Bot.

502 Resistance Physiology

In both natural and agricultural conditions plants are frequently exposed to extreme
environment which create stressful conditions for plants and have significant impact on their
physiology, development and survival. Despite modern genotypes and agro technology farmers
still experience crop failure due to bad weather.

Principle Environment Stresses

Environment stress

Biotic Abiotic
(Pest, Pathogen Competition) (Any Kind of Physiochemical Stress)

Temperature Water Radiation Chemical Miscellaneous

High Deficit (Drought) Infrared Salts/ ions Wind
Low Excess (Flooding) Visible Gases Pressure
(Frost & Chilling) Ultraviolet Herbicides Sound
Ionizing, Fertilizers Magnetic
X rays Insecticides Electrical Stress

Jacob Levit (1972-1980) suggested that biological stress is any change in environmental
conditions that might decrease or adversely change plant growth and development i.e. normal
functions. When stress acts upon plant it produces injury in different ways:

1.Direct Stress Injury: It is irreversible, also called direct plastic strain. The plants may be killed
by a brief exposure to stress for eg. Rapid freezing strain (frost)

2.Indirect Stress Injury: It is reversible, also called elastic strain. Injury is produced by a long
exposure to stress. For eg. Chilling, Wilting.

3.Secondary Stress injury: Stress may injure the plant by giving rise to 2 nd stress for eg. High
temperature may not be injurious by itself but may produce water deficit which may injure
the plant. Here relatively long exposure to primary stress is required.

How Do Plants Response to Stress

They can respond by stress response

Stress Response

Resistance Susceptible Avoidance

Acclimation Senescence Survival

Growth Death

Survival

SUSCEPTIBILITY:
Some plants may be injured by stress i.e. they exhibit one or more metabolic
disfunction. If the stress is moderate/short term the injury may be temporary & when
stress is removed plant is recovered. If stress is severe it may prevent flowering, seed
formation and induce senescence that leads to death.

AVOIDANCE:
Ephemerals (short life cycle) complete their life cycle before on set of stress for eg.
Desert plants and arctic annuals, which complete their life cycle during summers, to
reduce the stress impact even when the stress is present in the environment.

RESISTANCE:
It is the capacity of a plant to tolerate particular stress. Here the organism comes to the
thermodynamics equilibrium to stress (homeostasis). Internal conditions are in
equilibrium with conditions outside the plant for eg. Resurrection plants like some ferns
and flowering plants the foliage will survive air drying as to as little 7% without injury.
Plants resist stress by adaptation or acclimation

ADAPTATION:
There are heritable modifications in the structure or function that increase the fitness of
an organism in stressful environment. For eg. Morphological & physiological
modifications associated with CAM plants.

ACCLIMATIONS: (ADJUSTMENT)
These are non heritable physiological modifications induced by gradual exposure to
stress for eg. Chilling. The capacity to acclimation is genetic trait and the process of
acclimation to stress is known as hardening. The plants with capacity to acclimation are
hardy plants while the plants with minimum capacity to acclimation are non hardy
species.

WATER STRESS

Excess water: Flooding- reduces oxygen supply limits respiration, nutrient uptake.

Water deficit stress: Drought- desiccation, salt stress, osmotic stress.

DROUGHT: It causes stress in a plant if too little water in a suitable thermodynamic

state is available. Stress due to drought does not occur abruptly but develop slowly and
increase in intensity with time. Generally the term drought denotes a period without
rainfall during which the water content in soil is reduced to such an extent that plant
suffers from lack of water and where annual evaporation exceeds annual rainfall i.e.
arid region. Drought may also caused by freezing temperature.

CELLULAR ASPECTS OF WATER POTENTIAL

Water potential (W.P) = ψA + ψP + ψM + ψG

Water potential may be defined as difference in the potential energy of pure water
( which has zero W.P)and the water potential in some system such as that in plant cell
or soil. W.P in plant or soil can be broken down in 4 major components:

1. Osmotic potential
2. Pressure potential
3. Metric potential
4. Gravitational potential

OSMOTIC POTENTIAL: Major component of water potential in living cells and is

major driving force by which water moves into cell. O.P is due to solutes dissolved in
water. It is either 0 or –ve because solute reduce capacity of water to do work. Four
compounds can compose most of osmotic components in symplast (living cells)
inorganic ions, organic ions, non protein amino acids (prolein, glycedetane) and simple
CHO’s). In organic ions and organic acids are higher in vacuoles while non protein
amino acids and simple carbohydrates like sucrose and raffinose are in higher
concentration in cytoplasm.

