Chapter PP.

5 
PLANT RESPONSES TO INTERNAL AND EXTERNAL SIGNALS

1. SIGNAL TRANSDUCTION AND PLANT RESPONSES   

Signal-transduction pathways link internal and environmental signals to cellular responses

2. PLANT RESPONSES TO HORMONES   

Research on how plants grow toward light led to the discovery of plant hormones   
Plant hormones help coordinate growth, development, and responses to environmental stimuli

3. PLANT RESPONSES TO LIGHT   

Blue-light photoreceptors are a heterogeneous group of pigments   
Phytochromes function as photoreceptors in many plant responses to light   
Biological clocks control circadian rhythms in plants and other eukaryotes   
Light entrains the biological clock   
Photoperiodism synchronizes many plant responses to changes of season

4. PLANT RESPONSES TO ENVIRONMENTAL STIMULI OTHER THAN LIGHT   

Plants respond to environmental stimuli through a combination of developmental
and physiological mechanisms

5. PLANT DEFENSE: RESPONSES TO HERBIVORES AND PATHOGENS   

Plants deter herbivores with both physical and chemical defenses   
Plants use multiple lines of defense against pathogens
 1. SIGNAL TRANSDUCTION AND PLANT RESPONSES

* Signal-transduction pathways link internal and environmental signals to cellular responses

All organisms, including plants, have the ability to receive specific environmental and internal signals
and respond to them in ways that enhance survival and reproductive success. Plants, too, have cellular
receptors that they use to detect important changes in their environment, whether the change is an
increase in the concentration of a growth hormone, an injury from a caterpillar munching on leaves, or a
decrease in day length as winter approaches. In order for an internal or external stimulus to elicit a
physiological response, certain cells within the organism must possess an appropriate receptor, a
molecule that is sensitive to and affected by the specific stimulus. Upon receiving a stimulus, a receptor
initiates a specific series of biochemical steps, a signal transduction pathway, which couples reception
of the stimulus to the response of the organism. As you will learn, plants are sensitive to a wide range
of internal and external stimuli, and each of these stimuli initiates a specific signal transduction
pathway. We introduced general concepts of signal transduction in cells in Chapter 11. Here, we apply
those concepts to specific examples in plants
1.1. Signal-transduction pathways
link internal and environmental signals to cellular responses

In the back corner of a food cabinet, a long forgotten potato (a modified underground stem,
or tuber) has sprouted shoots from its "eyes" (axillary buds).

These shoots, however, scarcely resemble those of a typical plant. Instead of broad green
leaves and sturdy stems, these dark-sprouted shoots are ghostly pale, have stems that
are long and thin, bear leaves that are unexpanded, and give rise to roots that are reduced
(Fig 39.1a)

Once a shoot reaches the sunlight, its morphology and biochemistry undergo profound
changes collectively called greening: the elongation rate of the stems slows, the leaves
expand, the roots start to elongate, and the entire shoot begins to produce chlorophyll; in
short, it begins to resemble a typical plant (Fig 39.1b)

In the following discussion, we use this greening response as an example of how a plant
cell receives a signal -- in this case, light -- and how this reception is transduced into a
response (greening). Along the way we will explore how studies of mutants have provided
valuable insights into the roles played by various molecules in the 3 stages of cell-signal
processing: reception, transduction, and response (Fig 39.2).
 Fig 39-1. Light-induced greening of dark-sprouted potatoes
(a) A dark-grown potato has tall, spindly stems and nonexpanded leaves--a morphological
adaptation that enables the shoots to penetrate the soil. The roots are short, but little water is
lost by the shoots. (b) After a week’s exposure to natural daylight, the potato plant begins to
resemble a typical plant with broad green leaves, short sturdy stems, and long roots. This
transformation begins with the reception of light by a specific pigment
 Fig 39-2. Review of a general model for signal-transduction pathways
A hormone or other signal binding to a specific receptor stimulates the cell to produce
second messengers. Second messengers trigger the cell’s various responses to the
original signal. In this diagram, the receptor is on the surface of the target cell. In
other cases, hormones enter cells and bind to specific receptors inside
Reception

Signals, whether internal or external, are first detected by receptors, proteins that
undergo conformational changes in response to a specific stimulus.

The receptor involved in greening in plants is called phytochrome, which consists of a

light-absorbing pigment attached to a specific protein.

Unlike many receptors, which are built into the plasma membrane, the phytochrome that
functions in greening occurs in the cytoplasm

Phytochrome functions in light detection in the greening process.

Transduction

The greening response is triggered by extremely low levels of light.

Receptors such as phytochrome are sensitive to very weak environmental and chemical
signals.

How is the information from these extremely weak signals amplified, and how is their
reception transduced into a specific response by the plant? The answer is second
messengers -- small, internally produced chemicals that transfer and amplify the signal
from the receptor to proteins that cause the specific response

In the greening response, for example, each activated phytochrome may give rise to
hundreds of molecules of a second messenger, each of which, in turn, may lead to the
activation of hundreds of molecules of a specific enzyme.

