ Photoperiodism is a physiological response to photoperiod

1) Photoperiodism and Control of Flowering

▪ Some processes, including flowering in many species, require a certain photoperiod

▪ Plants that flower when a light period is shorter than a critical length are called
short-day plants

▪ Plants that flower when a light period is longer than a certain number of hours are
called long-day plants

▪ Flowering in day-neutral plants is controlled by plant maturity, not photoperiod

① Critical Night Length

▪ In the 1940s, researchers discovered that flowering and other responses to

photoperiod are actually controlled by night length, not day length

▪ Short-day plants are governed by whether the critical night length sets a minimum
number of hours of darkness

▪ Long-day plants are governed by whether the critical night length sets a maximum
number of hours of darkness

▪ Red light is the most effective color in interrupting the night length

▪ Phytochrome is the pigment that detects the red light

▪ A flash of red light followed by a flash of far-red light does not disrupt night length
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▪ Some plants flower after only a single exposure to the required photoperiod

▪ Other plants need several successive days of the required photoperiod

▪ Still others need an environmental stimulus in addition to the required photoperiod

▪ For example, vernalization is a pretreatment with cold to induce flowering

2) A Flowering Hormone?

▪ Photoperiod is detected by leaves, which cue buds to develop as flowers

▪ The flowering signal molecule is called florigen

▪ Florigen may be a protein governed by the FLOWERING LOCUS T (FT) gene

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Concept 39.4: Plants respond to a wide variety of stimuli other than
light

▪ Because of immobility, plants must adjust to a range of environmental

circumstances through developmental and physiological mechanisms

1. Gravity

▪ Response to gravity is known as gravitropism

▪ Roots show positive gravitropism; shoots show negative gravitropism

▪ Plants may detect gravity by the settling of statoliths, dense cytoplasmic

components

▪ Some Arabidopsis mutants that lack statoliths are still capable of gravitropism

▪ Dense organelles, in addition to starch granules, may contribute to gravity detection

2. Mechanical Stimuli

▪ The term thigmomorphogenesis refers to changes in form that result from

mechanical disturbance

▪ For example, rubbing stems of young plants a couple of times daily results
in plants that are shorter than controls

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 ▪ Thigmotropism is growth in response to touch

▪ It occurs in vines and other climbing plants

▪ Some plants undergo rapid leaf movements in response to mechanical stimulation

▪ For example, Mimosa pudica folds its leaflets and collapses in response to
touch

▪ The touch response results from the transmission of electrical impulses called action
potentials

3. Environmental Stresses

▪ Environmental stresses have a potentially adverse effect on survival, growth, and

reproduction

▪ Stresses can be biotic (living) or abiotic (nonliving)

-Biotic stresses include herbivores and pathogens

-Abiotic stresses include drought, flooding, salt stress, heat stress, and cold stress

1) Drought

▪ Plants may wilt or die when water loss by transpiration exceeds water absorption

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▪ During drought, plants reduce transpiration by closing stomata, reducing exposed
surface area, and in some species, shedding leaves

2) Flooding

▪ Waterlogged soils lack the air spaces needed to provide oxygen for cellular
respiration in roots

▪ Enzymatic destruction of root cortex cells creates air tubes that help plants survive
oxygen deprivation during flooding

▪ Some plants, such as mangroves, also produce aerial roots

3) Salt Stress

▪ Sodium and some other ions are toxic to plants in high concentrations

▪ Salt can also lower the water potential of the soil solution and reduce water uptake

▪ Plants respond to salt stress by producing solutes tolerated at high concentrations

▪ This process keeps the water potential of cells more negative than that of the soil
solution

4) Heat Stress

▪ Excessive heat can denature a plant’s enzymes

▪ Transpiration helps cool leaves by evaporative cooling

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▪ Heat-shock proteins are produced at temperatures above 40ºC to help protect
other proteins from heat stress

5) Cold Stress

▪ Cold temperatures decrease membrane fluidity

▪ Altering lipid composition of membranes is a response to cold stress

▪ Ice formation during freezing reduces water potential outside the cell

▪ Cells of many frost-tolerant species increase solute concentration of the cytoplasm

to reduce water loss from the cell

▪ Plants, and many other organisms, have antifreeze proteins that hinder the
formation of ice crystals