PRESSURE POTENTIAL: It is also key component and important in both living cells
and functioning xylem tissue ( nonliving & living). It can be –ve , 0 or +ve in value and
is a function of hydrostatic pressure of system. In a fully turgid cells the protoplast
press against the cell wall and the pressure pot. is +ve. When water is pulled through
the xylem vessels the pressure pot. is –ve and actually pulls the wall of vessel. So both
+ve & -ve pressure pot. exist in plants and both are essential for movement of water
with in plant and between soil, plants & atmosphere.

GRAVITATIONAL POTENTIAL: It results from pull of gravity on water and it is –ve

when water moves downward since the water is losing potential energy. It mostly
occurs in soil.

METRIC POTENTIAL: It is the result of cohesive forces that bind water to cell wall and
soil particles. It is always –ve because water has less –ve potential energy than free
water.

SOIL PLANT AIR CONTINAUM ( SPAC ):

The main driving force for water movement through plant is gradient of free energy for
water in soil solution to water vapour in the atmosphere. That column of water in plant
tissues is maintained by cohesive and adhesive properties of water molecule.

IMPORTANCE OF CELLULAR ASPECT OF WATER POTENTIAL

The W.P characteristics which is the best associated with water stress is change in
tissues water potential and not the absolute water potential. The water potential
component that changes most rapidly with changing water content is turgor pressure.
Turgor pressure is considered best indicator of plant water stress. The physiological
processes most sensitive to tissue water potential are cell expansion, cell wall growth,
protein synthesis and nitrate reduction.

FUNCTIONAL DISTURBANCES & PATTERNS OF INJURY DUE TO DROUGHT

1. CELL EXPANSION: Since the cell expansion is dependent upon pressure potential.
So developing cells will expand less and cell size will be less under drought
conditions. The reduction in cell size is dependent on the timing of water limitation
in relation to phenology of the plant. If this occurs during the beginning of growth
cycle leaf area will be reduced and carbon gain through out the growing season will
be reduced because of smaller leaves. If this occurs during inflorescence
development the no: of flowers will be reduced, all effort of reproduction may be
wasted. If occur during the fruit maturation , inflorescence will be normal and
entire plant mass will be unaffected but seed filling may be inhibited and fruit
abscission may be enhanced.

2. CELL ULTRA STRUCTURE: Removal of water leads to an increase in solute conc.

as protoplasm volume shrinks the normal bi layer membrane structure is
distracted, membrane becomes porous and selectivity is decreased. Mild water
limitation also disturb the structure of microbody releasing hydrolyzing enzymes in
cytoplasm. The presence of hydrolyzing enzymes disrupt the normal structure of
all cytosolic membranes for e.g. if tonoplast is degraded, vacuole fluid may empty
into cytoplasm and may damage the cytosolic proteins.

3. EFFECT ON PHOTOSYNTHESIS: This can be due to stomatal closure. The 1st

impact of drought is on stomatal closure which may be signal from root probably
ABA or because of low turgor pressure of guard cells. Stomatal closure induced by
water limitation causes a depletion of carbon dioxide in the intercellular spaces
once this CO2 has decreased relative to O2, photorespiration is stimulated and if
the light intensity is high enough photorespiration can utilize all energy providing
products of electron transport system. Under such conditions photosynthesis
inhibition can occur resulting in buildup of free radical of chloroplast.

4. NON STOMATAL EFFECTS: During the initial phases of water limitation stomatal
closure and non stomatal closure inhibition of photosynthesis occurs side by side.
 There are reports that non stomatal inhibition occurs 1 st thereby increasing level of
conc. of CO2 in the intercellular spaces and thus causing stomatal closure. Reduced
turgor may increase the permeability of outer chloroplast envelop which result in a
change in chloroplast pH and ion conc. The change in ion conc. and pH can affect
the activity of Rubisco. The degradation of chlorophyll increase and conc. of
binding protein complexes decrease during water stress, therefore the light
harvesting and electron transport associated with photo system II is decreased as
compared to PS I by water stress.