By such mechanisms, the second messenger of a signal transduction pathway leads to a

rapid amplification of the signal. In Chapter 11, we examined this role of second
messengers in general (Fig 11.12).

Here, let’s specifically consider the production of second messengers and their function in
the greening response (Fig 39.3)
Figure 11.12
Figure 11.11
 Fig 39-3. An example of signal transduction in plants:
the role of phytochrome in the greening response
The light signal is detected by the phytochrome receptor, which then activates at least two signal-
transduction pathways involving G-proteins. (1) One pathway leads to cGMP as a second messenger that
activates a protein kinase cascade (see Fig 11.11) (2) The other pathway leads to formation of a Ca2+1-
calmodulin complex that activates a specific protein kinase (3) Both pathways lead to expression of genes
for the proteins that function in the greening response. The main point of this diagram: Signal transduction
pathways in plants are variations of the general schemes you learned about in Chapter 11, involving such
components as G-proteins, second messengers, and protein kinases
Fig 39.3 refer that :

Phytochrome is such a receptor.

Light causes phytochrome to undergo a conformational change, and the phytochrome then
interacts with a specific G-protein.

During activation, guanosine diphosphate (GDP) that is bound to the inactive G-protein is
displaced by guanosine triphosphate (GTP).

The G-protein, now in its active form, in turn activates other enzymes in the signal-transduction
pathway that leads to greening

The cyclic nucleotides are second messengers that include cyclic adenosine monophosphate
(cyclic AMP) and cyclic guanosine monophosphate (cyclic GMP).

Experiments indicate that cyclic GMP (cGMP) is involved in the greening process

Changes in cytosolic Ca+ also play an important role in phytochrome signal transduction.
However, a wide range of hormonal and environmental stimuli can cause brief increases in
cytosolic Ca+.

In many cases, Ca2+ then binds directly to small proteins called calmodulins. The Ca+-
calmodulin complex then binds to and activates several enzymes, most notably certain types of
protein kinases.

Fig 39.3 that phytochrome activation during the greening mechanism results in both cGMP and
Ca+-calmodulin as second messengers
Response

Ultimately, a signal-transduction pathway leads to the regulation of one or more cellular activities.

The 2 main mechanisms by which a signaling pathway can activate an enzyme are by stimulating
transcription of mRNA for the enzyme or by activating existing enzyme molecules (post-translational
modification)

Transcriptional Regulation. Transcription factors bind directly to specific regions of DNA and control
the transcription of specific genes (see Fig 19.8).

In the case of phytochrome-induced greening, several transcription factors are activated by

phosphorylation in response to the appropriate light conditions. The activation of some of these
transcription factors depends upon cyclic GMP, whereas the activation of others requires Ca+-
calmodulin.

Post-Translational Modification of Proteins. Although the synthesis of new proteins by transcription

and translation are important molecular events associated with greening, so are post-translational
modifications of existing proteins.

Most often these existing proteins are modified by phosphorylation, the addition of a P group onto the
protein. The diverse proteins called protein kinases catalyze this phosphorylation of target proteins (see
Fig 11.11).

By such mechanisms, many signal pathways ultimately regulate the synthesis of new proteins, usually
by turning specific genes on or off (Fig 39.3)

The Greening Proteins. What sorts of proteins are either newly transcibed or activated by
phosphorylation during the greening process? Many are enzymes that function in photosynthesis
directly; others are enzymes involved in supplying the chemical precursors necessary for chlorophyll
production; still others affect the levels of plant hormones that regulate growth
Figure 19.8
Figure 11.11
 Fig 39-3. An example of signal transduction in plants:
the role of phytochrome in the greening response
The light signal is detected by the phytochrome receptor, which then activates at least two signal-
transduction pathways involving G-proteins. (1) One pathway leads to cGMP as a second messenger that
activates a protein kinase cascade (see Fig 11.11) (2) The other pathway leads to formation of a Ca2+1-
calmodulin complex that activates a specific protein kinase (3) Both pathways lead to expression of genes
for the proteins that function in the greening response. The main point of this diagram: Signal transduction
pathways in plants are variations of the general schemes you learned about in Chapter 11, involving such
components as G-proteins, second messengers, and protein kinases
 2. PLANT RESPONSES TO HORMONES

* Research on how plants grow toward light led to the discovery of plant hormones 

* Plant hormones help coordinate growth, development, and responses to

environmental stimuli

The word hormone is derived from a Greek verb meaning "to excite." Found in all multicellular
organisms, hormones are chemical signals that coordinate the parts of the organism. By
definition, a hormone is a compound that is produced by one part of the body and then
transported to other parts of the body, where it binds to a specific receptor and triggers responses
in target cells and tissues. Another important characteristic of hormones is that only minute
amounts are required to induce substantial change in an organism. Hormone concentrations or
the rates of their transport can change in response to environmental stimuli. Often the response of
a plant is governed by the interaction of two or more hormones .
2.1. Research on how plants grow toward light
led to the discovery of plant hormones

Any growth response that results in curvatures of whole plant organs toward or away from
stimuli is called a tropism (from the Greek tropos , turn).

