Plant Responses to

Internal & External

Signals39
Chapter
The Necessity of Plant Responses

• For appropriate growth

and development
Etiolation De-etiolation
• For survival and
reproduction
• For protection
• For adjustment in
accordance with the
change in time
• ….
Stimulus → → →
response: Signal
transduction
pathways are the
link

Reception: Internal and external signals are detected by receptors

(proteins that change in response to specific stimuli)

Transduction: Second messengers transfer and amplify signals from

receptors to proteins that cause specific responses

Response: Results in regulation of one or more cellular activities. In

many cases this involves the increased activity of certain enzymes
 An example of signal transduction in plants: the
role of phytochrome in the de-etiolation (greening)
response.

Phytochrome:
photoreceptor
locating in cytoplasm

Require both
pathways for
complete de-
etiolation
↑ 100-fold

- Chlorphyll biosynthesis
- Photosynthetic enzymes
- Phytohormone content (↓ auxin)
 Plant hormones help coordinate growth,
development and responses to stimuli

(phototropism)

(coconut milk,
maize zeatin)

(fungus of the
genus Gibberella
causes
this “foolish
seedling
disease.”)
(growth inhibitor):
the ratio of ABA
to one or more
growth hormones
determines the
final physiological
outcome

The levels of
ABA may increase
100-fold during seed
maturation, inhibiting
germination
and inducing the
Often, the ratio of production of proteins
ABA to GAs that help the seeds
determines withstand the extreme
whether seeds dehydration that
remain dormant or accompanies
germinate maturation.

Precocious germination = early germination

 (gas)
(coal gas
leakage and
premature leaf
abscission)

(brassinosteroid-
deficient mutant)

(biotic
defense)
(Jasmonate (JA) & methyl
jasmonate (MeJA))

Auxin, CK, GA, ABA and ET are

considered the five “classic” plant
hormones
• Plant hormones are produced in very low
concentrations, but a tiny amount of hormone
can have a profound effect on plant growth and
development.
• By affecting the division, elongation, and
differentiation of cells
• Various natural biologically active forms of the
same class of phytohormones have been
identified
• Each hormone has multiple effects, depending
on its site of action, its concentration, and the
developmental stage of the plant.
• Conversely, multiple hormones can influence a
single process.
• Plant hormone responses commonly depend
on both the amounts of the hormones involved
and their relative concentrations.

The regulation is complex due to overlapping

functions and cross-talks among hormonal
pathways (synergistic or antagonistic actions)
 Many plant biologists prefer the broader term plant
growth regulator to describe organic compounds,
either natural or synthetic, that modify or control one or
more specific physiological processes within a plant
FEW EXAMPLES
 Tropisms
– Any growth response that results in plant
organ curving toward or away from a stimulus
is called a tropism
– Often caused by hormones
Phototropism: light is the stimulus

What part of a grass coleoptile senses light, and how is the signal transmitted?

Positive
phototropism
Phototropism: light is the stimulus

1880

Bending: x x √ √
- The grass seedling ensheathed in its coleoptile could bend toward
light only if the tip of the coleoptile was present and exposed to light.
- The phototropic bending, however, occurred at a distance from the
site of light perception (the tip)
Phototropism: light is the stimulus

1913

Peter Boysen-Jensen demonstrated that the signal was a mobile

chemical substance, which is later known as auxin (indoleacetic
acid – IAA).
• In natural ecosystems, where plants may be crowded,
phototropism directs shoot growth toward the sunlight that
powers photosynthesis.

• This response results from a differential growth of cells on

opposite sides of the shoot; the cells on the darker side elongate
faster than the cells on the brighter side.

Coleoptiles curve toward light

because of a higher concentration
of auxin on the darker side of the
coleoptile.
 Frits Went’s experiment in 1926 to find out how
auxin causes a coleoptile to grow toward light
The coleoptile grew straight if the
chemical was distributed evenly.
EXPERIMENT
RESULTS If the chemical was distributed
unevenly, the coleoptile curved away
He placed coleoptiles in the dark from the side with the block, as if
and removed their tips, putting growing toward light, even though it
some tips on agar blocks that he was grown in the dark.
predicted would absorb the
chemical.
Excised tip placed
on agar block

Growth-promoting
On a control coleoptile, he placed chemical diffuses
into agar block
a block that lacked the chemical.

