RONA MAY S.

ESPERANZATE
MS BIOED
 theoretical basis of plant acclimation and plasticity
01 as a time-nested phenomenon

role of state transitions in response to changes

02 in light quality

OUTLINE
xanthophyll cycle and protection against
03 photodamage

mechanisms that allow plants to survive water

04 limitations
 low temperature acclimation and freezing tolerance
05

excitation pressure as a redox signal for retrograde

06 regulation of nuclear genes

OUTLINE
photosynthetic acclimation to high temperature
07

protective role of O2 as an alternative

08 photosynthetic electron acceptor during acclimation
 PLANT ACCLIMATION IS A TIME-DEPENDENT
PHENOMENON

• stress-tolerant plant species exposure

to a particular stress leads to
acclimation to that specific stress in a
time-dependent manner

• acclimation process in stress-resistant

species is usually reversible upon
removal of the external stress
1. Short term processes
2. Long term processes

 Time nested response

 Plant plasticity – the capacity to
regulate gene expression in
response to environmental
changes in a time nested manner
 STATE TRANSITIONS REGULATE ENERGY DISTRIBUTION
IN RESPONSE TO CHANGES IN SPECTRAL DISTRIBUTION

• Absorption cross-section
Referred as absorption coefficients and
the capacity of PSI and PSII to absorb
light

• State Transition
short-term mechanism to regulate
excitation energy distribution between
the two photosystems that is required to
maintain an efficient flow of electrons to
NADP+.
• Sunflecks
 Sudden transient
fluctuations influence rate
 Spot of direct sunlight
impinging on the leaf
through an open gap in the
canopy
 can last up to 20 minutes or
longer

RESULT:
Leaf may be subject to as much as
tenfold increase in energy, and could
have severe damaging effects on
PSII
Reversible phosphorylation of LHCII
• Protein Kinase
 Phosphorylation
 Activated when excess energy drives PSII ( state 2)

RESULT:
1. Increase the negative charge of protein causing LHCII
dissociate from PSII
2. Loosens the appression of thylakoid membranes
freeing a small portion of LHCII into PSI rich stroma

 State 2 – excitation of PSI

RESULT:
3. Plastoquinone reoxidized
4. Protein kinase is deactivated
5. Protein phosphates dephosphorylates the LHCII
6. LHCII migrate back into the appressed region
7. LHCII recombines with PSII
 CAROTENOIDS SERVE A DUAL FUNCTION: LIGHT
HARVESTING AND PHOTOPROTECTION

• β-carotene
• Small amounts of α-carotene
• Xanthophylls
 Lutein
 Violaxanthin
 Zeaxanthin

Principal Functions:
a. Light harvesting – transfer absorbed
light energy to chlorophyll
b. Photoprotection
• Any excess absorbed energy must be
dissipated. If not, the excess absorbed
energy may lead to photoinhibition of
photosynthesis
• ROS ( Reactive Oxygen Species)
 toxic forms of oxygen that may lead to
cell damage and ultimately cell death
• Ground state of 𝑂2 = 3𝑂2
• Prevents from chemically
reacting with most organic
molecules
• Singlet excited Chlorophyll
will be converted to triplet
excited chlorophyll
• Carotene can compete with
ground state oxygen and
prevent the formation of
singlet oxygen
OSMOTIC ADJUSTMENT IS A RESPONSE TO WATER STRESS

• Osmotic Adjustment
 response to water stress in many plants is a
decrease in osmotic potential resulting from an
• PROLINE
accumulation of solutes
 Amino acid that is sensitive to a particular
 net increase in solute concentration due to
stress
metabolic processes triggered by stress
 Exposure to water stress: loss of leaf
turgor and rapid accumulation of proline
RESULT:
 Sorbitol ( sugar alcohol) and betaine
1. More negative leaf water potential
(compatible solutes)
2. Helping the water movement into the leaf.
3. Helping partially wilted leaves to regain turgor
4. Enables plants to keep their stomata open and
continue taking up CO2 for photosynthesis
Osmotic adjuster
LOW TEMPERATURES INDUCE LIPID UNSATURATION AND
COLD REGULATED GENES IN COLD TOLERANT PLANTS

• increase in the proportion of

unsaturated fatty acids bound to lipids
• membrane can remain in a more fluid
and less gel-like state at lower
temperature which enhances
membrane stability and function at
these low temperatures (Transition
Temperature)
RESULT:
• Changes in mRNA transcription,
increases in protein synthesis, and
qualitative changes in the pattern of
proteins synthesized.
LEA-Proteins
cold-induced genes encode homologs of late
embryogenesis active proteins (LEA-proteins) that
are synthesized late in embryogenesis and during
dehydration stress.
Physical Properties of cold-regulated
genes:
1. They are unusually hydrophilic.
2. They remain soluble upon boiling in
dilute aqueous buffer.
3. They exhibit relatively simple amino
acid sequences that form amphipathic
α-helices.
Q10 FOR PLANT RESPIRATION VARIES AS A FUNCTION OF
TEMPERATURE

• Enzyme reactions typically are

considered to have a Q10
(temperature coefficient) of about 2,
which means that the rate of the
reaction doubles for each 10◦C rise in
temperature.
• The rate of reaction increases as
temperature until an optimum is
reached, beyond which the rate
usually declines sharply.

