Plant Nutrition

https://www.youtube.com/watch?
v=MnY_cCRELvs
 Plant Responses
Chapter 39
to Internal and
External Signals
Response to stimuli
• Plants, being rooted to the ground must
respond to whatever environmental change
comes their way
• For example, the bending of a grass seedling
toward light begins with the plant sensing the
direction, quantity, and color of the light
Signal Transduction stimulus    response

• Signal transduction pathways link signal

reception to response
• Plants have cellular receptors to detect
important changes in their environment
• For a stimulus to elicit a response the cell must
have an appropriate receptor
• Upon receipt of the stimulus the receptor starts
a series of biochemical steps that lead to a
response
 Potato Example
• A potato left growing in darkness will produce shoots that
do not appear healthy, and lack elongated roots
• These are morphological adaptations for growing in
darkness are referred to as etiolation
• After the potato is exposed to light, the plant undergoes
changes called de-etiolation, (greening) in which shoots
and roots grow normally

Before exposure to light. A

dark-grown potato has tall,
spindly stems and nonexpanded
leaves—morphological
adaptations that enable the
shoots to penetrate the soil. The
roots are short, but there is little
need for water absorption
because little water is lost by the
shoots.
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,
phytochrome.
Reception … Transduction … Response
CELL CYTOPLASM
WALL

  1 Reception 2 Transduction 3 Response

Activation
of cellular
Relay molecules responses

Receptor

Hormone or
environmental
stimulus
Plasma membrane

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
Greening…an example of signal transduction

1 Reception   2 Transduction 3 Response

Transcription
CYTOPLASM
factor 1 NUCLEUS
Specific
Plasma cGMP protein P
membrane kinase 1
Second messenger activated
produced Transcription
Phytochrome factor 2
activated
by light 2 One pathway uses cGMP as a
second messenger that activates P
Cell a specific protein kinase.The other
Specific
wall pathway involves an increase in
protein
cytoplasmic Ca2+ that activates
kinase 2
another specific protein kinase.
activated Transcription

Light
Translation

3 Both pathways
1 The light signal is lead to expression De-etiolation
detected by the of genes for proteins (greening)
Ca channel
2+
phytochrome receptor, that function in the response
opened
which then activates de-etiolation proteins
at least two signal (greening) response.
transduction pathways.
Ca2+
Tropisms
Plant Hormones and Tropisms
• Hormones: Chemical signals that coordinate
growth, development, and responses to stimuli
• The discovery of plant hormones came from
work with tropisms
– Any growth response that results in curvatures
of whole plant organs toward or away from a
stimulus is called a tropism
– Tropisms are often caused by hormones
Darwin’s experiments with Phototropisms

EXPERIMENT In 1880, Charles Darwin and his son Francis designed an experiment to determine
what part of the coleoptile senses light. In 1913, Peter Boysen-Jensen conducted an experiment to
determine how the signal for phototropism is transmitted.

RESULTS

Control Darwin and Darwin (1880) Boysen-Jensen (1913)

Shaded
side of
coleoptile

Light
Light
Light

Illuminated Tip Tip covered Tip Base covered Tip separated Tip separated
side of removed by opaque covered by opaque by gelatin by mica
coleoptile cap by trans- shield block
parent
cap

CONCLUSION In the Darwins’ experiment, a phototropic response occurred only when light could
reach the tip of coleoptile. Therefore, they concluded that only the tip senses light. Boysen-Jensen
observed that a phototropic response occurred if the tip was separated by a permeable barrier (gelatin)
but not if separated by an impermeable solid barrier (a mineral called mica). These results suggested
that the signal is a light-activated mobile chemical.
Went’s experiment
• In 1926, Frits Went EXPERIMENT In 1926, Frits Went’s experiment identified how a growth-promoting
chemical causes a coleoptile to grow toward light. He placed coleoptiles in the dark and
removed their tips, putting some tips on agar blocks that he predicted would absorb the

– Extracted the
chemical. On a control coleoptile, he placed a block that lacked the chemical. On others,
he placed blocks containing the chemical, either centered on top of the coleoptile to
distribute the chemical evenly or offset to increase the concentration on one side.

chemical RESULTS The coleoptile grew straight if the chemical was distributed evenly.
If the chemical was distributed unevenly, the coleoptile curved away from the side with
the block, as if growing toward light, even though it was grown in the dark.

messenger for Excised tip placed

on agar block

phototropism, Growth-promoting
chemical diffuses
into agar block

auxin, by
modifying earlier Control
(agar block
lacking
Agar block
with chemical
stimulates growth

experiments Control
chemical)
has no
effect
Offset blocks
cause curvature

CONCLUSION Went concluded that a coleoptile curved toward light because its dark
side had a higher concentration of the growth-promoting chemical, which he named auxin.
Plant Hormones
Plant hormones
• In general, hormones control plant growth and
development
– By affecting the division, elongation, and
differentiation of cells

