Defining Plant Stress
Ideal conditions: allows the plant to achieve its maximum growth and
reproductive potential as measured by plant weight, height, and seed
number, which together comprise the total biomass of the plant.
Stress:any environmental condition that prevents the plant from achieving
its full genetic potential.
For example, decrease in water availability would have deleterious effect
on growth because of reduce in water potential is by closing their stomata,
which reduce water loss by transpiration. It also reduces the CO2 uptake
which decrease the photosynthesis.
Physiological adjustment to abiotic stress involves trades offs between
vegetative and reproductive development.
Acclimation and Adaption
Acclimation:nonpermanent change in physiology or morphology of the
individual to improve response with exposure to environmental stress.
Epigenetic mechanism that alter expression of genes without changing the
genetic code
Adaptation:Fixed genetic change over many generations by selective
environmental pressure
Environmental Factors and their
biological impacts on plants
Environmental
Factor
Primary Effect Secondary effect
Water Deficit Water potential
reduction,
Cell dehydration,
Hydraulic
resistance
Reduced cell/leafexpansion, cellular/metabolic
activities, stomatal closure, photosynthetic inhibition,
leaf abscission, Altered Carbon partitioning,
Cytorrhysis, cavitation, membrane and protein
destabilization, ROS production, Ion cytotoxicity, cell
death
Salinity Water potential
reduction,
Cell dehydration,
Ion Cytotoxicity
Same as for Water deficit
Light Stress Photo inhibition
ROSproduction
Inhibition of PSIIrepair
Reduced CO2 fixation
Environmental Factors and their
biological impacts on plants
Environmental
Factor
Primary Effect Secondary effect
High Temperature Membraneand
protein
destabilization
Photosyntheticand respiratory inhibition, ROS
production, Cell death
Chilling Membrane
Destabilization
Membrane dysfunction
Flooding and soil
Compaction
Hypoxia,
Anoxia
Reduced respiration, Fermentativemetabolism,
inadequate ATP production, production of toxins by
anaerobic microbes, ROS production, stomatal
closure
Environmental Factors and their
biological impacts on plants
Environmental
Factor
Primary Effect Secondary effect
Freezing Water Potential
reduction,
Cell hydration,
Symplasticice
crystal formation
Same as for water deficit
Physical destruction
Trace element
toxicity
Disturbed cofactor
binding to proteins
and DNA,
ROSproduction
Disruption of metabolism
Mimic other essential metals
Mineral Nutrient
deficiencies
Reducedgrowth
and unavailable
for uptake
Ceases energy production
Environmental Factors and their
biological impacts on plants
Ozone and ultraviolet light generate ROS that cause lesions and induce
PCD
Combination of abiotic stresses induce unique signaling and metabolic
pathways
Combination of abiotic stresses have both positive as well as negative
impacts
Sequential exposure to different abiotic stresses sometimes confers cross
protection, for examples molecular chaperones and osmoprotectants for
ROC scavenging
Stress Sensing Mechanisms in Plants
Physical Sensing: mechanical effect of stress on plant such contraction of
plasm membrane
Biophysical Sensing: Change in protein or enzyme structure
Metabolic Sensing: accumulation of ROS
Biochemical Sensing: specialized protein to sense a particular stress, Ca
channel
Epigenetic Sensing: modification of DNA or RNA such change in chromatin
Signaling Pathways activated in
Response to abiotic Stress
Different pathways such as calcium, protein kinases, protein phosphatases,
ROS signaling, activation of transcriptional regulators, accumulations of
plant hormones
Stress specific signals that emerge from these pathways, in turn, activate or
suppress various network that may allow growth under stress conditions until
favorable conditions returns
Increase in the concentration of Ca and ROS are early signaling events
Ca regulates the transcription factors by binding directly or to form Ca
complexes.
Ca activates various protein kinases and phosphatases that regulate gene
expression either by phosphorylating (activating) or dephosphorylating
(inhibiting) transcriptional factors
Signaling Pathways activated in
Response to abiotic Stress
Steady state level of ROS is governed by the balance of ROS generating
and ROS scavenging reactions
ROS generation: Activities of Specialized oxidases
ROS Scavenging: Antioxidant molecules such as APX, CAT, SOD
ROS can trigger the opening of calcium channels which activate Ca
dependent protein kinases
For example, mitogen activated protein kinases regulates stresses (MAPK)
Acclimation to stress involves transcriptional
regulatory network called regulons
Transcriptional regulators or factors binds to specific DNA sequences and
activate or suppress the expression of genes.
Chloroplast genes respond to high intensity light by sending stress signals to
the nucleus
Epigenetic mechanisms and small RNAs provide additional protection
against stress
Hormonal interactions regulate normal development and abiotic stress
resonses
Developmental and Physiological
Mechanisms that protect plants
Plants adjust osmotically to drying soil by accumulating solutes
Submerged organs develop aerenchym tissue in response to hypoxia
Antioxidant and ROS scavenging pathways protect cells from oxidative stress
Molecular chaperones and molecular shields protect proteins and membranes
during abiotic stress (Heat shock Protein)
Plant can alter their membrane lipids in response to temperature and other
biotic stresses
Exclusion (block entry) and internal tolerance mechanisms allow plants to cope
with toxic ions: glycophytes and halophytes
Phytochelatins and other chelators contribute to internal tolerance of toxic
metal ions: chelating molecule have ligation sites
Plants use cryoprotectantmolecules and antifreeze proteins to prevent ice
crystal formations
ABA signaling during water stress
During water stress, ABA increases in leaves, which leads to stomatal closure
Stomata closure is due to reduction in turgor pressure that follows the massive efflux of K
and anions from guard cells
Activation of specialized ion efflux channels on the plasma membrane is required for such
a large scale loss of K and anions
Plasma membrane K efflux channels are voltage gated, they open only if plasma
membrane become depolarized.
ABA causes membrane depolarization by elevating the cytosolic Ca in two ways: transient
influx of Ca ions and release of Ca from internal stores
Increase in Ca open the Ca activated anion channel on the plasma membrane
Opening of anion channels allow Cl and malate ions to escape, moving down their
electrochemical gradients
ABA signaling during water stress
Outflow of negatively charged Cl and malate trigger the opening of voltage gated K efflux
channels
Elevated level of Ca cause K influx channels to close
ABA cause alkalization that further stimulate the opening of K efflux channels
ABA also inhibit the activity of the plasma membrane H-ATPase
During stomatal closure, surface area of guard cell contract 50 %. Extra membrane taken
up as small vesicles by endocytosis
Signal Transduction involves protein kinases and phosphatases
Plant can alter their morphology in
response to abiotic stress
Phenotype plasticity: plant activate developmental program that alter the
phenotype
Leaf area, leaf orientations, trichrome, cuticle, root: shoot ratio
Metabolic shifts enable plants to cope with variety of abiotic stresses
The process of recovery from the stress can be dangerous to the plant and
require a coordinated adjustment of plant metabolism and physiology
High level of ROS could form and damage cells.
Plants needs to remove recycle all the unneeded mRNAs and protein
Abiotic stress tolerance crop
Developing crops with enhanced tolerance to both biotic and abiotic
stress conditions is a major goal
Such crops would decrease the yield penalty and prevent annual losses of
billions of dollars