The growth of a shoot toward light is called positive phototropism (growth away from light
is negative phototropism).

Fig 39-4. Early experiments of phototropism. Only the tip of the coleoptile can sense the
direction of light, but the bending response occurs some distance below the tip. A signal of
some kind must travel downward from the tip. The signal can pass through a permeable
barrier (gelatin block) but not through a solid barrier (a mineral called mica), suggesting that
the signal for phototropism is a mobile chemical

Fig 39-5. The Went experiments. Some chemical (indicated by pink) that can pass into an
agar block from a coleoptile tip stimulates elongation of the coleoptile when the block is
substituted for a tip. If the block is placed off-center on the top of a decapitated coleoptile
kept in the dark, the organ bends as if responding to illumination from one side. The
chemical is the hormone auxin, which stimulates elongation of cells in the shoot

The classical hypothesis for what causes grass coleoptiles to grow toward light, based on
the work of the Darwins and Went, is that an asymmetrical distribution of auxin moving
down from the coleoptile tip causes cells on the darker side to elongate faster than cells on
the brighter side
 Fig 39-4. Early experiments of phototropism
Only the tip of the coleoptile can sense the direction of light, but the bending response occurs
some distance below the tip. A signal of some kind must travel downward from the tip. The
signal can pass through a permeable barrier (gelatin block) but not through a solid barrier (a
mineral called mica), suggesting that the signal for phototropism is a mobile chemical
 Fig 39-5. 
The Went experiments.
Some chemical (indicated
by pink) that can pass into
an agar block from a
coleoptile tip stimulates
elongation of the coleoptile
when the block is
substituted for a tip. If the
block is placed off-center
on the top of a decapitated
coleoptile kept in the dark,
the organ bends as if
responding to illumination
from one side. The
chemical is the hormone
auxin, which stimulates
elongation of cells in the
shoot
2.2. Plant hormones help coordinate
growth, development, and responses to environmental stimuli

TABLE 39.1 previews some of the major classes of plant hormones: auxin, cytokinins,
gibberellins, abscisic acid, ethylene, and brassinosteroids. Notice that all of the
hormones shown in TABLE 39.1 are relatively small molecules. Their transport from cell
to cell often involves passage across cell walls, a pathway that blocks the movement of
large molecules

In general, hormones control plant growth and development by affecting the division,
elongation, and differentiation of cells. Some hormones also mediate shorter-term
physiological responses of plants to environmental stimuli

Plant hormones are produced in very low concentrations, but a minute amount of
hormone can have a profound effect on the growth and development of a plant organ

Signal transduction pathways amplify the hormonal signal and connect it to a cell’s
specific responses.

Response to a hormone usually depends not so much on the absolute amount of that
hormone as on its relative concentration compared with other hormones
Auxin (IAA, IBA, NAA, 2.4D)

Produce: Embryo of seed, meristems of apical buds, young leaves

Function: Stimulates stem elongation (low concentration only) root growth, cell differentiation, and
branching; regulates development of fruit; enhances apical dominance; functions in phototropism and
gravitropism.

Moving: Auxin seems to be transported directly through parenchyma tissue, from one cell-cell. It
moves only from shoot tip to base, not in the reverse direction.

This unidirectional transport of auxin is called polar transport . Polar auxin transport requires energy.

Fig 39.6 illustrates how proton pumps in the plasma membrane, driven by ATP, couple metabolic
energy to auxin transport

The Role of Auxin in Cell Elongation. The apical meristem of a shoot is a major site of auxin
synthesis. As auxin from the shoot apex moves down to the region of cell elongation (see Chapter 35),
the hormone stimulates growth of the cells, probably by binding to a receptor built into the plasma
membrane

According to a proposal called the acid growth hypothesis, proton pumps play a major role in the
growth response of cells to auxin. In a shoot’s region of elongation, auxin stimulates plasma
membrane proton pumps, an action that, within minutes, both increases the voltage across the
membrane (membrane potential) and lowers the pH in the cell wall (Fig 39.7)

Auxin also alters gene expression rapidly, causing cells in the region of elongation to produce new
proteins within minutes
Lateral and Adventitious Root Formation. Auxins are used commercially in the
vegetative propagation of plants by cuttings