• Plant hormones are produced in very low

concentrations
– But a minute amount can have a profound
effect on the growth and development of a
plant organ
A Survey of Plant Hormones
Auxin
• The term auxin is used for any chemical
substance that promotes cell elongation in
different target tissues
• Auxin is involved in the formation and
branching of roots (Lateral and Adventitious
Root Formation)
• Auxin affects secondary growth by inducing
cell division in the vascular cambium and
influencing differentiation of secondary xylem
• Auxins as herbicides…an overdose of auxins
can kill eudicots (2,4-D is a synthetic auxin)
Cell elongation in response to auxin
• A model called the acid growth hypothesis suggests proton pumps
play a major role in the growth response of cells to auxin
3 Wedge-shaped expansins, activated
by low pH, separate cellulose microfibrils from
Cell wall cross-linking polysaccharides. The exposed cross-linking
enzymes polysaccharides are now more accessible to cell wall enzymes.
Cross-linking Expansin
4 The enzymatic cleaving
cell wall of the cross-linking
polysaccharides CELL WALL polysaccharides allows
the microfibrils to slide.
The extensibility of the
Microfibril cell wall is increased. Turgor
causes the cell to expand.
H2O
Cell
H+ Plasma wall
2 The cell wall H+ membrane
becomes more
H+
acidic.
H+
H+ H+ H+
H+

1 Auxin
increases the
Nucleus Cytoplasm
activity of
Vacuole
proton pumps. ATP Plasma membrane
H+ 5 With the cellulose loosened,
the cell can elongate.
Cytoplasm
Cytokinins
• Cytokinins
– Stimulate cell division

– Are produced in actively growing tissues such

as roots, embryos, and fruits
– Work together with auxin

– Retard the aging of some plant organs (anti-

aging effects)
Control of Apical Dominance
• Cytokinins, auxin, and other factors interact in
the control of apical dominance (The ability of a
terminal bud to suppress development of
axillary buds)

If the
terminal
bud is
removed
plants
become
bushier
“Stump” after
removal of
apical bud
Axillary buds

Lateral branches
Gibberellins
• Gibberellins have a variety of effects
– stem elongation

– fruit growth
– seed germination
Fruit Growth
• In many plants both auxin and gibberellins
must be present for fruit to set
• Gibberellins are used commercially in the
spraying of Thompson seedless grapes

Untreated Treated
Germination
• After water is imbibed, the release of gibberellins from
the embryo signals the seeds to break dormancy and
germinate
2 The aleurone responds by
1 After a seed synthesizing and secreting
imbibes water, the digestive enzymes that 3 Sugars and other
embryo releases hydrolyze stored nutrients in nutrients absorbed
gibberellin (GA) the endosperm. One example from the endosperm
as a signal to the is -amylase, which hydrolyzes by the scutellum
aleurone, the thin starch. (A similar enzyme in (cotyledon) are consumed
outer layer of the our saliva helps in digesting during growth of the
endosperm. bread and other starchy foods.) embryo into a seedling.

Aleurone
Endosperm

-amylase Sugar
GA

GA
Water

Radicle
Scutellum
(cotyledon)
Brassinosteroids
• Brassinosteroids
– Are similar to the sex hormones of animals

– Induce cell elongation and division

Abscisic Acid effects
1. Seed dormancy
• Seed dormancy has great survival value because it ensures
that the seed will germinate only when there are optimal
conditions

2. Drought tolerance
– Through a variety of mechanisms (For example, an
increasing amt of ABA in leaves will cause the stomata to
close to reduce water loss)

3. Inhibits growth
Ethylene
• Produced in response to
stresses such as drought,
flooding, mechanical EXPERIMENT Germinating pea seedlings were placed in the
dark and exposed to varying ethylene concentrations. Their growth
pressure, injury, and was compared with a control seedling not treated with ethylene.

infection RESULTS All the treated seedlings exhibited the triple

response. Response was greater with increased concentration.

• The Triple Response to

Mechanical Stress
– allows a growing shoot
to avoid obstacles
during soil penetration

1. Stems elongate less

rapidly 0.00 0.10 0.20 0.40 0.80

Ethylene concentration (parts per million)

2. Stems thicken
CONCLUSION Ethylene induces the triple response in pea seedlings,
3. Stems grow horizontally with increased ethylene concentration causing increased response.
Other Ethylene effects
• 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 production triggers
the ripening process

• Leaf Abscission: a change in the balance of

auxin and ethylene controls leaf abscission
(the process that occurs in autumn when a leaf
falls)
Plant Responses to Light
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
EXPERIMENT Researchers exposed maize (Zea mays) coleoptiles to violet, blue, green, yellow,
orange, and red light to test which wavelengths stimulate the phototropic bending toward light.