On others, he placed blocks Control Agar block

(agar block with chemical
containing the chemical, either lacking stimulates growth

centered on top of the coleoptile chemical)

has no
Offset blocks
to distribute the chemical evenly Control effect
cause curvature

or offset to increase the

concentration on one side.
 Expansins are enzymes that break
Cell elongation in response to auxin: the cross-links (hydrogen bonds)
between cellulose microfibrils and
the acid growth hypothesis other cell wall constituents,
loosening the wall’s fabric.

Auxin stimulates growth only over a certain concentration range, from about 10-8 to 10-4 M.

At higher concentrations, auxin may inhibit cell elongation, by inducing production of

ethylene, a hormone that generally hinders growth.
 Apical dominance

• The ability of a terminal bud to suppress

development of axillary buds

• CKs, auxin, strigolactones interact in the

control of apical dominance
Control of Apical Dominance

Bud dormancy = paradomancy

Auxin added to
decapitated stem

“Stump” after
removal of
Axillary buds apical bud Applying a gelatin capsule
containing auxin to the
Lateral branches
stump prevents the lateral
branches from growing
If the terminal bud is removed
plants become bushier
Control of Apical Dominance- Underlying mechanisms

1. Direct inhibition hypothesis: auxin and CKs act

antagonistically in regulating axillary bud growth
- auxin transported down the shoot from the apical bud
directly inhibits axillary buds from growing, causing a
shoot to lengthen.
- Meanwhile, CKs entering the shoot system from roots
counter the action of auxin by signaling axillary buds to
begin growing.
- Thus, the ratio of auxin and CKs was viewed as the
critical factor in controlling axillary bud inhibition.
Control of Apical Dominance- Underlying mechanisms

2. Indirect inhibition hypothesis: auxin’s effects

are partially indirect (more updated)
- The polar flow of auxin down the shoot triggers the
synthesis of strigolactones, which repress bud growth.
- Moreover, another signal, perhaps an electrical one,
appears to cause buds to begin growing much earlier
than can be explained by disrupted auxin flow.
Control of Apical Dominance- Underlying mechanisms

3. The hypothesis is further revised: Recent

discoveries indicate bud dormancy in response to
intrinsic and environmental factors is linked to
enhanced growth of the main shoot and reduced
sugar level in the buds

- by diverting sugars away from buds (i.e. a growing stem

is a strong sink for sugars .
Roles of Gibberellin on stem elongation,
fruit growth and seed germination

GA are best known for

stimulating stem and leaf
growth by enhancing cell
elongation and cell division.

One hypothesis proposes that

they activate enzymes that
loosen cell walls, facilitating
entry of expansin proteins.
Thus, GAs act in concert with
auxin to promote stem
elongation
Roles of Gibberellin on stem elongation,
fruit growth and seed germination
In many plants, both auxin
and gibberellins must be
present for fruit to develop.

GA makes the individual

grapes grow larger, the
internodes of the grape bunch
elongate, allowing more space
for the individual grapes.

By enhancing air circulation

between the grapes, this
increase in space also makes it
harder for yeasts and other
microorganisms to infect the
fruit.
Roles of Gibberellin on stem elongation, fruit growth
and seed germination
Roles of ABA in
drought tolerance

By affecting second messengers (e.g. Ca2+), ABA

causes K+ channels in the plasma membrane of
guard cells to open
→ leading to a massive loss of K+ ions.
→ osmotic loss of water reduces guard cell turgor
→ leads to closing of the stomatal pores

https://doi.org/10.3390/plants10122774
Roles of ABA in
drought tolerance

https://doi.org/10.1016/bs.abr.2019.05.001
 Ethylene effects in responses to mechanical stress

• The Triple Response:

EXPERIMENT Germinating pea seedlings were placed in the
1. Stems elongate less dark and exposed to varying ethylene concentrations. Their growth
was compared with a control seedling not treated with ethylene.
rapidly RESULTS All the treated seedlings exhibited the triple
response. Response was greater with increased concentration.
2. Stems thicken (make it
stronger)
3. Stems grow horizontally

→ allows a growing shoot to

avoid obstacles during soil
0.00 0.10 0.20 0.40
penetration 0.80

Ethylene concentration (parts per million)