Auxins as Herbicides. Synthetic auxins, such as 2,4-dinitrophenol (2,4-D), are widely

used as herbicides

Other Effects of Auxin. In addition to stimulating cell elongation for primary growth, auxin
affects secondary growth by inducing cell division in the vascular cambium and by
influencing the differentiation of secondary xylem
 Fig 39-6. 
Polar auxin transport: a
chemiosmotic model.
In growing shoots,
auxin is transported
unidirectionally, from
the apex down the
shoot. Along this
pathway, the hormone
enters a cell at the
apical end, exits at the
basal end, diffuses
across the wall, and
enters the apical end of
the next cell
Fig 39-7. Cell elongation in response to auxin: the acid growth hypothesis
Cytokinins (BA, kinetin, 2iP, zeatin, TDZ)

Produce: Synthesized in roots and

Moving: Transported to other organs

Function: Affect root growth and differentiation; stimulate cell division and growth;
stimulate germination; delay senescence

Despite much effort, the enzyme that produces cytokinins has neither been purified
from plants nor has the gene that encodes for it been identified . Regardless of the
source of cytokinins, plant cells do have cytokinin receptors

Control of Cell Division and Differentiation. Cytokinins are produced in actively

growing tissues, particularly in roots, embryos, and fruits

Control of Apical Dominance. Cytokinins, auxin, and other factors interact in the

control of apical dominance, the ability of the terminal bud to suppress the development
of axillary buds (Fig 39.8)

Anti-Aging Effects. Cytokinins retard the aging of some plant organs by inhibiting

protein breakdown, by stimulating RNA and protein synthesis, and by mobilizing
nutrients from surrounding tissues
 Fig 39-8. Apical dominance
(a) Auxin from the apical bud inhibits the growth of axillary buds. This favors elongation of
the shoot’s main axis. Cytokinins, which are transported upward from roots, counter auxin,
stimulating the growth of axillary buds. This explains why, in many plants, axillary buds near
the shoot tip are less likely to grow than those closer to the roots. (b) Removal of the apical
bud from the same plant enabled lateral branches to grow.
Gibberellins (GA3)

Produce: Meristems of apical buds and roots, young levels, embryo

Moving: Transported to other organs

Function: Promote seed and bud germination, stem elongation, and leaf growth; stimulate
flowering and development of fruit; affect root growth and differentiation

In 1926, the Japanese plant pathologist E. Kurosawa discovered that a fungus of the genus
Gibberella caused this "foolish seedling disease" (Fig 39.9).

Stem Elongation. Roots and young leaves are major sites of gibberellin production. Gibberellins
stimulate growth in both the leaves and the stem, but they have little effect on root growth. In
stems, gibberellins stimulate cell elongation and cell division (Fig 39.10)

Fruit Growth. In many plants, both auxin and gibberellins must be present for fruit to set. The
most important commercial application of gibberellins is in the spraying of Thompson seedless
grapes (Fig 39.11). The hormone makes the individual grapes grow larger, a trait prized by the
consumer, and it makes the internodes of the grape bunch elongate, allowing more space for the
individual grapes.

Germination. The embryo of seeds is a rich source of gibberellins. After water is imbibed, the
release of gibberellins from the embryo signals the seeds to break dormancy and germinate

Gibberellins support the growth of cereal seedlings by stimulating the synthesis of digestive
enzymes such as α-amylase that mobilize stored nutrients (see Fig 38.13).
 Fig 39-9. "Foolish seedling disease" in rice
The spindly rice plants on the right are infected with the fungus Gibberella .
The pathogen secretes gibberellins, a growth stimulant that uninfected plants (left) also
produce, but in smaller quantity
 Fig 39-10. 
Treating pea dwarfism
with a growth hormone.
Contrast the untreated
dwarf seedlings on the
left (control) with the
dwarf seedlings on the
right, which were treated
five days earlier with an
application of 5mmg of
gibberellin
 Fig 39-11. The effect of gibberellin treatment on seedless grapes
The grape bunch on the left is an untreated control. The bunch on the right is growing
from a vine that was sprayed with a gibberellin during fruit development
Figure 38.13
Abscisic Acid (ABA)

Produce: Leaves, stems, roots, green fruit

Moving: Transported to other organs

Function: Inhibits growth; closes stomata during water stress; counteracts breaking of
dormancy

ABA generally slows down growth. Often ABA antagonizes the actions of the growth
hormones, and it is the ratio of ABA to one or more growth hormones that determines the final
physiological outcome

Seed Dormancy. Seed dormancy has great survival value because it ensures that the seed will
germinate only when there are optimal conditions of light, temperature, and moisture

The high levels of ABA in maturing seeds inhibit germination and induce the production of
special proteins that help the seeds withstand the extreme dehydration that accompanies
maturation. Many types of dormant seeds will germinate when ABA is removed or inactivated in
some way. The seeds of some desert plants break dormancy only when heavy rains wash ABA
out of the seed