5. DARK RESPIRATION AND CARBOHYDRATES METABOLISM: As water

limitation progresses photosynthesis decrease before respiration. Consequently the
ratio between photosynthesis and respiration decrease because due to drought the
stomata close and there is no movement of CO2 from outside. Only CO2 is released
during respiration is used therefore there is little photosynthesis.

6. SOURCE SINK RELATION AND ROOT SHOOT RATIO: There is also change in
the source sink ratio during water stress. Low CO2 assimilation by leaves and
increased respiration in mesophyll cells of leaves reduce the gradient of sucrose
between source leaves and sinks. The reduced gradient from source to sink causes
the reduction in carbohydrate flow in the phloem. The timing of water limitation
relatives developmental status of plant affects allocation pattern. Water stress that
occurs during early growth phases caused by large shift in root shoot ratio because
there is more growth of roots and water stress during later stages has little or no
effect on the root shoot ratio but flowering and seed production is reduced and fruit
abortion is increased.

7. NITROGEN METABOLISM: Nitrate and ammonia accumulation decrease during

water limitation. Flow of nitrogen from roots to leaves slows and high conc. of NO3
and NH4 build up in water stressed roots than in roots of well watered plant.
Reduction of nitrogen is not due to specific effect of water stress on transport
proteins or accumulation mechanism but rather on change in nitrogen use and
result in conditions in plants that inhibit nitrogen accumulation kinetics.

HOW DO PLANTS COPE UP WITH WATER STRESS

Drought escape:
By rapid phenological development: They complete their life cycle when water
availability is high e.g. Argemone mexicana.

Developmental plasticity or deciduousness: In Mediterranean & desert climate leaf

population are developed in winter months when water availability is high. In later
spring & summer pre-drawn water potential decrease and most of leaves abscise.
Deciduous species survive the period of water stress in dormant stage. In desert plant
welwitchia only 2 leaves are produced before flowering.

By Extended dormancy: Many plants survive by perennating organs like tubers, bulbs
and corms.

Drought Tolerance with low water potential: The ability to tolerate low water
avalability. Means that plant can continue metabolic processes during period of water
limitation. The mechanism used by species to continue metabolism under low tissue
water potential are different from those used by plants maintaining water potential
high during water limitation. Plants used no: of techniques to maintain metabolic
activity under low water potential
i)Osmotic adjustment:
a. True osmotic adjustment which is also called as TYPE I occurs when actual no: of
osmotically active solute increase in symplasm without any change in proportion
of water in symplasm.
b. Type II osmotic adjustment: This occurs when proportion of water in symplasm
decrease without any change in absolute no: of solute dissolved.

c. Type III osmotic adjustment: It occurs when both the absolute no: of dissolved
solutes changes and symplastic water fraction changes.

Symplast v/s Apoplast volume

When the symplast water fraction decreases relative to apoplastic fraction, solutes are
held in smaller fraction of tissue water. This increase osmotic conc. without require
more solutes & may help in maintaining turgor potential at lower water potential. The
adjustment in symplatic water fraction normally occurs by the construction of new
tissue. The addition of mucilage to apoplast can reduce the symplast water potential.
Eg. Opuntia.

Desiccation tolerance: The lower vascular plants have ability to tolerate complete
desiccation and regain metabolic activity rapidly on rehydration. Lichens, algae & ferns.
In these plants water evaporates from tissues, cytoplasm shrinks result in higher conc.
of cytoplasm and large spaces inside the cell wall. Three criteria must be met by species
of desiccation tolerance
1. The cellular metabolism must be so designed that higher conc. does not affect the
basic structure of cell.
2. There should be few plasmodesmatal connections because cell shrinkage would
break connection.
3. The cell wall must be able to withstand extensive dehydration without losing
structure.