RESULTS
The graph below shows phototropic effectiveness (curvature per photon) relative
to effectiveness of light with a wavelength of 436 nm. The photo collages show coleoptiles before
and after 90-minute exposure to side lighting of the indicated colors. Pronounced curvature occurred
only with wavelengths below 500 nm and was greatest with blue light.
Phototropic effectiveness relative to 436 nm

1.0

0.8

0.6

0.4

0.2

0
400 450 500 550 600 650 700

Wavelength (nm)

Light

Time = 0 min.

Time = 90 min.

CONCLUSION

The phototropic bending toward light is caused by a photoreceptor that is

sensitive to blue and violet light, particularly blue light.
Light Receptors (two major classes)
• Blue-light photoreceptors
– Control hypocotyl elongation, stomatal
opening, and phototropism

• Phytochromes
– Regulate many of a plant’s responses to light
throughout its life. (such as seed germination)
Seed Germination Experiment
EXPERIMENT During the 1930s, USDA scientists briefly exposed batches of lettuce seeds to red
light or far-red light to test the effects on germination. After the light exposure, the seeds were placed in
the dark, and the results were compared with control seeds that were not exposed to light.
RESULTS The bar below each photo indicates the sequence of red-light exposure, far-red light
exposure, and darkness. The germination rate increased greatly in groups of seeds that were last exposed
to red light (left). Germination was inhibited in groups of seeds that were last exposed to far-red light (right).

Dark (control)

Red Dark Red Far-red Dark

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

CONCLUSION 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 were reversible.
Phytochrome switch
• Phytochromes exist in two photoreversible states (isomers)
with conversion of Pr (red absorbing) to Pfr (far-red
absorbing) triggering many developmental responses
• When seeds are exposed to adequate sunlight for the first
time, it is the appearance of Pfr that triggers germination
Pr Pfr
Red light
Responses:
Synthesis seed germination,
control of
flowering, etc.
Far-red
light

Slow conversion Enzymatic

in darkness destruction
(some plants)
Phytochromes and Shade Avoidance
• The phytochrome system also provides the
plant with information about the quality of light
• In the “shade avoidance” response of a tree
– The phytochrome ratio shifts in favor of Pr when a tree
is shaded. (amount of Pr greater than amount of Pfr)

– This causes the tree to allocate more resources to

growing taller (vertical growth) and less to branching
– Lateral branching occurs in plentiful direct sunlight
because the phytochrome ratio favors Pfr (Pfr >Pr)
Biological Clocks and Circadian Rhythms
• Many plant processes oscillate during the day
– For example, many legumes lower their leaves
in the evening and raise them in the morning
(these are called sleep movements)

Noon Midnight
Circadian rhythms
• cyclical responses to environmental stimuli

• approximately 24 hours long

• can be entrained (set) to exactly 24 hours by

the day/night cycle by daily signals from the
environment
• Human examples include: blood pressure,
body temperature, alertness, sex drive,
metabolic rate, etc. etc.
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

• Photoperiod, the relative lengths of night and

day is the environmental stimulus plants use
most often to detect the time of year
• Photoperiodism
– Is a physiological response to photoperiod
Photoperiodism and Control of Flowering
• Flowering in many species requires a certain
photoperiod
• Short-day plants (generally flower in late
summer, fall, or winter) (mums… poinsettias)
• Long-day plants (flower in late spring or early
summer) (lettuce…iris)
• Day-neutral plants are unaffected by
photoperiod and flower at a certain stage of
maturity regardless of day length at the time
(tomato…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
EXPERIMENT During the 1940s, researchers conducted experiments in which periods of darkness were
interrupted with brief exposure to light to test how the light and dark portions of a photoperiod affected flowering
in “short-day” and “long-day” plants.
RESULTS
Darkness

Flash of
light

24 hours
Critical
dark
period
Light

(a) “Short-day” plants (b) “Long-day” plants

flowered only if a period of
continuous darkness was
longer than a critical dark
period for that particular
species (13 hours in this
example). A period of
darkness can be ended by a
brief exposure to light.
flowered only if a
period of continuous
darkness was shorter
than a critical dark
period for that
particular species (13
hours in this example).

CONCLUSION The experiments indicated that flowering of each species was determined by a critical period of
darkness (“critical night length”) for that species, not by a specific period of light. Therefore, “short-day” plants are
more properly called “long-night” plants, and “long-day” plants are really “short-night” plants.
Test for presence of a flowering hormone
EXPERIMENT To test whether there is a flowering hormone, researchers conducted an
experiment in which a plant that had been induced to flower by photoperiod was grafted to
a plant that had not been induced.

