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Chapter 39
Plant Sensory Systems
BIOLOGY
Thirteenth Edition
Raven, Johnson, Mason, Losos,
Duncan
© 2023 McGraw Hill, LLC. All rights reserved. Authorized only for instructor use in the classroom.
No reproduction or further distribution permitted without the prior written consent of McGraw Hill, LLC.
Lecture Outline
39.1 Responses to Light
39.2 Responses to Gravity
39.3 Responses to Mechanical
Stimuli
39.4 Responses to Water and
Temperature
39.5 Hormones and Sensory
Systems
©Educational Images/Getty Images
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Responses to Light
Pigments are molecules that are capable of absorbing light
energy
• Some are used in photosynthesis
• Others detect light and mediate the plant’s response to it.
Photomorphogenesis
• Nondirectional, light-triggered development.
Phototropisms
• Directional growth responses to light.
Both compensate for inability to move
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Phytochrome
The pigment-containing protein phytochrome (P) is present in
all groups of plants
The molecule exists in 2 interconvertible forms
• Pr – absorbs red light at 660 nm.
• Biologically inactive.
• Converted to Pfr when red photons
available.
• Pfr – absorbs far-red light at 730 nm.
• Active form.
• Converted back to Pr when far-red photons available.
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How Phytochrome Works
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Pfr Regulation
• Amount of Pfr is also regulated by degradation
• Ubiquitin tags Pfr for transport to the proteasome
• Process of tagging and recycling Pfr is precisely regulated
to maintain needed amounts of phytochrome in the cell
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Components of Phytochrome
Phytochrome (P) consists of two parts:
• Chromophore which is light-receptive and causes
conformation changes
• Apoprotein which facilitates expression of light-response
genes
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Signaling Pathways
Phytochromes are involved in many signaling pathways that
lead to gene expression
• Pr is found in the cytoplasm.
• When it is converted to Pfr it enters the nucleus.
• Pfr binds with other proteins that form a transcription
complex, leading to the expression of light-regulated
genes.
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Pfr Regulates Gene Expression
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Protein Kinase-Signaling Pathways
• Phytochrome also works through protein kinase-signaling
pathways
• When Pr is converted to Pfr , its protein kinase domain
causes autophosphorylation or phosphorylation of another
protein
• This initiates a signaling cascade that activates
transcription factors leading to expression of light-
regulated genes
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Phosphorylation Initiates Signaling Cascade
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Different Phytochromes
Although we refer to phytochrome as a single molecule,
there are different forms that appear to have specific
functions
Five forms have been characterized in Arabidopsis
• PHYA through PHYE.
• Each form plays an overlapping but distinct role in the light
regulation of growth and development.
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Involvement of Phytochrome
Phytochrome is involved in
Seed germination.
• Inhibited by far-red light and stimulated by red light in many
plants.
• Only germinate when exposed to direct sunlight.
Shoot elongation.
• Etiolation is an energy conservation strategy where plants divert
energy to internode elongation when growing in the dark
• Caused by a lack of red light.
• Assume normal morphology when Pfr increases.
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Detection of plant crowding.
• Plants measure the amount of far-red light bounced back from
neighboring plants.
• The closer together plants are,
• the more far-red they perceive.
• The more likely they are to grow taller, outcompeting
others for sunshine
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Etiolation Regulation
©Niko Geldner, UNIL
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Response to Blue Light
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Phototropism
• Tropisms are directional growth responses
• Phototropisms contribute to the variety of forms we see
within a species as shoots grow toward light
• Connect environmental signal with cellular perception of
the signal, transduction into biochemical pathways, and
ultimately an altered growth response
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Blue-Light Receptors
Blue-light receptor phototropin 1 (PHOT1)
• 2 light-sensing regions – change conformation in response
to blue light.
• Stimulates the kinase region of PHOT1 to
autophosphorylate.
• Regulates the flux of auxin in shoots.
• By adjusting the distribution of this hormone, it changes the parts of
the shoot that are actively elongated.
• Elongation on one side of a shoot can lead to bending toward the
light.
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PHOT 1
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Circadian Clocks
“Around the day” rhythms are particularly common among
eukaryotes
Have four characteristics:
1. Continue in absence of external inputs
2. Must be about 24 hours in duration
3. Cycle can be reset or entrained
• Phytochrome action entrains daily cycles.
• Plants in darkness may become desynchronized.
