STRESS PHYSIOLOGY

INTRODUCTION
• Biological stress is a potentially harmful or injurious effect of unfavourable
environmental conditions
• Stress is an adverse environmental force or influence that tends to inhibit
the normal functioning of biological systems, with a disadvantageous effect
on plant life
• Water stress, temperature stress, chilling stress, salt stress, etc. are
examples
• Stress factors include drought, very low soil moisture, extremes of cold and
hot temperatures, desiccating atmosphereic conditions, prolonged
cloudiness, high wind velocities, high levels of UV radiation, excessive soil
salinity, severe air pollution, snow etc.
• An environmental factor, that is stressful for one plant may not be stressful
for another.
 Biological effects of stress
• The biological effect of stresses is generally called strain.
• Strains are of two main categories – elastic and plastic.
• Elastic strains are temporary and reversible. So, the affected plants can return to
normalcy when the stress ceases to exist. Example the reduction in
photosynthesis in very low light, and the resumption of normal photosynthetic
rate when bright light is again available.
• Plastic strains are permanent and irreversible. So, the affected plant will not
return to normalcy even when the stress ceases to exist. Examples are the
damages caused by extreme water deficit, drought, frost, heat, high salinity, etc.
• The commonest harmful effects of stresses include marked metabolic
dysfunction, reduction in photosynthesis, arrested growth, stomatal closure,
wilting of leaves and shoots, etc.
• If the stress is severe, it may prevent flowering and seed-setting, or may
endanger the survival of the plant.
 Response of plants to stress

• Plants react to stresses and avoid or overcome their effects by

gradually developing several kinds of biochemical and
physiological counter mechanisms, adjustments and
specializations.
• If the stress is mild, moderate, or less severe, and short living, its
harmful effects may be transient or temporary and the plant may
recover when the stress is over.
• However, when the intensity or severity of stresses approaches
the extreme level, plants may suffer from serious damages, or
may succumb to death.
• The innate ability of plants to avoid, escape, endure, or resist
stresses is known as hardiness or acclimation.
• Acclimation differs from adaptation, even though both are
effective means for achieving tolerance against a particular
stress.
• Adaptation is a genetically determined and heritable structural
or functional modification which can increase the fitness and
survival value of organisms in a stressful environment.
• It is achieved through natural selection over many generations.
• On the other hand, acclimation is a non-heritable physiological
modification, induced by environmental stress.
• It occurs during the life of an individual and enables it to survive
and reproduce in the stressful environment.
Plants survive stresses by several strategies

