07-Mar-21

Chapter 39

Plant Signals
and Behavior

Lecture Presentations by
Nicole Tunbridge and
© 2018 Pearson Education Ltd.
Kathleen Fitzpatrick

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

© 2018 Pearson Education Ltd.

1
 07-Mar-21

Figure 39.1

© 2018 Pearson Education Ltd.

Figure 39.1a

© 2018 Pearson Education Ltd.

2
 07-Mar-21

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

© 2018 Pearson Education Ltd.

Figure 39.2a

(a) Before exposure to light

© 2018 Pearson Education Ltd.

3
 07-Mar-21

Figure 39.2b

(b) After a week’s exposure to

natural daylight

© 2018 Pearson Education Ltd.

Figure 39.2

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

to natural daylight

© 2018 Pearson Education Ltd.

4
 07-Mar-21

 A potato’s response to light is an example of cell

signal processing

 The stages are reception, transduction, and

response

© 2018 Pearson Education Ltd.

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

© 2018 Pearson Education Ltd.

5
 07-Mar-21

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

© 2018 Pearson Education Ltd.

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
© 2018 Pearson Education Ltd.

6
 07-Mar-21

Figure 39.4_1

1 Reception

CYTOPLASM
Plasma
membrane

Phytochrome
activated
by light
Cell
wall

Light

Ca2+

© 2018 Pearson Education Ltd.

Figure 39.4_2

1 Reception 2 Transduction

CYTOPLASM
Plasma cGMP Protein
membrane kinase 1
Second
Phytochrome messenger
activated produced
by light
Cell
wall Protein
kinase 2

Light

Ca2+ channel
opened

Ca2+

© 2018 Pearson Education Ltd.

7
 07-Mar-21

Figure 39.4_3

1 Reception 2 Transduction 3 Response

Transcription
CYTOPLASM factor 1 NUCLEUS
Plasma cGMP Protein P
membrane kinase 1
Second
messenger Transcription
Phytochrome
produced factor 2
activated
by light P
Cell
wall Protein
kinase 2
Transcription
Light
Translation

Ca2+ channel De-etiolation

opened (greening)
response
proteins
Ca2+

© 2018 Pearson Education Ltd.

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

© 2018 Pearson Education Ltd.

8
 07-Mar-21

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

© 2018 Pearson Education Ltd.

 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

© 2018 Pearson Education Ltd.

9
 07-Mar-21

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

© 2018 Pearson Education Ltd.

De-etiolation (“Greening”) Proteins

 De-etiolation activates enzymes that

 Function in photosynthesis directly

 Supply the chemical precursors for chlorophyll

production

 Affect the levels of plant hormones that regulate

growth

© 2018 Pearson Education Ltd.

10
 07-Mar-21

Concept 39.2: Plant hormones help coordinate

growth, development, and responses to stimuli
 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

© 2018 Pearson Education Ltd.

 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
© 2018 Pearson Education Ltd.

11
 07-Mar-21

A Survey of Plant Hormones

 The major plant hormones include

 Auxin
 Cytokinins
 Gibberellins
 Abscisic acid
 Ethylene
 Brassinosteroids
 Jasmonates
 Strigolactones

© 2018 Pearson Education Ltd.

Auxin

 Any response resulting in curvature of organs toward

or away from a stimulus is called a tropism (tropos:
turn)

 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

© 2018 Pearson Education Ltd.

12
 07-Mar-21

 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

© 2018 Pearson Education Ltd.

Figure 39.5a

Control
Shaded
side

Light

Illuminated
side

© 2018 Pearson Education Ltd.

13
 07-Mar-21

Figure 39.5b

Darwin and Darwin: Phototropism occurs only

when the tip is illuminated.

Light

Tip Opaque Trans- Opaque

removed cap parent shield over
cap curvature

© 2018 Pearson Education Ltd.

Figure 39.5c

Boysen-Jensen: Phototropism
occurs when the tip is separated by
a permeable barrier but not an
impermeable barrier.

Light

Gelatin Mica
(permeable) (impermeable)

© 2018 Pearson Education Ltd.

14
 07-Mar-21

Figure 39.5
Results
Control
Shaded
side

Light

Illuminated
side

Darwin and Darwin Boysen-Jensen

Light Light

Tip Opaque Trans- Opaque Gelatin Mica

removed cap parent shield over (permeable) (impermeable)
cap curvature
Data from C. R. Darwin, The power of movement in plants, John Murray, London (1880). P. Boysen-Jensen,
Concerning the performance of phototropic stimuli on the Avenacoleoptile, Berichte der Deutschen
Botanischen Gesellschaft (Reports of the German Botanical Society) 31:559–566 (1913).
© 2018 Pearson Education Ltd.

 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

© 2018 Pearson Education Ltd.

15
 07-Mar-21

 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

© 2018 Pearson Education Ltd.

Figure 39.6

Results
Cell 1

100 µm Cell 2

Epidermis

Cortex

Phloem

Xylem 25 µm
Basal end
Pith of cell

Data from L. Gälweiler et al., Regulation of polar auxin transport by AtPIN1 in

Arabidopsis vascular tissue, Science 282:2226–2230 (1998).

© 2018 Pearson Education Ltd.

16
 07-Mar-21

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

© 2018 Pearson Education Ltd.

 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

© 2018 Pearson Education Ltd.

17
 07-Mar-21

Figure 39.7

Cell wall before auxin activates Loosening of cell wall, enabling elongation
proton pumps
Cellulose 1 Auxin increases 2 Low pH activates
microfibrils activity of proton expansins (red).
pumps.

