Biology: Plant Signals and Behavior

BIOLOGY TENTH EDITION
Global Edition
Campbell • Reece • Urry • Cain • Wasserman • Minorsky • Jackson
39
Plant Signals
and Behavior
Lecture Presentation by
Nicole Tunbridge and
Kathleen Fitzpatrick
© 2015 Pearson Education Ltd

Stimuli and a Stationary Life
a) Plants receive signals from the environment and

respond by altering growth and development
a)For example, the bending of a dodder seedling toward
a host plant occurs in response to chemicals released
by the host

Figure 39.1

Figure 39.1a

Concept 39.1: Signal transduction pathways link signal
reception to response
a) A potato left growing in darkness produces shoots

that look unhealthy, and it lacks elongated roots
b) These are morphological adaptations for growing in
darkness, collectively called etiolation
c) After exposure to light, a potato undergoes changes
called de-etiolation, in which shoots and roots grow
normally

Figure 39.2
(a) Before exposure to light (b) After a week’s exposure

to natural daylight

a) A potato’s response to light is an example of cell
signal processing
b) The stages are reception, transduction, and response

Figure 39.3
CELL CYTOPLASM
WALL
1 Reception 2 Transduction 3 Response
Receptor Relay proteins and Activation

of cellular
second messengers responses
Hormone or
environmental
stimulus Plasma membrane

Reception
a) Internal and external signals are detected by

receptors, proteins that change in response to
specific stimuli
b) In de-etiolation, the receptor is a phytochrome
capable of detecting light

Transduction
a) Second messengers transfer and amplify signals

from receptors to proteins that cause responses
b) Two types of second messengers play an important
role in de-etiolation: Ca2+ ions and cyclic GMP
(cGMP)
c) The phytochrome receptor responds to light by
a)Opening Ca2+ channels, which increases Ca2+ levels in
the cytosol
b)Activating an enzyme that produces cGMP
Figure 39.4-3
1 Reception 2 Transduction 3 Response
Transcription
CYTOPLASM
factor 1 NUCLEUS
Plasma cGMP Protein P
membrane kinase 1
Second
Phytochrome messenger Transcription
factor 2
activated produced
by light P
Cell
wall Protein
kinase 2
Transcription
Light
Translation
Ca2+ channel De-etiolation

opened (greening)
response
proteins
Ca2+

Response
a) A signal transduction pathway leads to regulation of

one or more cellular activities
b) In most cases, these responses to stimulation involve
increased activity of enzymes
c) This can occur by transcriptional regulation or post-
translational modification

Post-Translational Modification of Preexisting Proteins
a)Post-translational modification involves

modification of existing proteins in the signal
response
b)Modification often involves the phosphorylation of
specific amino acids
c) The second messengers cGMP and Ca2+ activate
protein kinases directly
d)Protein phosphatases “switch off” the signal
transduction pathways by dephosphorylating
proteins
Transcriptional Regulation
a)Specific transcription factors bind directly to

specific regions of DNA and control transcription
of genes
b)Some transcription factors are activators that
increase the transcription of specific genes
c) Other transcription factors are repressors that
decrease the transcription of specific genes

De-Etiolation (“Greening”) Proteins
a) De-etiolation activates enzymes that

a)Function in photosynthesis directly
b)Supply the chemical precursors for chlorophyll
production
c)Affect the levels of plant hormones that regulate growth

Concept 39.3: Responses to light are critical for plant
success
a)Light cues many key events in plant growth and

development
b)Effects of light on plant morphology are called
photomorphogenesis

a) Plants detect not only presence of light but also its
direction, intensity, and wavelength (color)
b) A graph called an action spectrum depicts relative
response of a process to different wavelengths
c) Action spectra are useful in studying any process that
depends on light

a)Different plant responses can be mediated by the
same or different photoreceptors
b)There are two major classes of light receptors:
blue-light photoreceptors and phytochromes

Blue-Light Photoreceptors
a) Various blue-light photoreceptors control hypocotyl

elongation, stomatal opening, and phototropism

Figure 39.15
1.0 436 nm
Phototropic effectiveness
0.8 Refracting prism
0.6
0.4
White
light
0.2
0
400 450 500 550 600 650 700
Wavelength (nm)
(a) Light wavelengths below 500 nm induce (b) Blue light induces the most curvature of
curvature. coleoptiles.