Drought tolerance with high water potential

It occurs when species are able to maintain a metabolism during water limitation
without experiencing low water potential in their tissue.
1) REDUCTION OF WATER LOSS: If a plant can reduce total water use by canopy
soil water will be conserved. In any given set of environment with specific leaf
morphology, there is a theoretical optimum stomatal behavior that maximize the
water use efficiency. The relationship between transpiration and photosynthesis
may be expresses as
Instantaneous change in transpiration
λ = -------------------------------------------------
Instantaneous change in net photosynthesis
When water availability is low λ is low and this promote a mid day depression in
stomatal conductance. This mid day depression improves water use efficiency at low
water availability.

2) ENHANCEMENT OF WATER ACCUMULATION: Species can maintain high water

potential by utilizing large proportion of soil water. Root systems that travel deep
into soil for e.g. phreatophytes (deep root system) tap root species with 5-10 m deep
roots. They exploit water resources unused by most other species. Deep roots acts as
hydraulic lift i.e. when deep roots are saturated with water upper roots are in dry
soil. The water potential gradient between roots and soil pulls water from roots into
soil and increase in the water availability. Some cacti have extensive root system on
upper surface of soil this ensures maximum absorption of rain water, during
drought some cacti cut off connection with their roots because in dry soils extensive
surface of roots will tend to draw water out of plant.
In Tillandsia (Peru) air plant. They absorb water from fog because there is little or no
rainfall in desert area. These are hemi- parasitic species which utilize resources from
host plants.
 FLOODING

It is prolonged period of rainfall or over irrigation combined with poor internal soil
drainage causes flooding leading to soil water logging.

It is detrimental occurrence because of its impact on agricultural and human habitation.

6 % of worlds terrestrial ecosystem commonly experience flooding in agricultural filed
irrigation can cause flooding for short period. Natural flooding by excessive rainfall or
rising river water can cause extensive damage to crops.

Environmental attributes of flooding conditions

The most important detrimental characteristics of flooding ecosystem of plant is oxygen

partial pressure in the root zone. When the soil is flooded the oxygen cannot diffuse in
fast enough through water to compensate for root and soil micro-organisms respiration.
A few cm of soil surface can retain some oxygen i.e. hypoxic in comparison to deeper
layers which has no oxygen.
Now these changes in rhizosphere are important
1. Roots are particularly sensitive to anaerobic conditions.
2. Anaerobic conditions support a different microbial community compared with
aerobic conditions and this severely affects nutrient relation of soil. Anaerobic
bacteria are less efficient than aerobic bacteria in decomposing organic matter.
Therefore flooded soils have large amount of organic matter because anaerobic
bacteria do not decompose as fast as aerobic. The low water potential of flooded
soils also cause a reduced availability of oxidized nutrient such as NO3, all other
nutrient availability is also effected for eg. Fe & S. CO2 & certain organic acid are
reduced to form methane. In most wet lands the thin layer of aerobic condition at
surface of soil results in aerobic and anaerobic interphase. This interphase is
particularly important for transformation of major nutrients such as nitrogen.

In aerobic soils nitrification is the dominant N transformation process, accomplished by

aerobic bacteria. In contrast de-nitrification i.e. conversion of nitrate to nitrous oxide
and di nitrogen (N2) occurs through action of anaerobic bacteria. Ammonia in the
anaerobic sediments moves up into aerobic zone where it is converted to NO3. NO3
diffuse down into anaerobic sediments where it is converted to N or N2O which diffuse
up into atmosphere. The N becomes an important limiting nutrient because N is lost by
nitrification and de-nitrification process set up by aerobic and anaerobic interphase.

EFFECT OF FLOODING
1. On Respiration: The most immediate effect of anaerobic soil conditions on plants
is reduction of respiration in roots. In aerobic respiration 36 molecules of ATP are
produced and in anaerobic respiration only 2 ATP molecules are produced. ATP :
ADP ratio, the conc. of ATP and energy is decreased in root cells under flooded
conditions. Ultra structure and functioning of mitochondria are affected rapidly
after anoxia begins. However after transfer back to aerobic conditions the ATP:ADP
increase rapidly and structure of mitochondria returns to normal. Shortly under low
O2 levels alternate cyanide resistant respiration pathway becomes operative. A high
activity of cyanide resistant respiration pathway may reduce rate of fermentation,
reduction in the rate of ethanol build up and generate enough oxidized NAD to
keep the glycolysis active.