Abscisic acid induces dormancy in seeds. When its action is blocked--in this case, by a
mutation affecting an ABA-regulated transcription factor--precocious germination results (Fig
39.12)

Drought Stress. ABA is the primary internal signal that enables plants to withstand drought.
When a plant begins to wilt, ABA accumulates in leaves and causes stomata to close rapidly,
reducing transpiration and preventing further water loss.
 Fig 39-12. 
Precocious
germination of mutant
maize seeds.
Abscisic acid induces
dormancy in seeds.
When its action is
blocked--in this case,
by a mutation affecting
an abscisic acid-
regulated transcription
factor--precocious
germination results
Ethylene

Produce: Tissues of ripening fruits, nodes of stems, aging leves and flowers

Moving: Transported to other organs

Function: Promotes fruit ripening, opposes some auxin effects; promotes or inhibits growth and
development of roots, leaves, and flowers, depending on species

The Triple Response to Mechanical Stress: Using Mutants to Dissect a Signal-Transduction

Pathway. As the stem pushes against the obstacle, the mechanical stress in its delicate tip induces the
seedling to start producing ethylene. Ethylene, in turn, instigates the seedling to perform a growth
maneuver called the triple response that enables it to circumvent the obstacle.

The 3 parts of this response, which you can see in Fig 39.13, are a slowing of stem elongation, a
thickening of the stem (which makes it stronger), and a curvature that causes the stem to start growing
horizontally

Ethylene-insensitive (ein) mutants fail to undergo the triple response after exposure to ethylene (Fig
39.14a). The phenotype of such ethylene-overproducing (eto) mutants can be restored to wild-type by
treating the seedlings with inhibitors of ethylene synthesis. Still other mutants, called constitutive triple-
response (ctr) mutants, undergo the triple response in air but do not respond to inhibitors of ethylene
synthesis (Fig 39.14b). The affected gene in ctr mutants turns out to code for a protein kinase . Fig
39.15 summarizes the responses of ein , eto , and ctr mutants to ethylene and ethylene synthesis
inhibitors

Apoptosis: Programmed Cell Death. Consider the shedding of a leaf in autumn or the death of an
annual after flowering. The cells, organs and plants that are genetically programmed to die on a
particular schedule do not simply shut down their cellular machinery and await death. Rather, the onset
of programmed cell death, called apoptosis, is one of the busiest times in a cell’s life, requiring new
gene expression.
Leaf Abscission. The loss of leaves each autumn is an adaptation that keeps deciduous trees from
desiccating during winter when the roots cannot absorb water from the frozen ground

When an autumn leaf falls, the breaking point is an abscission layer located near the base of the petiole.
The small parenchyma cells of this layer have very thin walls, and there are no fiber cells around the
vascular tissue. The abscission layer is further weakened when enzymes hydrolyze polysaccharides in
the cell walls. Finally, the weight of the leaf, with the help of wind, causes a separation within the
abscission layer . Even before the leaf falls, a layer of cork forms preventing pathogens from invading
the plant (Fig 39.16).

A change in the balance of ethylene and auxin controls abscission

Fruit Ripening. Immature (unripe) fruits that are tart, hard, and green become edible at the time of seed
maturation. A burst of ethylene production in the fruit triggers this ripening

A chain reaction occurs during ripening: ethylene triggers ripening, and ripening, in turn, triggers
even more ethylene production -- one of the rare examples of positive feedback in physiology (Fig 1.8).
The result is a huge burst in ethylene production

Given the importance of ethylene in the post-harvest physiology of fruits, the genetic engineering of
ethylene signal transduction pathways has potentially important commercial applications

Brassinosteroids

Produced in seeds, fruits, shoots, leaves, and floral buds. Function in inhibits root growth; retards leaf
abscission; promotes xylem differentiation

It is important to remember that these plant hormones (chemical signals) are components of control
systems that tune a plant’s growth, development, reproduction, and physiology to the environment
 Fig 39-13. Ethylene induces the triple response in pea seedlings
Either artificial treatment with the gaseous hormone ethylene or natural production of the hormone
triggered by mechanical stress causes the stems of germinating pea seedlings to elongate less rapidly,
to become thicker, and to grow horizontally--the triple response. This growth response is an adaptation
that helps the seedling circumvent obstacles encountered during soil penetration. This experiment
shows the extent of the triple response in pea seedlings treated with different concentrations of
ethylene
Fig 39-14. Ethylene triple-response mutants
 Fig 39-15. 
Ethylene signal-
transduction
mutants can be
distinguished by
their different
responses to
experimental
treatments
 Fig 39-16. 
Abscission of
a maple leaf.
Abscission is controlled
by a change in the
balance of ethylene and
auxin. The abscission
layer can be seen here
as a vertical band at the
base of the petiole. After
the leaf falls, a protective
layer of cork becomes
the leaf scar that helps
prevent pathogens from
invading the plant (LM)
Figure 1.8
 3. PLANT RESPONSES TO LIGHT