Phytochrome Photoreceptors
a) Phytochromes are pigments that regulate many of a

plant’s responses to light throughout its life
b) These responses include seed germination and
shade avoidance

Phytochromes and Seed Germination
a)Many seeds remain dormant until light and other

conditions are near optimal
b)In the 1930s, scientists at the U.S. Department of
Agriculture determined the action spectrum for
light-induced germination of lettuce seeds

a) Red light increased germination, while far-red light
inhibited germination
b) The effects of red and far-red light are reversible; the
final light exposure determines the response
c) The photoreceptors responsible for the opposing
effects of red and far-red light are phytochromes

Figure 39.16
Results
Red Dark Red Far-red Dark
Dark (control)
Red Far-red Red Dark Red Far-red Red Far-red

a)Phytochromes exist in two photoreversible states,
with conversion of Pr to Pfr triggering many
developmental responses
b)Red light triggers the conversion of Pr to Pfr
c) Far-red light triggers the conversion of Pfr to Pr
d)The conversion of Pr to Pfr is faster than the
reverse process
e)Sunlight, containing both red and far-red light,
increases the ratio of Pfr to Pr and triggers
germination
Figure 39.17
Responses to Pfr:
Red light • Seed germination
• Inhibition of vertical
Synthesis Pr Pfr growth and stimu-
lation of branching
Far-red • Setting internal clocks
light • Control of flowering
Slow conversion Enzymatic

in darkness destruction
(some species)

Phytochromes and Shade Avoidance
a)The phytochrome system also provides the plant

with information about the quality of light
b)Leaves in the canopy absorb red light
c) Shaded plants receive more far-red than red light
d)In the “shade avoidance” response, the
phytochrome ratio shifts in favor of Pr when a tree
is shaded
e)This shift induces vertical growth
Biological Clocks and Circadian Rhythms
a) Many plant processes oscillate during the day in

response to light and temperature changes
b) Many legumes lower their leaves in the evening and
raise them in the morning, even when kept under
constant light or dark conditions

Figure 39.18
Noon 10:00 PM

a) Circadian rhythms are cycles that are about 24
hours long and are governed by an internal “clock”
b) Circadian rhythms can be set to exactly 24 hours by
the day/night cycle
c) The clock may depend on synthesis of a protein
regulated through feedback control and may be
common to all eukaryotes

The Effect of Light on the Biological Clock
a) Phytochrome conversion marks sunrise and sunset,

providing the biological clock with environmental cues

Photoperiodism and Responses to Seasons
a)Photoperiod, the relative lengths of night and day,

is the environmental stimulus plants use most
often to detect the time of year
b)Photoperiodism is a physiological response to
photoperiod

Photoperiodism and Control of Flowering
a) Some processes, including flowering in many

species, require a certain photoperiod
b) Plants that flower when a light period is shorter than a
critical length are called short-day plants
c) Plants that flower when a light period is longer than a
certain number of hours are called long-day plants
d) Flowering in day-neutral plants is controlled by plant
maturity, not photoperiod

Critical Night Length
a) In the 1940s, researchers discovered that flowering
and other responses to photoperiod are actually
controlled by night length, not day length

a) Short-day plants are governed by whether the critical
night length sets a minimum number of hours of
darkness
b) Long-day plants are governed by whether the critical
night length sets a maximum number of hours of
darkness

Figure 39.19
24 hours
(a) Short-day
(long-night) plant
Light Darkness
Flash
Critical of light
dark period
(b) Long-day
(short-night) plant
Flash of light
a) Red light can interrupt the nighttime portion of the
photoperiod
b) A flash of red light followed by a flash of far-red light
does not disrupt night length
c) Action spectra and photoreversibility experiments
show that phytochrome is the pigment that detects
the red light

Figure 39.20
24 hours
R FR
R FR R
R FR R FR
Short-day Long-day
Critical dark period (long-night) (short-night)
plant plant
a) Some plants flower after only a single exposure to the
required photoperiod
b) Other plants need several successive days of the
required photoperiod
c) Still others need an environmental stimulus in
addition to the required photoperiod
a)For example, vernalization is a pretreatment with cold
to induce flowering

A Flowering Hormone?
a)Photoperiod is detected by leaves, which cue

buds to develop as flowers
b)The flowering signal molecule is called florigen
c) Florigen may be a protein governed by the
FLOWERING LOCUS T (FT) gene