4. Clock can compensate for differences in temperature
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Gravitropism
• Response of a plant to the gravitational field of the Earth
• Shoots exhibit negative gravitropism (growth away from
gravity)
• Roots have a positive gravitropic response (growth toward
gravity)
©Ray F. Evert
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Responses to Gravity
Four general steps lead to a gravitropic response:
1. Gravity is perceived by the cell
2. A mechanical signal is transduced into a physiological
signal
3. Physiological signal is transduced inside the cell and to
other cells
4. Differential cell elongation occurs in the “up” and “down”
sides of root and shoot
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Root and Shoot Responses to Gravity
In shoots, gravity is sensed along the length of the stem in
endodermal cells surrounding the vascular tissue
• Signaling occurs toward the outer epidermal cells
In roots, the cap is the site of gravity perception
• Signaling triggers differential cell elongation and division in
the elongation zone
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Response to Gravity in Shoots
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Response to Gravity in Roots
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Stem Response to Gravity
Auxin accumulates on lower side of the stem
• Results curvature of the stem upward.
Two Arabidopsis mutants, scarecrow (scr) and short root
(shr) do not show a normal gravitropic response
• Due to lack of a functional endodermis and its gravity-
sensing amyloplasts.
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scr and shr Mutants
©Jee Jung and Philip Benfey
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Root Response to Gravity
Gravity-sensing cells are located in the root cap
Cells that actually undergo asymmetrical growth are in the
distal elongation zone (closest to root cap)
Auxin may be involved, but exact mechanism is still unknown
• A gravitropic response still occurs is the distal elongation
zone if auxin transport is suppressed.
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Responses to Touch
• Thigmomorphogenesis: permanent form change in
response to mechanical stresses
• Thigmotropism: directional growth of a plant or plant part
in response to contact
• Thigmonastic responses: independent of the direction of
the stimulus and usually produced by changes in turgor
pressure
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Thigmotropism
Ed Reschke/Getty Images
The thigmotropic responses of these twining stems causes
them to coil around the object with which they have come
into contact.
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Changes in Turgor Pressure
• Some touch-induced plant movements involve reversible
changes in turgor pressure
• If water leaves turgid cells, they may collapse, causing
plant movement
• If water enters a limp cell, it becomes turgid and may also
move
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Mimosa pudica
The sensitive plant (Mimosa pudica) has compound leaves
with swollen structures called pulvini at the base of each
leaflet
• When leaves are stimulated by touch, wind, heat, or
possibly strong light, an electrical signal is generated.
• The electrical signal is translated into a chemical signal,
with potassium ions being pumped from the cells in one
half of a pulvinus to the intercellular spaces in the other
half
• Water follows by osmosis.
• Decreased interior turgor pressure causes the leaf to fold.
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Thigmonastic Response
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Tracking the Sun
• Some turgor movements are triggered by light
• Leaves may track the sun, orienting their blades at right
angle to the light
• This movement maximizes photosynthesis
Ronda Kimbrow/iStock/Getty Images
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Sleep Movement in Bean Leaves
During day, pulvini are turgid and leaves are horizontal.
• Maximizes photosynthetic surface area.
At night, the pulvini lose turgor and leaves become more vertical.
• Reduces water loss from transpiration.
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Dormancy
Dormancy results in the cessation of growth during
unfavorable conditions
• Deciduous trees: buds dormant, apical meristems are
protected inside enfolding scales.
• Perennial herbs: spend winter underground
• Annuals: winter as seeds.
• Often begins with abscission – dropping of leaves.
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Abscission
Abscission involves changes that occur in an abscission
zone at the petiole’s base
Hormonal changes lead to differentiation
• Protective layer – consists of several layers of suberin-
impregnated cells on stem side.
• Separation layer – consists of 1 to 2 layers of swollen,
gelatinous cells on leaf side.
• Pectins in the middle lamellae will break down
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Leaf Abscission
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Color Change During Leaf Abscission
Fuse/Getty Images
As the abscission zone develops, the green chlorophyll present in the
leaf breaks down, revealing the yellows and oranges of other pigments,
such as carotenoids, that previously had been masked. At the same time,
red and blue pigments called anthocyanins and betacyanins may
accumulate, all contributing to an array of fall colors in leaves.
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Seed Dormancy
Seeds allow plant offspring to wait until conditions for
germination are optimal
Triggers to break seed dormancy
• Increased levels of abscisic acid (ABA).
Favorable temperatures, day length, and amounts of water
can release buds, underground stems and roots, and seeds
from a dormant state
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Establishing Seed Dormancy
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Responses to Chilling
Lipid composition of a plant’s membranes can help predict
whether the plant will be sensitive or resistant to chilling
• The more unsaturated the membrane lipids are, the more
resistant the plant is to chilling.
• When chilling occurs, the enzyme desaturase changes
saturated lipids into unsaturated ones.