• Stress avoidance
• Stress escape (stress evation)
• Stress tolerance (stress endurance)
Stress avoidance
• The internal environment of the plant is modified in
such a way that cells become safe from any stress
problem
• In this case, plants respond to stresses by somehow
reducing the impact of the stress
Stress escape
• In stress escape, plants germinate, grow and flower
very quickly during the favourable season and
complete their life cycle before the onset of the next
unfavourable season.
• Ephemeral plants and some arctic annuals are best
stress escapers.
 Stress tolerance
• In stress tolerance, plants develop high tolerance or endurance
to withstand the harmful effects of stresses. In this case, the
plant comes to a state of thermodynamic equilibrium with the
stress.
• In such an equilibrium, the internal conditions of the plant are
in equilibrium with the external environmental conditions.
• High drought tolerance of some desert plants is an example.
• Stress tolerance often increases with previous exposure to
stresses which makes the plant hardy, acclimated, or
conditioned to survive
 WATER STRESS
• Water is indispensable for the life of plants. So, its extreme deficiency or
high excessiveness may serve as stress factors.
• Sometimes, the rate of transpiration far exceeds the rate of water
absorption.
• This also creates a temporary water stress in plants. But, it causes only the
closure of stomata, reducing gas exchange; the plant will not suffer from
any lasting or serious problem.
• However, extreme scarcity of water causes prolonged water stress and very
seriously affects the physiological functions, growth and reproduction of
plants.
• It will adversely affect shoot growth, root growth, leaf expansion, flowering
and fruit-setting.
• This is because extreme deficiency of water within the plant body destroys
proteins, increases solute concentration, makes cell sap and intracellular
fluid highly concentrated or hypertonic and reduces the water potential of
intracellular and intercellular fluids.
• Water stress results either from an excess of water or from extreme water
deficit.
• So, it includes water surplus stress and water deficit stress. Excess water
results from flooding.
• Flooding stress is simultaneously an oxygen stress also, as far as plants are
concerned.
• This is primarily due to the insufficient supply of oxygen to roots.
• Oxygen deficit, in turn, very seriously affects respiration, nutrient uptake
and other critical root functions.
• Water deficit stress is far more common than water surplus
stress. So, in reality, water stress represents water deficit
stress.
• Under natural conditions, water stress results from
drought or lack of rainfall.
• So, it is often termed drought stress. Water stress is equally
a component of salt stress and osmotic stress also.
• The commonest causes of water stress include
• (i) sub-optimal availability of soil moisture and
• (ii) high rate of transpiration which is in excess of the rate
of absorption of soil water by roots.
 Consequence of water stress
• Dehydration of protoplasm and deleterious molecular and
physiological changes.
• Changes in the molecular organization of enzymes
• Differential response of different physiological processes
• Closure of stomata
• Decrease in the rate of net photosynthesis
• Decrease in the rate of translocation of photosynthate from
leaves to other parts
• Marked reduction in vegetative growth
• Wilting of plants and the drooping and sagging of leaves and
other parts.
Adaptations of plants against water stress
• Short life cycle for stress avoidance.
• This involves rapid seed germination and vegetative growth and early
flowering and seed-setting, when water is available
• Development of deep roots to seek out available soil water and
thereby to encounter drought stress.
• Thick cuticle, sunken stomata, low water content of seeds, etc. to
encounter drought.
• Orientation of the leaves parallel to sun rays to minimize the
absorption of solar radiation and thereby to reduce transpirational
water loss.
• High water stress tolerance and the ability for minimal photosynthesis
even at extremely low levels of leaf water potential.
• Deciduous plants shed their leaves during dry season, paving the way for the
appearance of small new leaves in which the rate of transpiration would be
much lower. This, in turn, reduces water loss and drought stress.
• Succulent plants store enough water in their tissues for protection from
drought.
• Non succulent xerophytes develop high water tolerance and great drought
endurance
• Some annual desert plants escape drought by remaining as dormant seeds.
On the advent of rainy season, seeds germinate and seedlings grow fast.
• Small cells, high nucleic acid content, high sugar content and low starch
content in drought tolerant plants.
• Early flowering and the production of tiny fruits and seeds during prolonged
drought. This is believed to be an adaptation to minimize water utility for
survival.
 SALT STRESS
• Salt stress refers to the excessive concentrations of mineral ions in rhizosphere and
plants and their effect on the life of plants.
• Higher plants, growing in salt marshes, along sea-shores, on the banks of estuaries,
etc., have to encounter the perennial problems of very high salt concentration and
excessive salinity in the soil.
• High salinity is characteristic of inland deserts and the shores of inland lakes also.
• In inland deserts, evaporation exceeds precipitation and there is only very little
leaching.
• So, minerals accumulate in the surface soil, especially Na+, Cl-, Ca2+, SO42_ and
carbonates.
• In those inland lakes which have no outlets (e.g., Dead sea), salts accumulate as water
evaporates.
• In heavily irrigated agricultural fields, there is extensive water loss through evapo-
transpiration. This results in the accumulation of salts in the surface soil
• Based on the tolerance against salt stress, two groups of plants can
be recognized, namely halophytes and glycophytes or non-
halophytes.
• Halophytes are highly salt-tolerant plants which can grow and
flourish in saline soils.
• On the other hand, glycophytes are plants with less salt-tolerance.
• They often exhibit growth retardation, leaf discolouration, loss of
dry weight, etc., when soil salinity goes beyond the threshold level.
• Halophytes are of two main kinds, euryhaline halophytes and
stenohaline halophytes.
• The former can tolerate a wide range of soil salinity, whereas the
latter can tolerate only a narrow range of soil salinity
 Effects of salt stress
• Salt injury.
• Abnormally higher concentrations of salts in the cell sap may
adversely affect the water relations and metabolism of plants.
• They may interfere with the functioning of certain enzymes. This,
in turn, may lower the metabolic rate.
• Heavy accumulation of chloride ions within plants may
considerably lower the rates of absorption and transpiration of
water.
• It may also inflict direct injuries, such as leaf edge burns, necrotic
leaf spots, bleaching of chlorophyll, etc.
Salt stress can cause damage to plants in these ways:

• High salt accumulation in general, and high sodium accumulation in

particular, will abnormally alter soil structure.
• This is by decreasing the porosity of the soil and adversely affecting the
aeration of the soil. These alternations harmfully affect plant life.
• High salt concentration contributes much to water stress, generates low soil
water potential and causes a form of physiological drought.
• Under these conditions, plants find it very difficult to take up water and
nutrients from the soil.
• High salt accumulation, especially that of Na+ and Cl-, causes toxic effects in
plants. This is particularly common among glycophytes.
• The toxic effects include disorganization of membranes, inhibition of
enzymes, metabolic dysfunction, suppression of growth, reduction of carbon
assimilation, etc.
 Physiological adaptations against salt stress

• They are primarily meant for coping with the high salinity
of soil and water.
• They include high salt tolerance, high rate of salt
elimination, high rate of water loss, and high ability to
absorb salty water with easiness.
• Many species have evolved coping strategies like salt
avoidance, salt accumulation, salt endurance (salt
tolerance) and salt regulation
• Salt regulation is effected by two main mechanisms,
namely salt exclusion and salt excretion
Salt avoidance
Plants accomplish salt avoidance by:
• Limiting germination, growth and reproduction
to specific favourable seasons
• By sending roots to non saline regions
• By limiting the uptake of salts
Salt exclusion
• Salt exclusion is found in some mangrove plants, such
as Rhizophora.
• In Rhizophora, there is a salt-excluding mechanism in
the root system.
• It allows only the selective passage of only water and
essential nutrients to the tree, and blocks the free
passage of salts.
• Thus, Rhizophora does not absorb salt, but actively
exclude it from its roots.
 Salt excretion
• Salt excretion is the secretion and elimination of excess
salts by the salt glands in the leaves.
• It occurs through the exudation of plant sap which
contains dissolved salts (e.g. Avicennia).
• In some desert plants 9e.g. Tamarix), excretions of excess
salts form heavy incrustations of salts on leaf surface.
 Salt accumulation
• Salt accumulation is a form of salt evation. In this case, salts
accumulate in specific cells of the plant ( e.g., Sonneratia, Atriplex).
• Active accumulation of salts increases the osmotic concentration
of the cell sap and thereby enables the plants to easily absorb
saline water.
• It has been shown that the bark and mature leaves of Sonneratia
are the store-house of several salts and elements.
• In some halophytes, Na, K, Ca, Mg etc form granular
accumulations in some leaf cells and pith.
 Salt endurance
• Salt endurance is the high tolerance of plants to the toxic
effects of high salt concentration within their cells.
• It is found in those plants whose protoplasm can endure
high salt contents without the danger of metabolic
disorder and tissue damage.
• Plants having only very narrow range of salt tolerance
(stenohaline) are called salt-sensitive plants and those
with wide range of salt tolerance (euryhaline) are called
salt-insensitive plants.
TEMPERATURE STRESS
• Upper and lower extremes of temperature are stress
factors for many plants.
• Temperature stress includes both cold-stress and heat
stress.
• The former involves extremely low temperature, and
the latter involves extremely high temperature or
heat.
 Cold stress
• Very low temperature is harmful to many plants, especially during autumn and
winter.
• Cold stress is of two kinds, namely chilling stress and frost stress or freezing stress.
• The former results from low temperatures a little above the freezing level of plants,
whereas the latter results from low temperatures well below the freezing level of
plants.
• Frost stress often causes ice formation within plant tissues.
• The tolerance or resistance of plants against cold stress varies with different species.
• Some plants have only very low cold tolerance and so they are highly sensitive to
cold.
• Some others have moderate tolerance to cold and they are sensitive only to
extremely low temperatures.
• Still others are totally cold-resistant and they are insensitive to cold.
 Chilling stress