Figure 39.21
24 hours 24 hours 24 hours
Graft
Short-day Long-day plant Long-day

plant grafted to plant
short-day plant

Concept 39.4: Plants respond to a wide variety of
stimuli other than light
a) Because of immobility, plants must adjust to a range

of environmental circumstances through
developmental and physiological mechanisms

Gravity
a)Response to gravity is known as gravitropism

b)Roots show positive gravitropism; shoots show
negative gravitropism
c) Plants may detect gravity by the settling of
statoliths, dense cytoplasmic components

Figure 39.22
Statoliths
20 µm
(a) Primary root of maize (b) Statoliths settling to

bending gravitropically the lowest sides of
(LMs) root cap cells (LMs)

a) Some mutants that lack statoliths are still capable of
gravitropism
b) Dense organelles, in addition to starch granules, may
contribute to gravity detection

Mechanical Stimuli
a) The term thigmomorphogenesis refers to changes

in form that result from mechanical disturbance
b) Rubbing stems of young plants a couple of times
daily results in plants that are shorter than controls

Figure 39.23

a) Thigmotropism is growth in response to touch
b) It occurs in vines and other climbing plants
c) Another example of a touch specialist is the sensitive
plant Mimosa pudica, which folds its leaflets and
collapses in response to touch
d) Rapid leaf movements in response to mechanical
stimulation are examples of transmission of electrical
impulses called action potentials

Figure 39.24
(a) Unstimulated state (leaflets spread (b) Stimulated state (leaflets folded)
apart)

Environmental Stresses
a) Environmental stresses have a potentially adverse

effect on survival, growth, and reproduction
b) Stresses can be abiotic (nonliving) or biotic (living)
c) Abiotic stresses include drought, flooding, salt stress,
heat stress, and cold stress
d) Biotic stresses include herbivores and pathogens

Drought
a) During drought, plants reduce transpiration by closing

stomata, reducing exposed surface area, and in some
species, shedding leaves

Flooding
a) Enzymatic destruction of root cortex cells creates air

tubes that help plants survive oxygen deprivation
during flooding

Figure 39.25
Vascular
cylinder
Air tubes
Epidermis
(a) Control root (b) Experimental root

100 µm 100 µm
(aerated) (nonaerated)

Salt Stress
a) Salt can lower the water potential of the soil solution

and reduce water uptake
b) Plants respond to salt stress by producing solutes
tolerated at high concentrations
c) This process keeps the water potential of cells more
negative than that of the soil solution

Heat Stress
a)Excessive heat can denature a plant’s enzymes

b)Transpiration helps cool leaves by evaporative
cooling
c)Heat-shock proteins help protect other proteins
from heat stress

Cold Stress
a)Cold temperatures decrease membrane fluidity

b)Altering lipid composition of membranes is a
response to cold stress
c) Freezing causes ice to form in a plant’s cell walls
and intercellular spaces
d)Many plants, as well as other organisms, have
antifreeze proteins that prevent ice crystals from
growing and damaging cells

Concept 39.5: Plants respond to attacks by pathogens
and herbivores
a) Plants use defense systems to deter herbivory,

prevent infection, and combat pathogens

Defenses Against Pathogens
a) A plant’s first line of defense against infection is the

barrier presented by the epidermis and periderm
b) Pathogens can enter through wounds or natural
openings, such as stomata

Immune Responses of Plants
a) The first line of immune defense depends on the

plant’s ability to recognize pathogen-associated
molecular patterns (PAMPs)
b) These molecular sequences are specific to certain
pathogens
c) PAMP recognition starts a chain of signaling events
leading to the production of antimicrobial chemicals
and toughening of the cell wall

a) Plants and pathogens have engaged in an
evolutionary arms race
b) Pathogens that have evolved the ability to deliver
effectors into plant cells can suppress PAMP-
triggered plant immunity
c) Effectors are pathogen-encoded proteins that cripple
the host’s innate immune system
d) A second level of plant immune defense evolved in
response to these pathogens

a) Effector-triggered immunity results from the action of
hundreds of disease resistance (R) genes
b) Each R protein is activated by a specific effector
c) R proteins activate plant defenses by triggering signal
transduction pathways
d) These defenses include the hypersensitive response
and systemic acquired resistance