* Blue-light photoreceptors are a heterogeneous group of pigments 

* Phytochromes function as photoreceptors in many plant responses to light 

* Biological clocks control circadian rhythms in plants and other eukaryotes 

* Light entrains the biological clock Photoperiodism synchronizes many plant responses to

changes of season

Light is an especially important environmental factor in the lives of plants. In addition to being
required for photosynthesis, light also cues many key events in plant growth and development.
These effects of light on plant morphology are what plant biologists call photomorphogenesis.
Light reception is also important in allowing plants to measure the passage of days and
seasons. Action spectra reveal that red and blue light are the most important colors in regulating
a plant’s photomorphogenesis. These observations led researchers to two major classes of light
receptors: a heterogeneous group of blue-light photoreceptors and a family of photoreceptors
called phytochromes that absorb mostly red light
3.1. Blue-light photoreceptors are a heterogeneous group of pigments

In the 1990s, molecular biologists analyzing Arabidopsis mutants found that plants actually use at least
three completely different types of pigments to detect blue light: cryptochromes (for the inhibition of
hypocotyl elongation), phototropin (for phototropism), and a carotenoid-based photoreceptor called
zeaxanthin (for stomatal opening).

Fig 39-17. 
Action spectrum for blue-light-
stimulated phototropism.
(a) Phototropic bending toward
light is controlled by the blue-light
photoreceptor called phototropin.
In this action spectrum for
phototropism in maize (Zea mays
) coleoptiles, you can see that
only light of wavelengths below
500 nm (blue and ultraviolet) are
effective in inducing curvature.
(b) The upper photograph in each
pair was taken at the beginning
of the experiment; the lower
photos were taken after a 90-
minute exposure to side lighting
of the indicated colors.
3.2. Phytochromes function as photoreceptors
in many plant responses to light
The Phytochrome Switch and Seed Germination

The successful sprouting of many types of small seeds, such as lettuce, requires that they
germinate only when conditions, especially the light environment, are near optimal . The effects
of red (660nm) increase and far-red (730nm) light decrease germination, are reversible (Fig
39.18)

The photoreceptor responsible for these opposing effects of red and far-red light is a
phytochrome (Fig 39.19)

The chromophore of a phytochrome reverts back and forth between 2 isomeric forms. This Pr
< Pfr interconversion acts as a switching mechanism that controls various light-induced events
in the life of the plant (Fig 39.20)

The Phytochrome Switch and Shade Avoidance

The phytochrome system also provides the plant with information about the quality of light.
Sunlight includes both red and far-red radiation

The "shade-avoidance" response of a tree that requires relatively high light intensity

Direct sunlight increases the proportion of Pfr, which stimulates branching and inhibits vertical
growth
 Fig 39-18. Phytochrome regulation of lettuce seed germination
The control seeds, on the left, were kept in the dark. The various batches
of experimental seeds received flashes of light. The bars below the photos
indicate the sequences of red and far-red light flashes
Fig 39-19. Structure of a phytochrome
 Fig 39-20. Phytochrome: a molecular switching mechanism
Absorption of red light causes the bluish Pr to change to the blue-greenish Pfr. Far-
red light reverses this conversion. In most cases, it is the Pfr form of the pigment that
switches on physiological and developmental responses in the plan
3.3. Biological clocks control circadian rhythms
in plants and other eukaryotes

Many legumes lower their leaves in the evening and raise them in the morning (Fig 39.21)

Physiological cycles with a frequency of about 24-hours and that are not directly paced by
any known environmental variable are called circadian rhythms (from the Latin circa,
approximately, and dies, day)

Circadian rhythms are truly prompted by an internal clock, or are they merely daily
responses to some subtle but pervasive environmental cycle

If an organism is kept in a constant environment, its circadian rhythms deviate from a 24-
hour period (a "period" is the duration of one cycle)

A leading hypothesis is that biological timekeeping may depend on synthesis of a protein

that regulates its own production through feedback control.

This protein may be a transcription factor that inhibits transcription of the gene that
encodes for the transcription factor itself.

the concentration of this transcription factor may accumulate during the first half of the
circadian cycle, and then decline during the second half, due to self-inhibition of its own
production
Fig 39-21. Sleep movements of a bean plant. Leaf position at noon (top) and at midnight
The movements are caused by reversible changes in the turgor pressure of cells on
opposing sides of the pulvini, swollen regions of the petiole
3.4. Light entrains the biological clock

What entrains the biological clock to precisely 24 hours every day?

The answer is light.

Both phytochrome and blue-light photoreceptors can entrain circadian rhythms in

plants, but our understanding of how phytochrome does this is more complete

Interactions between phytochrome and the biological clock enable plants to measure the
passage of night and day in nature.