CAUSE: Thermal denaturation

• Q10 increases linearly upon short-
term increases in temperature from
40 to 10 ℃

2 primary factors
1. effect of temperature on the Vmax of
enzymes involved in respiratory
carbon metabolism as well as on the
maximum rate of respiratory electron
transport
2. effect of temperature on substrate
availability
LIGHT REGULATES NUCLEAR GENE EXPRESSION AND
PHOTOACCLIMATION

• leaves of high-light plants are usually

• Photoacclimation a pale green or yellow-green
 process whereby adjustments are compared to a dark green phenotype
made to the structure and function of of the same species grown at low light
the photosynthetic apparatus in
response changes in growth
irradiance

Consequence: pigment composition

which results in an altered visible
phenotype.
 Algae
• Low light phenotype
• High light phenotype  Decrease in total chlorophyll content
 modulation of the size  Decrease in light harvesting efficiency
 Modulation of composition of the light-
harvesting complex (LHCII) of PSII
 change in Rubisco content

RESULT:
 content of LHCII decreases on a leaf
area basis
 growth irradiance increases
 increased xanthophyll cycle activity
 increased photoprotection through
NPQ.
• DBMIB (2,5-dibromo-6-isopropyl-3-
methyl-1,4-benzoquinone)
 Inhibits electron transport chain at
cytochrome b6f complex

RESULT:
1. net accumulation of PQH2
2. PQ pool remains largely reduced
3. transcription of the nuclear Lhcb
genes coding for the major LHCII
polypeptides is repressed
4. inhibition in biosynthesis of LHCII

yellow phenotype typical of high-

light grown algal cells
• DCMU (3-(3,4-dichlorophenyl)-1,1-
dimethylurea
 Inhibits electron transport chain at
PSII

RESULT:
1. unable to reduce PQ to PQH2
2. PQ accumulates because any PQH2
pool is oxidized by PSI.

Green phenotype
 Phenotypic response to growth
under high light
• Chronic photoinhibition
• If persists over the long-term,
may lead to cell death.
ACCLIMATION TO DROUGHT AFFECTS SHOOT–ROOT
RATIO AND LEAF AREA

• Long term effects of Water deficit

1. Reduction in vegetative growth
2. Shoot growth
3. Growth of leaves
4. Reduction of leaf expansion

Reduced leaf expansion is beneficial

to a plant under conditions of water
stress because it leads to a smaller leaf
area and reduced transpiration.
 An early response to water
for the reduction of
deficits is closure of stomata to
CO2, the continuous consumption
conserve water. In some plants
of O2 by Rubisco requires the
this may lead to low internal leaf
continuous regeneration of RuBP
CO2 concentrations which will
by the Calvin Cycle
limit photosynthetic capacity
 COLD ACCLIMATION MIMICS PHOTOACCLIMATION

• low temperatures results in enhanced • Growth of cold-tolerant at low

freezing tolerance is measured as temperature stimulates photosynthetic
LT50 (freezing temperature at capacity with minimal changes in
which 50 percent of a population of photosynthetic efficiency or in the
plants are killed) ratios of chlorophyll a/chlorophyll b.
• enhanced freezing tolerance is
strongly correlated with the expression
of cor genes
• decreased sensitivity to
photoinhibition exhibited by these cold
acclimated plants mimics
photoacclimation to high light
 CAUSES:
1. Cold acclimation enhances the transcription and translation of genes
encoding major regulatory enzymes of stromal and cytosolic carbon
metabolism such as Rubisco, chloroplastic FBPase, cytosolic FBPase, and
sucrose-P synthase (SPS), as well as increased fructan biosynthesis in the
vacuole.
RESULT: higher total enzyme activity and a higher flux of carbon through the
sucrose biosynthetic pathway.
2. Coupled to higher rates of carbon export from the leaves in the light due to
enhanced sink activity
3. Cold acclimation suppresses photorespiration which also enhances net
carbon gain.
 FREEZING TOLERANCE IN HERBACEOUS SPECIES IS A
COMPLEX INTERACTION BETWEEN LIGHT AND LOW
TEMPERATURE

Herbaceous species grown at low temp • Photosynthesis continues during the

• short, compact growth habit, thicker cold acclimation period
leaves due to an increase in leaf
mesophyll cell size
• increase in the number of palisade
cell layers,
• increase in cell cytoplasm associated
with a decrease in leaf water content.
cold-acclimated state and maximum freezing tolerance
 COLD ACCLIMATED PLANTS SECRETE ANTIFREEZE
PROTEINS
Antifreeze proteins (AFPs)
• inhibit ice crystal growth by binding to
the surface of a growing ice crystal via
hydrogen bonding between specific
hydrophilic amino acids present in the
AFP and water within the crystal lattice
of ice.
WINTER RYE ACCLIMATION
• ethylene, is produced.
• Ethylene induces the transcription of the family
• of genes that encode AFPs.
• Upon translation, the AFPs are secreted via the
endoplasmic reticulum, Golgi bodies, and
vesicles that fuse with the plasma membrane
• deposited on the surface of the cell wall where
they inhibit ice crystal formation.
 OXYGEN MAY PROTECT DURING ACCLIMATION TO
VARIOUS STRESSES
• O2 can also act as alternative electron
acceptor for photosynthetic electron
transport.
• The photoreduction of oxygen by PSI is
called the Mehler reaction and results
in the production of another toxic,
reactive oxygen species known as a
superoxide radical.
• An effective system for the removal of
superoxide is the ubiquitous enzyme
superoxide dismutase (SOD)
• Hydrogen peroxide reduced to water in the
chloroplast by sequential reduction with
ascorbate (vitamin C), glutathione, and
NADPH
Asada-Halliwell Pathway
• Reduction of H2O2 is necessary in order
to prevent its reaction with𝑂−2 to form the
highly toxic hydroxyl radical (OH·)
• capacity to reduce O2 in the dark through
the chlororespiratory pathway
• results in the reduction of the thylakoid PQ
pool in the dark.