The Hypersensitive Response
a) The hypersensitive response

a)Causes cell and tissue death near the infection site
b)Induces production of enzymes that attack the
pathogen
c)Stimulates changes in the cell wall that confine the
pathogen

Figure 39.26
Infected tobacco leaf with lesions
Signal 5
4
Hypersensitive Signal 6
3 response transduction
pathway
2 Signal transduction pathway
7 Acquired
resistance
1
R protein
Pathogen
Effector protein
Hypersensitive Systemic acquired

response resistance
Systemic Acquired Resistance
a) Systemic acquired resistance causes systemic

expression of defense genes and is a long-lasting
response
b) Methylsalicylic acid is synthesized around the
infection site and carried in the phloem to other
remote sites where it is converted to salicylic acid
c) Salicylic acid triggers the defense system to respond
rapidly to another infection

Defenses Against Herbivores
a) Herbivory, animals eating plants, is a stress that

plants face in any ecosystem
b) Plants counter excessive herbivory with physical
defenses, such as thorns and trichomes, and
chemical defenses, such as distasteful or toxic
compounds

Molecular-Level Defenses
a) Chemical compounds including terpenoids, phenolics,
and alkaloids can be produced to deter attackers

Figure 39.27a
MAKE CONNECTIONS:
Levels of Plant Defenses Against Herbivores
Molecular-Level Cellular-Level Tissue-Level
Organ-Level

Cellular-Level Defenses
a) Cells may be specialized to form trichomes, store
chemical deterrents, or produce irritants

Tissue-Level Defenses
a) Leaves may be toughened with sclerenchyma tissue

Organ-Level Defenses
a) Leaves can be modified into spines and bristles;
stems into thorns
b) Some species have leaves that appear partially
eaten, others have structures that mimic insect eggs

Organismal-Level Defenses
a) Plants may respond to attack by altering flowering
time

Figure 39.27b
MAKE CONNECTIONS:
Levels of Plant Defenses Against Herbivores
Organismal-Level
Community-Level
Population-Level

Population-Level Defenses
a) Some plants release chemicals in response to
herbivore attack that trigger defense responses in
nearby members of the population
b) Others populations synchronously produce massive
amounts of seeds after long intervals

Community-Level Defenses
a) Some plants “recruit” predatory animals that help
defend against specific herbivores

Biology: Plant Signals and Behavior

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BIOLOGY TENTH EDITION

Campbell • Reece • Urry • Cain • Wasserman • Minorsky • Jackson

© 2015 Pearson Education Ltd

a) Plants receive signals from the environment and

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a) A potato left growing in darkness produces shoots

© 2015 Pearson Education Ltd

(a) Before exposure to light (b) After a week’s exposure

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

1 Reception 2 Transduction 3 Response

Receptor Relay proteins and Activation

© 2015 Pearson Education Ltd

a) Internal and external signals are detected by

© 2015 Pearson Education Ltd

a) Second messengers transfer and amplify signals

1 Reception 2 Transduction 3 Response

Ca2+ channel De-etiolation

© 2015 Pearson Education Ltd

a) A signal transduction pathway leads to regulation of

© 2015 Pearson Education Ltd

a)Post-translational modification involves

a)Specific transcription factors bind directly to

© 2015 Pearson Education Ltd

a) De-etiolation activates enzymes that

© 2015 Pearson Education Ltd

a)Light cues many key events in plant growth and

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a) Various blue-light photoreceptors control hypocotyl

© 2015 Pearson Education Ltd

0.8 Refracting prism

© 2015 Pearson Education Ltd

a) Phytochromes are pigments that regulate many of a

© 2015 Pearson Education Ltd

a)Many seeds remain dormant until light and other

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

Red Dark Red Far-red Dark

Red Far-red Red Dark Red Far-red Red Far-red

© 2015 Pearson Education Ltd

Slow conversion Enzymatic

© 2015 Pearson Education Ltd

a)The phytochrome system also provides the plant

a) Many plant processes oscillate during the day in

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a) Phytochrome conversion marks sunrise and sunset,

© 2015 Pearson Education Ltd

a)Photoperiod, the relative lengths of night and day,

© 2015 Pearson Education Ltd

a) Some processes, including flowering in many

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

© 2015 Pearson Education Ltd

a)Photoperiod is detected by leaves, which cue

© 2015 Pearson Education Ltd