Does a RESULTS
Plant subjected to photoperiod Plant subjected to photoperiod

flowering that induces flowering that does not induce flowering

Graft

hormone
exist
(florigen)?
Time
(several
weeks)

CONCLUSION Both plants flowered, indicating the transmission of a flower-inducing

substance. In some cases, the transmission worked even if one was a short-day plant
and the other was a long-day plant.
Experimental evidence for a flowering hormone

24 hours 24 hours 24 hours

Graft

Short-day Long-day plant Long-day

plant grafted to plant
short-day plant
Meristem Transition and Flowering
• Whatever combination of environmental cues
and internal signals is necessary for flowering
to occur …the outcome is the transition of a
bud’s meristem from a vegetative to a flowering
state
Plant response to
Non-Light stimuli
Gravity
• Response to gravity is gravitropism

• Roots show positive gravitropism

• Stems show negative gravitropism

Statoliths
• Plants may detect gravity by the settling of statoliths
(specialized plastids containing dense starch grains) to
lower portions of cells.
• How does it work?...maybe because of their density they
enhance gravitational sensing in some way?

Statoliths
20 m

(a) (b)
Gravitropism

https://www.youtube.com/watch?v=mYZXax8V_L0

https://www.youtube.com/watch?v=5JXm1USHlQY

https://www.youtube.com/watch?v=RzD4skFeJ7Y
Response to Mechanical Stimuli
• Thigmomorphogenesis
refers to the changes in
form that result from
mechanical perturbation
– Rubbing the stems of
young plants a couple
of times daily results in
plants that are shorter
than controls

Rubbed Un-rubbed
Thigmotropism
•Growth in response to touch occurs in vines and
other climbing plants.

https://www.youtube.com/watch?v=dTljaIVseTc
Rapid leaf movement in response to mechanical stimulation

https://www.youtube.com/watch?v=bczox-dDKP0
Response 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

• Drought
– During drought plants respond to water deficit
by reducing transpiration
– Deeper roots continue to grow
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
Vascular
cylinder

Air tubes

Epidermis

100 m 100 m
(a) Control root (aerated) (b) Experimental root (nonaerated)
Other stresses
• Salt Stress…Plants respond to salt stress by
producing compatible solutes (solutes tolerated at
high concentrations) which keeps the water
potential of cells more negative than that of the
soil solution
• Heat Stress… Heat-shock proteins help plants
survive heat stress by protecting important
molecules from denaturation
• Cold Stress…Altering lipid composition of
membranes to maintain fluidity of membranes is
one response to cold. Increasing levels of solutes
(like sugar) in the cells helps some frost-tolerant
plants to avoid freezing
Plant Defenses
 Defenses Against Herbivores
• Plants counter excessive herbivory
– With physical defenses such as thorns

– With chemical defenses such as distasteful or

toxic compounds
– Recruitment of predatory animals
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
Defenses Against Pathogens
• A plant’s first line of defense against infection
– Is the physical barrier of the plant’s “skin,” the
epidermis and the periderm

• Once a pathogen invades a plant

– The plant mounts a chemical attack as a
second line of defense that kills the pathogen
and prevents its spread
– The second defense system is enhanced by
the plant’s inherited ability to recognize certain
pathogens
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 by the protein products (receptors)
of specific plant disease resistance (R) genes

https://www.youtube.com/watch?v=L5vogXP4n1c
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
• A hypersensitive response against an
avirulent pathogen seals off the infection and
kills both pathogen and host cells in the region
of the infection
4 Before they die,
infected cells
release a chemical
signal, probably
salicylic acid.
3 In a hypersensitive
response (HR), plant
cells produce anti-
microbial molecules, 5 The signal is
seal off infected 4 Signal 5
distributed to the
areas by modifying rest of the plant.
their walls, and
then destroy Signal 6
themselves. This 3 Hypersensitive
transduction
localized response response 6 In cells remote from
pathway
produces lesions the infection site,
and protects other Signal transduction the chemical
parts of an infected 2 pathway Acquired initiates a signal
leaf. 7 resistance transduction
pathway.
2 This identification
step triggers a
signal transduction 1 7 Systemic acquired
pathway. Avirulent resistance is
1 Specific resistance is pathogen activated: the
based on the production of
binding of ligands molecules that help
from the pathogen protect the cell
to receptors in R-Avr recognition and Systemic acquired against a diversity
plant cells. hypersensitive response resistance of pathogens for
several days.
Systemic 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 Responses to Pathogen Invasions

https://www.youtube.com/watch?v=zW18wCNAivQ