As the effects of the initial ET pulse lessen,

the stem resumes vertical growth. If it again
contacts a barrier, another burst of ET is
released, and horizontal growth resumes.
Other Ethylene effects
(along with enzymatic breakdown)

• Apoptosis (programmed cell death): a burst of

ethylene is associated with the programmed
destruction of cells, organs, or whole plants

• Fruit Ripening: a burst of ET production

triggers the ripening process
→ Postharvest management concern
Other Ethylene effects
(along with enzymatic breakdown)
• Leaf Abscission (the loss of leaves, usually in
autumn): a change in the balance of ET and
auxin controls leaf abscission
– Before dying leaves abscise, many essential elements are
salvaged from them and stored in stem parenchyma cells.
– These nutrients are recycled back to developing leaves the
following spring

Even before the leaf falls, a

layer of cork forms a
protective scar on the twig
side of the abscission layer,
preventing pathogens from
invading the plant.
Responses to light are
critical
for plant success
Plant Responses to Light

• Light cues many key events in plant growth

and development.
• Light reception is important for measuring the
passage of days and seasons
• Effects of light on plant morphology is called
photomorphogenesis
• Plants not only detect the presence of light but
also its direction, intensity, and wavelength
(color)
 Action Spectra for Phototropic response
EXPERIMENT Researchers exposed maize (Zea mays) Action spectra are
coleoptiles to violet, blue, green, yellow,
orange, and red light to test which useful in studying any
wavelengths stimulate the phototropic process that depends
bending toward light.
on light

This action spectrum illustrates that only

light wavelengths below 500 nm (blue and
violet light) induce curvature.

CONCLUSION
The phototropic bending toward light is caused
by a photoreceptor that is sensitive to blue and violet
light, particularly blue light.
Photoreceptors (two major classes)

1. Blue-light receptors
– Control hypocotyl elongation, stomatal opening, and
phototropism
– Include phototropins (regulate phototropism) and
cryptochromes (regulate stem elongation, flowering)

2. Phytochromes (red/far-red light receptors)

– Regulate many of a plant’s responses to light
throughout its life (such as seed germination and
shade avoidance)
doi: 10.1105/tpc.000646
Phytochromes & Seed
Germination

Red light stimulated

germination, and far-
red light inhibited
germination.

The final exposure

was the determining
factor. The effects of
red and far-red light
The germination rate
were reversible. increased greatly in
Germination was
inhibited in groups of
groups of seeds that were
seeds that were last
last exposed
exposed to far-red light.
to red light.
 Red light
Synthesis

Phytochrome Far-red
switch Pr light Pfr
Slow conversion
Enzymatic
in darkness
destruction
(some plants)

• Phytochromes exist in 2 photoreversible states (isomers:)

Pr (red-absorbing form) and Pfr (far-red- absorbing form)
• Plants synthesize Pr, and if seeds are kept in the dark, the
pigment remains almost in the Pr form.
• Sunlight contains both red light and far-red light, but the
conversion to Pfr is faster than the conversion to Pr.
• When seeds are exposed to adequate sunlight, the
production and accumulation of Pfr triggers germination
 Phytochromes and Shade Avoidance

• Phytochrome also provides the

plant with information about the
light quality

• In the “shade avoidance”

response of a tree
– The phytochrome ratio shifts in
favor of Pr when a tree is
shaded. (Pr > Pfr)
→ The tree to allocate more
resources to grow taller Chlorophyll in leaves
and less branch of the canopy absorb
red light and allow
DOI:10.3389/fpls.2017.01181
DOI:10.1104/pp.122.3.621
far-red light to pass
Biological Clocks and Circadian Rhythms

• Many plant processes undergo a daily

oscillation.
• Examples: transpiration, the synthesis of
certain enzymes
• Some of these cyclic variations are responses
to the changes in light levels and temperature
that accompany the 24-hour cycle of day and
night
• Biological clock: An internal We can interfere
timekeeper that controls an with a biological
organism’s biological rhythms. The rhythm, but the
biological clock marks time with or underlying
without environmental cues but clockwork continues
often requires signals from the to tick!
environment to remain tuned to an
appropriate period.