• At freezing ice crystals form and the cell dies from
dehydration
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Supercooling – some plants have the ability to survive at
temperatures as low as −40°C
• Limits ice crystal formation to extracellular spaces.
Antifreeze proteins prevent ice crystals from forming
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Responses to High Temperatures
Plants produce heat shock proteins (HSPs) if exposed to
rapid temperature increases
• HSPs stabilize other proteins.
Plants can survive otherwise lethal temperatures if they are
gradually exposed to increasing temperature
• Acquired thermotolerance.
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Hormones and Sensory Systems
Hormones are chemicals produced in one part of an
organism and transported to another part where they exert a
response
Generally produced in small, often minute quantities
In plants, hormones are not produced by specialized tissues
Eight major kinds of plant hormones
• Auxin, cytokinins, gibberellins, strigolactones,
brassinosteroids, oligosaccharins, ethylene, and abscisic
acid.
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Auxin
Effects of auxin discovered in 1881 by Charles and Francis
Darwin
• They reported experiments on the response of growing
plants to light.
• Grass seedlings do not bend if the tip is covered with a lightproof
cap.
• They do bend when a collar is placed below the tip.
Thirty years later, Peter Boysen-Jensen and Arpad Paal
demonstrated that a chemical causes the shoots to bend
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Shoot Tips Perceive Light
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Frits Went
• In 1926, Frits Went performed an experiment that
explained all of the previous results
• The chemical signal, which he named auxin, accumulated
on the side of an oat seedling away from light
• Auxin promoted these cells to grow faster than those on
the lighted side
• Cell elongation causes the plant to bend towards light
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Frits Went’s Experiment
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Auxin Causes Cell Elongation
Plant cells that are in the shade have more auxin and grow
faster than cells on the lighted side, causing the plant to bend
toward light.
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Winslow Briggs
• Winslow Briggs later demonstrated that auxin molecules
migrate away from the light into the shaded portion of the
shoot
• Barriers inserted in a shoot tip revealed equal amounts of
auxin in both the light and dark sides of the barrier
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Winslow Briggs Experiment
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Auxins
• Indoleacetic acid (IAA) is the most common natural auxin
• Probably synthesized from tryptophan
• The synthetic auxin 2,4-D is a widely used herbicide
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Mediating Auxin
Two families of proteins mediate auxin-induced changes in
gene expression
• Auxin response factors (ARFs).
• Bind to DNA and can enhance or suppress transcription
• Aux/IAA proteins.
• Bind and repress proteins that activate the expression of ARF
genes.
• ARF genes are activated when Aux/IAA proteins are degraded
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TIR1 is the auxin receptor
• Auxin binds directly to a protein called the transport
inhibitor response protein 1 (TIR1)
• It is part of the SCF protein complex.
• Auxin binds to TIR1 if Aux/IAA proteins are present
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Steps from Auxin Perception to Auxin-Induced Gene
Expression
1. Auxin binds TIR1 in the SCF complex
2. Activated SCF complex tags Aux/IAA proteins with
ubiquitin
3. Aux/IAA proteins are degraded in the proteasome
4. Aux/IAA proteins are no longer available to bind and
repress ARF transcriptional activators
5. ARF transcription factors faciliate transcription of auxin-
response-genes
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How Auxins Work
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Auxin Perception Sites
• Unlike with animal hormones, a specific signal is not sent
to specific cells, eliciting a predictable response
• Most likely, multiple auxin perception sites are present
• Auxin is also unique among the plant hormones in that it is
transported toward the base of the plant
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Auxin and Plant Cell Walls
One of the direct effects of auxin is an increase in the
plasticity of the plant cell wall
• Works only on young cell walls lacking extensive
secondary cell wall formation
Acid growth hypothesis
• Cells actively transport hydrogen ions from the cytoplasm
into the cell wall space
• The resulting drop in pH activates enzymes that can break
the bonds between cell wall fibers
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Acid Growth Hypothesis
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Synthetic Auxins
Naphthalene acetic acid (NAA) and indolebutyric acid (IBA)
have many uses in agriculture and horticulture
• Prevent fruit drop in apples and berries.
• Promote flowering and fruiting in pineapples.
• 2,4-dichlorophenoxyacetic acid (2,4-D) is a herbicide
commonly used to kill weeds.
• Applied in high concentrations, ceases all axial growth in broad-
leaves dicots.
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Cytokinins
• Plant hormone that, in combination with auxin, stimulates cell
division and differentiation
• Appear to be derivatives of adenine
• Common synthetic cytokinins include kinetin and 6-benzylamino
purine.