Elongation
PLANT CELL H+ H+ H+
H+
Nucleus Vacuoles WALL H+

H+
H+ H+

H+
H+

H+

H+
H+

Cellulose microfibril Cross-linking 3 Polysaccharides are cleaved by cell wall-loosening

polysaccharide enzymes (purple), loosening the microfibrils (brown).

© 2018 Pearson Education Ltd.

 Auxin also alters gene expression and stimulates a

sustained growth response

© 2018 Pearson Education Ltd.

18
 07-Mar-21

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

© 2018 Pearson Education Ltd.

 The activity of the vascular cambium is under control

of auxin transport

 Organization of female angiosperm gametophytes is

likely regulated by an auxin gradient

© 2018 Pearson Education Ltd.

19
 07-Mar-21

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

© 2018 Pearson Education Ltd.

 Developing seeds produce auxin, which promotes

fruit development

 Greenhouse tomatoes produce few seeds

 Spraying synthetic auxins on greenhouse tomatoes

improves fruit development

© 2018 Pearson Education Ltd.

20
 07-Mar-21

Cytokinins

 Cytokinins are so named because they stimulate

cytokinesis (cell division)

© 2018 Pearson Education Ltd.

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

© 2018 Pearson Education Ltd.

21
 07-Mar-21

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

 Removal of the apical bud increases sugar

availability and decreases auxin and strigolactone
levels, initiating axillary bud growth
© 2018 Pearson Education Ltd.

Figure 39.8
Topmost axillary buds grow
and take over as the new
apical bud.

Apical bud

Limited growth
of axillary buds
Plant with apical Plant with apical
bud intact bud removed
© 2018 Pearson Education Ltd.

22
 07-Mar-21

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

© 2018 Pearson Education Ltd.

Gibberellins

 Gibberellins have a variety of effects, such as stem

elongation, fruit growth, and seed germination

© 2018 Pearson Education Ltd.

23
 07-Mar-21

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

© 2018 Pearson Education Ltd.

Figure 39.9a

(a) Rosette form (left) and

gibberellin-induced bolting (right)
© 2018 Pearson Education Ltd.

24
 07-Mar-21

Fruit Growth
 In many plants, both auxin and gibberellins must be
present for fruit to develop

 Gibberellins are used in spraying of Thompson

seedless grapes

© 2018 Pearson Education Ltd.

Figure 39.9b

(b) Grapes from control vine

(left) and gibberellin-treated
vine (right)

© 2018 Pearson Education Ltd.

25
 07-Mar-21

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

© 2018 Pearson Education Ltd.

Figure 39.10

Aleurone
Endosperm 1 2 3

α-amylase Sugar
GA

GA
Water

Scutellum Radicle
(cotyledon)

© 2018 Pearson Education Ltd.

26
 07-Mar-21

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

© 2018 Pearson Education Ltd.

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

© 2018 Pearson Education Ltd.

27
 07-Mar-21

Figure 39.11
Red mangrove
(Rhizophora mangle)
seeds

Coleoptile

Maize mutant

© 2018 Pearson Education Ltd.

Figure 39.11a
Red mangrove
(Rhizophora mangle)
seeds

© 2018 Pearson Education Ltd.

28
 07-Mar-21

Figure 39.11b

Coleoptile

Maize mutant

© 2018 Pearson Education Ltd.

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”

© 2018 Pearson Education Ltd.

29
 07-Mar-21

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

© 2018 Pearson Education Ltd.

The Triple Response to Mechanical Stress

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

© 2018 Pearson Education Ltd.

30
 07-Mar-21

Figure 39.12

0.00 0.10 0.20 0.40 0.80

Ethylene concentration (parts per million)

© 2018 Pearson Education Ltd.

 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

© 2018 Pearson Education Ltd.

31
 07-Mar-21

Figure 39.13

ein mutant
ctr mutant

(a) ein mutant (b) ctr mutant

© 2018 Pearson Education Ltd.

Figure 39.13a

ein mutant

(a) ein mutant

© 2018 Pearson Education Ltd.

32
 07-Mar-21

Figure 39.13b

ctr mutant

(b) ctr mutant

© 2018 Pearson Education Ltd.

Senescence
 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

© 2018 Pearson Education Ltd.

33
 07-Mar-21

Leaf Abscission
 A change in the balance of auxin and ethylene
controls leaf abscission, the process that occurs in
autumn when a leaf falls

© 2018 Pearson Education Ltd.

Figure 39.14

0.5 mm

Protective layer Abscission layer

Stem Petiole

© 2018 Pearson Education Ltd.

34
 07-Mar-21

Figure 39.14a

0.5 mm

Protective layer Abscission layer

Stem Petiole

© 2018 Pearson Education Ltd.

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

© 2018 Pearson Education Ltd.

35
 07-Mar-21

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

© 2018 Pearson Education Ltd.

 Jasmonates, including jasmonate (JA) and methyl

jasmonate (MeJA) play important roles in plant
defense and development
 They are produced in response to wounding and are
involved in controlling plant defenses

© 2018 Pearson Education Ltd.

36
 07-Mar-21

 Jasmonates also regulate many other physiological

processes, including
 Nectar secretion
 Fruit ripening
 Pollen production
 Flowering time
 Seed germination
 Root growth
 Tuber formation
 Mycorrhizal symbiosis
 Tendril coiling
© 2018 Pearson Education Ltd.

 Strigolactones are xylem-mobile chemicals that

 Stimulate seed germination
 Suppress adventitious root formation
 Help establish mycorrhizal associations
 Help control apical dominance
 Strigolactones are named for parasitic Striga plants
 Striga seeds germinate when host plants exude
strigolactones through their roots

© 2018 Pearson Education Ltd.

37