Shut off from

jet lag
environmental
cues, the
plant becomes
desynchronized.
• Circadian rhythm: A
physiological cycle of about 24 Molecular
hours that persists even in the mechanisms
absence of external cues. underlying
circadian rhythms
are oscillations in
the transcription of
certain genes.
Biological Clocks and Circadian Rhythms
– For example, many legumes lower their leaves
in the evening and raise them in the morning
(these are called sleep movements)
The Effect of Light on the Biological Clock

• The factor that entrains the biological clock to

precisely 24 hours every day is light.
• Both phytochromes and blue-light
photoreceptors can entrain circadian rhythms
in plants
The Effect of Light on the Biological Clock

• Phytochrome conversion marks sunrise and

sunset providing the biological clock with
environmental cues
An increase of red light during the day causes Pfr to accumulate,
while the amount of Pr accumulates in dim light or darkness

→ Interactions between phytochrome and the biological

clock enable plants to measure the passage of night
and day, thus the change over the course of the year
→ Adjust plant activities in synchrony with the seasons
• Photoperiod (the relative lengths of night and
day)
- the interval in a 24-hour period during which
an organism is exposed to light
- is the environmental stimulus plants use
most often to detect the time of year

• Photoperiodism
– Is a physiological response to photoperiod
(e.g. flowering)
• Short-day plants (generally flower in late summer,
fall, or winter): require a light period shorter than a
critical length to flower
– Example: Chrysanthemums

• Long-day plants (flower in late spring or early

summer): require the light period longer than a
certain number of hours to flower
– Examples: spinach, lettuce

• Day-neutral plants: are unaffected by photoperiod

and flower at a certain stage of maturity
– Examples: tomato, rice, dandelion
 Critical Night Length
• In the 1940s, researchers discovered that flowering
and other responses to photoperiod
– Are actually controlled by night length, not day length

Short-day (long-night)
- “Short-day” plants are more plant. Flowers when
night exceeds a critical
properly called “long-night” dark period.
plants,

- and “long-day” plants are really

“short-night” plants. Long-day (short-night)
plant. Flowers only if the
night is shorter than a
critical dark period.
It appears that plants
use their biological
clock, entrained by
night length with the
help of phytochrome,
to tell the season of
the year.

A flash of red (R) light shortens the dark

period. A subsequent flash of far-red
(FR) light cancels the red flash’s effect.
Test for presence of a flowering hormone (florigen)

Although flowers
form from apical or
axillary bud
meristems,
it is leaves that
detect changes in
photoperiod and
produce signaling
molecules that cue
buds to develop as
flowers.

It now appears that florigen is a protein.

A gene called FLOWERING LOCUS T (FT) is activated in leaf cells
during conditions favoring flowering, and the FT protein travels through
the symplasm to the shoot apical meristem, initiating the transition of a
bud’s meristem from a vegetative to a flowering state.
Plant responses to
Non-Light stimuli
Gravity

• Response to gravity is gravitropism

• Occur as soon as the seed germinates
• Roots show positive gravitropism
• Stems show negative gravitropism
 In roots, statoliths are
located in certain cells of the root cap

Statoliths
20 m

(a) (b)
DOI:10.1104/pp.103.032169 The root cap comprises central columella cells and surrounding root cap cells

• Uitility of statoliths is a way for gravity detection

• Statoliths are dense cytoplasmic components (e.g. plastids
with dense starch grains) to lower portions of cells.
Working model of the dual role of auxin in mediating root
gravitropism. The red lines indicate the role of auxin in
mediating gravity perception by regulating starch granule
accumulation, and the black lines indicate the role of auxin
in mediating gravitropic response.

https://doi.org/10.1111/nph.15932
Response to Mechanical Stimuli
Rubbing the stems of
young plants a couple of
• Thigmomorphogenesis times daily results in plants
that are shorter than
refers to the changes in controls
form that result from
mechanical perturbation

Rubbed Un-rubbed
 Response to Mechanical Stimuli

• Thigmotropism refers
directional growth in
response to touch

• Growth in response to touch These grasping organs usually

grow straight until they touch
occurs in vines and other something; the contact
climbing plants. stimulates a coiling response
caused by differential growth of
cells on opposite sides of the
• Getting mechanical tendril.
supports
Responses to Environmental Stresses

• Environmental stresses
– Have a potentially adverse effect on a plant’s survival, growth,
and reproduction
– Can have a devastating impact on crop yields in agriculture
– Climate change enhances the threats
Responses
to
Drought
Stresses