The relative lengths of night and day, however, change over the course of the year (except
at the Equator). Plants use this change to mark the passage of seasons.
3.4. Photoperiodism synchronizes
many plant responses to changes of season

The environmental stimulus plants use most often to detect the time of year is the
photoperiod, the relative lengths of night and day. A physiological response to photoperiod,
such as flowering, is called photoperiodism

Photoperiodism and the Control of Flowering

Short-day plant: need short day to flower. Long-day plant: need long day to flower.
Neutral-day plant : any time to flower

Critical Night Length. The actual number of hours in the critical night length is specific to
each species of plant.

Red light is the most effective color in interrupting the nighttime portion of the photoperiod.
Action spectra and photoreversibility experiments show that phytochrome is the pigment
that receives the red light (Fig 39.23)

Plants measure night length very accurately; some short-day plants will not flower if
night is even one minute shorter than the critical length. Some plant species always flower
on the same day each year
Fig 39-22. Photoperiodic control of flowering
Fig 39-23. Reversible effects of red and far-red light on photoperiodic response
A flash of red light shortens the dark period. A subsequent flash of far-red light
cancels the effect of the red flash
Is There a Flowering Hormone ?

Buds produce flowers, but it is leaves that detect photoperiod. If all leaves are removed,
however, the plant is blind to photoperiod.

When the photoperiodic requirement for flowering is met, leaves must send a signal to the buds
cuing them to develop as flowers.

Most plant physiologists believe the flowering signal is a hormone or some change in the
relative concentrations of two or more hormones (Fig 39.24).

Meristem Transition from Vegetative Growth to Flowering

Whatever combination of environmental cues (such as photoperiod or vernilization) and internal

signals (such as hormones) is necessary for flowering to occur,

the outcome is the transition of a bud’s meristem from a vegetative state to a flowering state

This transition requires changes in the expression of genes that regulate pattern formation

Meristem-identity genes that specify that the bud will now form a flower instead of a vegetative
shoot must first be switched on.

Then the organ-identity genes that specify the spatial organization of the floral organs --
sepals, petals, stamens, and carpels -- are activated in the appropriate regions of the meristem
(see Fig 21.19 and 35.12).
 Fig 39-24. 
Experimental evidence
for a flowering
hormone(s).
If a plant that has been
induced to flower by
photoperiod is grafted to
a plant that has not been
induced, both plants
flower, indicating the
transmission of a flower-
inducing substance. This
works in some cases
even if one is a short-day
plant and the other is a
long-day plant
Figure 21.19
Figure 35.12
 4. PLANT RESPONSES
TO ENVIRONMENTAL STIMULI OTHER THAN LIGHT

* Plants respond to environmental stimuli through

a combination of developmental and physiological mechanisms

Plants can neither migrate to a watering hole when water is scarce nor seek shelter when the
weather is too windy. A seed, landing upside down in the soil, cannot maneuver itself into an upright
position. Because of their immobility, plants must adjust to a wide range of environmental
circumstances through developmental and physiological mechanisms. Natural selection has refined
these responses. Light is so important in the life of a plant that we devoted the entire previous
section to a plant’s reception of and response to this one environmental factor. In this section, we
examine responses to some of the other environmental stimuli that a plant commonly faces in its
"struggle for existence."
4.1. Plants respond to environmental stimuli
through a combination of developmental and physiological mechanisms

Responses to Gravity

Auxin plays a major role in gravitropic responses.

Plants may tell up from down by the settling of statoliths, specialized plastids containing
dense starch grains, to the lower portions of cells (Fig 39.25)

The Ca+ and auxin accumulate on the lower side of the root’s zone of elongation.

At high concentration, auxin inhibits cell elongation, an effect that slows growth on the lower
side of the root. The more rapid elongation of cells on the upper side causes the root to curve
as it grows. This tropism continues until the root is growing straight down
Fig 39-25. The statolith hypothesis for root gravitropism
These corn roots were placed on their sides and
photographed before (top) and 1.5 hours after initiation of the
gravitropic response. The light micrographs show the
locations of statoliths, modified plastids, within cells of the root
cap. The settling of statoliths to the low points of these cells
may be a gravity-sensing mechanism that leads to the
redistribution of auxin and the differential rates of elongation
by cells on opposite sides of the root
Responses to Mechanical stimuli

Plants are very sensitive to mechanical stress: The term thigmomorphogenesis (from the
Greek thigma, touch) refers to the changes in form that result from mechanical
perturbation.

Rubbing the stems of a young plant a few times results in plants that are shorter than
controls (Fig 39.26).