▪ Frost tolerance can be increased in some crop plants by engineering antifreeze

genes into their genomes

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Concept 39.5: Plants respond to attacks by pathogens and
herbivores

▪ Plants have formed mutually beneficial interspecific interactions with many species
including mycorrhizal fungi and animal pollinators

▪ Plants are also subject to attack by herbivorous animals and pathogenic viruses,
bacteria, and fungi

▪ Defense systems have evolved to deter herbivory, prevent infection, and combat
pathogens

1. Defenses Against Pathogens

▪ A plant’s first line of defense against infection is the barrier presented by the
epidermis and periderm

▪ Pathogens can enter through wounds or natural openings, such as stomata

▪ Two types of immune response defend the plant after pathogens enter: PAMP-
triggered immunity and effector-triggered immunity

1) PAMP-Triggered Immunity

▪ The first line of immune defense depends on the plant’s ability to recognize
pathogen-associated molecular patterns (PAMPs)

▪ These molecular sequences are specific to certain pathogens

▪ PAMP recognition starts a chain of signaling events leading to the production of

antimicrobial chemicals and toughening of the cell wall

2) Effector-Triggered Immunity

▪ Plants and pathogens have engaged in an evolutionary arms race

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▪ Pathogens that have evolved the ability to deliver effectors into plant cells can
suppress PAMP-triggered plant immunity

▪ Effectors are pathogen-encoded proteins that cripple the host’s innate immune
system

▪ A second level of plant immune defense evolved in response to these pathogens

▪ Effector-triggered immunity results from the action of hundreds of disease

resistance (R) genes

▪ Each R protein is activated by a specific effector

▪ R proteins activate plant defenses by triggering signal transduction pathways

▪ These defenses include the hypersensitive response and systemic acquired

resistance

① The Hypersensitive Response

▪ Local cell and tissue death at and near the infection site is the hypersensitive
response

▪ This response induces production of enzymes and chemicals that attack the
pathogen

▪ It also stimulates changes in the cell wall that confine the pathogen

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 3) Systemic Acquired Resistance

▪ Systemic acquired resistance arises from the plant-wide expression of defense

genes. It provides protection from a diversity of pathogens for several days

▪ Methylsalicylic acid is synthesized around the infection site and carried in the
phloem to other remote sites where it is converted to salicylic acid

▪ Salicylic acid triggers the defense system to respond rapidly to another infection

▪ Plant disease epidemics can alter plant community structure when they occur in
natural populations

▪ In crop plants, they have the potential to cause starvation and suffering in human
populations

▪ Plant biologists are stockpiling the seeds of wild relatives of crop plants to preserve
the genetic diversity required to curb future plant epidemics

2. Defenses Against Herbivores

▪ Herbivory, animals eating plants, can restrict plant growth because energy is
diverted into defense

▪ Plants counter excessive herbivory with defenses that can be observed at multiple
levels of biological organization

① Molecular-Level Defenses

▪ Chemical compounds including terpenoids, phenolics, and alkaloids can be

produced to deter attackers

② Cellular-Level Defenses

▪ Cells may be specialized to form trichomes, store chemical deterrents, or produce

irritants

③ Tissue-Level Defenses

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▪ Leaves may be toughened with sclerenchyma tissue

④ Organ-Level Defenses

▪ Leaves can be modified into spines and bristles

▪ Some species have leaves that appear partially eaten; others have structures that
mimic insect eggs

⑤ Organismal-Level Defenses

▪ Plants may alter their physiology in response to attack by herbivores

▪ For example, tobacco plants alter their flowering time from night to morning
in response to feeding by hawk-moth larvae

⑥ Population-Level Defenses

▪ Some plants release chemicals in response to herbivore attack that trigger defense
responses in nearby members of the population

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 ▪ Other plant populations use masting, synchronous mass seed production following
long intervals, to ensure some seeds survive herbivory

⑦ Community-Level Defenses

▪ Some plants “recruit” predatory animals that help defend against specific herbivores

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