– Reducing canopy size, transpiration, photosynthesis

– Increase root growth, antioxidations
doi.org/10.1007/978-981-10-9029-5_1
Responses to Flooding
• Waterlogged soil lacks air spaces to provide oxygen for
cellular respiration in roots.
• Oxygen deprivation stimulates ethylene production which
then leads too…Enzymatic destruction of cells and
creation of air tubes “snorkels” that provide oxygen to
submerged roots
Responses to other abiotic stresses

• Salt Stress…producing compatible solutes

(solutes tolerated at high concentrations) which
keeps the water potential of cells more negative
than that of the soil solution; ion
exclusion/storing, antioxidation

• Heat Stress… Heat-shock proteins used to

protect important molecules from denaturation

• Cold/freezing Stress…Altering lipid composition

of membranes to maintain fluidity of membranes;
increasing levels of solutes (like sugar) in the cells
helps dealing with dehydration problem
 Plant defense against attacks by
herbivores and pathogens
nematode
virus

bacteria

herbivore
 Plant Defensive Approaches against herbivores

• Physical barriers
– Thorns and trichomes

• Chemicals
– distasteful or toxic compounds (e.g. terpenoids,
phenolics, and alkaloids)
– Recruitment of predatory animals following being
wounded
Recruitment of Predatory animals

4 Recruitment of
parasitoid wasps
that lay their eggs
within caterpillars

3 Synthesis and
release of
volatile attractants

1 Wounding 1 Chemical
in saliva

2 Signal transduction
pathway
 Other defenses

The leaf glands mimic the

presence of insect eggs on
their leaves, dissuading
insects from laying eggs
there

Mimics
The leaf of the snowflake plant
looks as if it has been partially
eaten, making it less attractive
Other defenses

Changing flowering
times from night to
dawn to protect
tobacco plants
(Nicotiana
attenuata) against
herbivory of hawk-
moths

Flowering at night Flowering at

and hawk-moth dawn and
as pollinator hummingbird as
pollinator
Plant Defensive Approaches Against Pathogens
• Physical barriers
– epidermis and the periderm

• Chemicals
– Once a pathogen invades, the plant mounts a
chemical attack that kills the pathogen and
prevents its spread
– This defense system is enhanced by the
plant’s inherited ability to recognize certain
pathogens via pathogen-associated molecular
patterns (PAMPs) (i.e. elicitors)
Pathogens

• A virulent pathogen
– Is one that a plant has little specific defense
against

• An avirulent pathogen
– Is one that may harm but not kill the host plant
Gene-for-gene recognition

• Involves recognition of pathogen-produced

molecules (effectors) by the protein products
(receptors) of specific plant disease resistance
(R) genes
Avirulent pathogen
• A pathogen is avirulent if it has a specific Avr
gene corresponding to a particular R allele in
the host plant
Receptor coded by R allele
Signal molecule (ligand)
from Avr gene product

Avr allele

Avirulent pathogen
Plant cell is resistant

(a) If an Avr allele in the pathogen corresponds to an R allele

in the host plant, the host plant will have resistance,
making the pathogen avirulent. R alleles probably code for
Figure 39.30a receptors in the plasma membranes of host plant cells. Avr alleles
produce compounds that can act as ligands, binding to receptors
in host plant cells.
Virulent pathogen
• If the plant host lacks the R gene that
counteracts the pathogen’s Avr gene
– Then the pathogen can invade and kill the
plant
R

No Avr allele;
virulent pathogen
Plant cell becomes diseased

Avr allele

No R allele;
Virulent pathogen plant cell becomes diseased

Virulent pathogen

No R allele;
plant cell becomes diseased

(b) If there is no gene-for-gene recognition because of one of

the above three conditions, the pathogen will be virulent,
causing disease to develop.
Hypersensitive Response against avirulent pathogens

• Seals off the infected site: kills both

pathogen and host cells in the region of the
infection, prevent the spread

• Produce antimicrobial chemicals called

phytoalexins (fungicidal and bactericidal)
• Thicken plant cell wall (formation of lignin
and cross-linkings)
• Send the message to distant organs for
systematic acquired resistance
Systemic Acquired Resistance (SAR)

– Is a set of generalized defense responses in

organs distant from the original site of infection
– Is triggered by the signal molecule salicylic
acid (which activates plant defenses throughout the
plant before infection spreads)
Plant Response against avirulent pathogens
Each R protein can be activated by its specific effector
- END-