• Chilling temperature is seriously harmful to many plants of the tropical and

subtropical regions.
• So, they are easily susceptible to chilling injury and hence are called chill-
sensitive plants.
• Exposure to a very low temperature of 1 to 5 degree celsius for 24 to 36
hours may be highly damaging or even fatal for them.
• Examples are the crop plants, such as rice, beans, peanut, cotton, etc.
Ornamental plants, such as Coleus, Croton, Passiflora etc., are also chill-
sensitive or chill-intolerant.
• Many others can withstand chilling temperatures and so they are called
chill-resistant or chill-tolerant plants. Maize, sorghum, tomato, pumpkin,
etc. are examples.
 Damages caused by chilling stress
• Stunted growth of plants
• Chlorosis and lesions in leaves
• Water-soaked appearance of foliage
• Wilting and ultimate death of the plant in extreme cases
These occur due to
• Disorganization and loss of function of cell membranes
• Inhibition of photosynthesis, protein synthesis and
carbohydrate translocation
• Increased protein degradation
• The membranes of chill-sensitive or chill-intolerant plants
will have high proportions of saturated fatty acids, while
those of chill-insensitive on chill resistant plants will have
high proportions of unsaturated fatty acids.
• At chilling temperatures, the saturated fatty acids of the
membranes of chill-sensitive plants are believed to solidify,
disrupting the normal functioning of the membrane.
• On the other hand, the higher proportions of unsaturated
fatty acids in the membranes of chill-resistant plants will
maintain the fluidity and the normal functioning of the
membranes even at chilling temperatures
 Frost stress
• Below freezing temperature it is lethal or seriously injurious to a
number of plants, especially to those of the temperate region.
• In such frost-sensitive plants, temperatures well below the
freezing point of water will cause ice formation in intercellular
spaces and xylem vessels.
• Ice formation is a slow and gradual process and it occurs once the
plant has become sufficiently supercooled.
• It begins with ice nucleation, a process in which ice crystals are
formed around some large polysaccharides and proteins of the
cell wall.
• These ice crystals gradually enlarge by drawing water from the
adjoining cells. As a result, the neighbouring cells get dehydrated.
• This dehydration causes enzymatic dysfunctioning, metabolic
disorders a and structural disorganization.
• These harmful effects are collectively called frost injury.
• In some cases, frost injury occurs during ice formation and
dehydration.
• But in others, it occurs during the subsequent thawing and re-
hydration of the dehydrated cells, when ice melts and water
reenters the cells.
• Rapid thawing and rehydration, most usually, may cause distortion
and death of cells.
• It is claimed that frost-resistance of plants in due to the
synthesis of certain proteins, called anti-frost proteins.
• They probably bind with ice crystals and check their
enlargement.
• They are also believed to stabilize cell membranes during
the dehydration of cells consequent to supercooling.
• Some sugars and proteins probably function as
cryoprotectants to control the enlargement of ice crystals
and to protect the cells from frost-injury
 Heat stress & its consequences

• Many plants can withstand and survive high temperature. But

some which cannot tolerate a wide range of high temperature are
prone to heat stress.
• Heat stress has both direct and indirect effects on plants.
• The direct consequences of heat stress include changes in the
kinetics of metabolism, respiratory degradation, protein
degradation, lipid degradation, etc.
• Indirect effects include inhibition of photosynthesis and
respiration, denaturation of proteins and nucleic acids, injury and
dysfunctioning of cell membranes, metabolic disturbances due to
starvation, very low rate of metabolism, high mobilization of
toxins within cells, etc.
 Adaptations for resistance against heat stress
• Increased production of enzymes to make up the loss caused by
the heavy destruction of enzymes by heat.
• Excessive synthesis of thermostable heat-shock proteins
• High proportion of saturated fatty acids in the membrane lipids to
keep up the stability and fluidity of cell membranes
• Vertical orientation of leaves to reduce the area of direct exposure
to intense solar radiation
• Reflective leaf hairs and waxy leaf surface to reflect solar radiation
• Small and deeply dissected leaves to minimize the marginal
thickness of leaves and thereby to maximize heat loss through
convection and conduction
 Heat shock proteins
• HSPs are a unique group of low-molecular weight heat-sensitive proteins
which help plants to endure heat stress.
• They serve as molecular chaperones and thereby prevent the misfolding and
denaturation of proteins during heat stress (chaperones are specialized
proteins which bring about the correct folding of individual polypeptide
chains to attain their stable and functional final configuration).
• Also, the HSPs allow the denatured proteins to refold correctly to their
functional form.
• Any stress caused by heat, toxins, free radicals, heavy metals, etc. will
promote the increased production of HSPs as a counter measure to check
the effects of heat stress.