Some plant species have become, over the course of their evolution, "touch specialists"

Directional growth in response to touch is called thigmotropism, and it allows the vines to
take advantage of whatever mechanical supports it comes across as it climbs upward
toward a forest canopy

Touch specialists are plants that undergo rapid leaf movements in response to mechanical
stimulation. For example, when the compound leaf of the sensitive plant Mimosa is
touched, it collapses and its leaflets fold together (Fig 39.27)
 Fig 39-26. 
Altering gene expression
by touch in Arabidopsis .
The shorter plant on the left
was touched twice a day.
The unmolested plant
(right) grew much taller
 Fig 39-27. 
Rapid turgor movements by
the sensitive plant
(Mimosa pudica ).
(a) In the unstimulated plant,
leaflets are spread apart. (b)
Within a second or two of
being touched, the leaflets
have folded together. (c) In
these light micrographs of a
leaflet pair in the closed
(stimulated) state, you can
see the motor cells in the
sectioned pulvini (motor
organs). The curvature of a
pulvinus is caused by motor
cells on one side losing
water and becoming flaccid
while cells on the opposite
side retain their turgor
Responses to Stress

Drought. On a bright, warm, dry day, a plant may be stressed by a water deficiency
because it is losing water by transpiration faster than the water can be restored by uptake
from the soil

Flooding. An overwatered houseplant may suffocate because the soil lacks the air spaces
that provide oxygen for cellular respiration in the roots (Fig 39.28)

Heat Stress. Excessive heat can harm and eventually kill a plant by denaturing its
enzymes and damaging its metabolism in other ways

Cold Stress. One problem plants face when the temperature of the environment falls is a
change in the fluidity of cell membranes

Freezing is a more severe version of cold stress. At subfreezing temperatures, ice

forms in the cell walls and intercellular spaces of most plants.
 Fig 39-28. A developmental response of corn roots to flooding and oxygen deprivation
(a) A transverse section of a control root grown in an aerated hydroponic medium. (b) An
experimental root grown in a nonaerated hydroponic medium. Ethylene-stimulated
apoptosis (programmed cell death) creates the air tubes
 5. PLANT DEFENSE:
RESPONSES TO HERBIVORES AND PATHOGENS

* Plants deter herbivores with both physical and chemical defenses 

* Plants use multiple lines of defense against pathogens

Plants do not exist in isolation, but interact with many other species in their communities. Some of
these interspecific interactions--for example, the associations of plants with fungi in mycorrhizae
(see Chapter 37) or with insect pollinators (see Chapter 38)--are mutually beneficial. Most of the
interactions that plants have with other organisms, however, are not beneficial to the plant. As
primary producers, plants are at the base of most food webs and are subject to attack by a wide
range of plant-eating (herbivorous) animals. A plant is also subject to infection by a diversity of
pathogenic viruses, bacteria, and fungi that have the potential to damage tissues or even kill the
plant. Plants counter these threats with defense systems that deter herbivory and prevent infection
or combat pathogens that do manage to infect the plant
5.1. Plants deter herbivores with both physical and chemical defenses

Fig 39-29. 
A corn leaf recruits
a parasitoid wasp as
a defensive response
to an herbivore, an
army-worm caterpillar
5.2. Plants use multiple lines of defense against pathogens

A plant’s first line of defense against infection is the physical barrier of the plant’s "skin," the
epidermis of the primary plant body and the periderm of the secondary plant body

Gene-for-Gene Recognition

Plants are generally resistant to most pathogens. Pathogens against which a plant has little
specific defense are said to be virulent.

Specific resistance to a plant disease is based on what is called gene-for-gene recognition,

because it depends on a precise match-up between a genetic allele in the plant and an allele in
the pathogen.

This occurs when a plant with specific dominant resistance alleles (R ) recognizes those
pathogens that possess complementary avirulence (Avr ) alleles (Fig 39.30)

Even if a plant is infected by a virulent strain of a pathogen -- one for which that particular plant
has no genetic resistance -- the plant is able to mount a localized chemical attack in response
to molecular signals released from cells damaged by the infection.

Plant + virulent  chemical attack (elicitors / oligosaccharides)  phytoalexin (antimicroial

compounds)

Molecules called elicitors, often cellulose fragments called oligosaccharins released by cell-
wall damage, induce the production of antimicrobial compounds called phytoalexins.

Infection also activates genes that produce PR proteins (for pathogenesis-related). Some of
these proteins are antimicrobial, attacking molecules in the cell wall of a bacterium
 Fig 39-30. 
Gene-for-gene resistance of
plants to pathogens.
(a) Resistance occurs when
the plant has a particular
dominant R allele that
corresponds to a specific
dominant Avr allele in the
pathogen. R genes probably
code for specific receptors. Avr
genes produce compounds
that function in the pathogen
but also act as ligands that
bind specifically to the host-
plant cell’s receptors. Disease
occurs if there is no gene-for-
gene recognition because (b)
the pathogen has no dominant
Avr allele matching an R allele
in the plant, (c) the plant has
no dominant R allele matching
an Avr allele in the pathogen,
or (d) both pathogen and plant
lack alleles upon which
recognition could be based