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Role of Cytokinins
Produced in the root apical meristems and developing fruits
Transported throughout plant
In all plants, cytokinins, working with other hormones, seem
to regulate growth patterns
Promote the growth of lateral buds into branches
Inhibit the formation of lateral roots
• Auxin promotes their formation.
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Hormones and Lateral Buds
©Prof. Malcolm B. Wilkins, Botany Dept, Glasgow University
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Cytokinins and Auxins
Plant tissue can form shoots, roots, or an undifferentiated mass
depending on the relative amounts of auxin and cytokinin
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Other Effects of Cytokinins
Promote the synthesis or activation
of cytokinesis proteins
Also function as antiaging
hormones
Agrobacterium inserts genes that
increase rate of cytokinin and auxin
production
• Causes massive cell division.
• Formation of crown gall tumor.
Custom Life Science Images/Alamy Stock Photo
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Strigolactones
Derived from carotenoids
Move upward from the roots
Inhibit axillary bud growth
• Affect extent of branching
Regulate vascular cambium in coordination with auxin
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Gibberellins
Named after the fungus Gibberella fujikuroi which causes
rice plants to grow very tall
Gibberellins belong to a large class of over 100 naturally
occurring plant hormones
• All are acidic and abbreviated GA (for gibberellic acid).
• Have important effects on stem elongation.
• Enhanced if auxin present.
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Gibberellin Deficiencies in Dwarf Plants
Omikron/Science Source
The mutant pea plant on the left is defective in gibberellin biosynthesis,
and thus has a shorter stem. This type of mutant can be rescued by
applying gibberellins to the shoot tip, allowing it to grow to normal height.
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Gibberellins and Grapes
Hasten seed germination
Used commercially to extend internode length in grapes
• Result is larger grapes.
©Amnon Lichter, The Volcani Center
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Gibberellin as a Signal
• GA is used as a signal from the embryo that turns on
transcription of genes encoding hydrolytic enzymes in the
aleurone layer
• When GA binds to its receptor, it frees GA-dependent
transcription factors from a repressor
• These transcription factors can now directly affect gene
expression
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Brassinosteroids
• First discovered in the pollen of Brassica spp.
• Are structurally similar to steroid hormones
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Brassinosteroid Characteristics
Functional overlap with other plant hormones, especially
auxins and gibberellins
Broad spectrum of physiological effects
• Elongation, cell division, stem bending, vascular tissue
development, delayed senescence, membrane
polarization, and reproductive development.
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Oligosaccharins
• Are complex plant cell wall carbohydrates that have a
hormone-like function
• Can be released from the cell wall by enzymes secreted
by pathogens
• Signal defense responses, such as the hypersensitive
response (HR)
• In peas, oligosaccharins inhibit auxin-stimulated
elongation of stems
• While in regenerated tobacco tissue, they inhibit roots and
stimulate flowers
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Ethylene
Gaseous hydrocarbon (H2C − CH2)
Auxin stimulates ethylene production in the tissues around
the lateral bud and thus restricts their growth
Ethylene also suppresses stem and root elongation
Major role in fruit development – hastens ripening
• Transgenic tomato plant can’t make ethylene.
• Shipped without ripening and rotting.
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Regulation of Fruit Ripening
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Abscisic Acid
• Synthesized mainly in mature green leaves, fruits, and root
caps
• Little evidence that this hormone plays a role in abscission
• Induces formation of dormant winter buds
• Counteracts gibberellins by suppressing bud growth and
elongation
• Counteracts auxin by promoting senescence
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Effects of Abscisic Acid
Necessary for dormancy in seeds
• Prevents precocious germination called vivipary.
Important in the opening and closing of stomata
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Effects of Abscisic Acid on Winter Buds
(a) Evelyn Jo Johnson/ McGraw Hill
ABA plays a role in the formation of these winter buds of an
American basswood. These buds will remain dormant for the
winter and the bud scales – modified leaves – will protect the buds
from dessication
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Effects of Abscisic Acid on Bud Seeds
(b) ©From D. R. McCarty, C. B. Carson, P. S. Stinard, and D. S. Robertson, “Molecular analysis of viviparous-1: an abscisic acid-insensitive mutant of maize,”
The Plant Cell, 1 (5): 523–532 ©1989 American Society of Plant Biologists
ABA is necessary for the dormancy of seeds. This viviparous
mutant in maize is deficient in ABS, and as a result the
embryos begin germinating on the developing cob.
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Effects of Abscisic Acid on Stomata
(c) Ed Reschke/Getty Images
Abscisic acid also affects the closing of stomata like this one
by influencing the movement of potassium ions out of guard
cells.
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