2. Photosynthetic Carbon Balance: Root zone flooding causes closure of stomata in

sensitive plant species because shoot respond to stress in roots & this leads to
reduction of photosynthesis. Also photosynthetic enzymes are inhibited by presence
of H2S (product of anaerobic respi.) carbohydrates translocation is inhibited and
there is build up of inhibitory growth regulators such as ABA & ethylene. There is
complete shut down of photosynthesis therefore anaerobic respiration strongly
increases and carbohydrates are depleted leading to their starvation during
flooding. Carbohydrates starvation is enhanced further in roots because
translocation of carbohydrates from leaves to roots is inhibited during flooded
conditions.

3. Water relation: Flooding with both saline & fresh water effect plant water relation.
Flooding increase atmospheric humidity which along with stomatal closure reduces
transpiration. Stomatal closure is caused by a root signal which is combination of
reduced cytokinins synthesis & transport along with an increased transport of ABA
& ethylene, this induces stomatal closure. There is decrease of permeability of roots
to water, wilting can be a transient respons to flooding, this wilting can lead to death
of root system which was build during anaerobic condition. However ethylene
induces the formation of adventitious roots, these roots develop further hypoxic
zone near soil surface. Stomata may reopen because root function is shifted to newly
developed adventitious roots. However the water conductance of these roots is
lower than aerobic roots because in these conditions extensive aerenchyma is
formed at the expense of water conducting xylem.

4. Nutrient relation: Flooding reduces transpiration and uptake of nutrients through

xylem. Anaerobes are unable to provide adequate ATP for active nutrient
 mechanism thus severly limits nutrient accumulation in flooded conditions. When
rhizosphere O2 conc.falls below the critical oxygen pressure ( the oxygen partial
pressure that decrease during cellular respiration and increase in respiratory
quotient) respiration is inhibited & nutrient uptake slows down. There is rapid set
of chlorosis in sensitive plants after flooding because of reduced nutrient uptake &
iron toxicity. Extended flooding changes the Fe in to soluble forms of ferrous ions,
many micro nutrients are reduced making them unavi. For e.g. S is reduced to
sulphide.

5. Mechanical effects of flooding: In flood plain ecosystem flooding can cause

significant damage to plants by force of rapidly moving water. There can be
extensive siltation which can bury stem, branches & erode leaves.

6. Effect of anaerobic microorganism: Anaerobic microorganism derive their energy

from reduction of NO3 to NO2 or dinitrogen oxide (N2O) and molecular nitrogen.
N2O & nitrogen both these gases are lost to atmosphere and the process is called
denitrification. As the conditions becomes more reducing a ferric ions are reduced to
ferrous ions and because of its greater solubility ferrous can rise to toxic conc. if soil
remains anaerobic for many years. anaerobic microorganism may also reduce
sulphate ions to hydrogen sulphide which is respiratory poison. Unpleasant odour
of water logged soils is due to bacterial metabolites such as acetic acid, butyric acid
along with suplhur compound in water logged soils.

ADAPTATIONS FOR FLOODING

1. Vigorous growth in flooding tolerant plant: In wet land species for e.g. water lily
submergence stem traps endogenous ethylene & this hormone stimulates cell
elongation of petiole extending it quickly to the water surface so that leaf reaches
air. Internodes of deep water rise to trap ethylene. In case of pond, weed
Potamageton which is aquatic monocot plant, stem elongation is insensitive to
ethylene instead elongation is promoted even under anaerobic conditions by
acidification of surrounding water caused by accumulation of respiratory CO2.

2. Development of interconnected gas filled channels: Plants which are acclimated

to wet conditions the stem & root develop longitudinal interconnected gas filled
channels that provides a low resistance pathway for movement of O2 & other
 gases. In many wet land species such as rice cells are separated by prominent gas
filled spaces called arenchyma that develop in roots independent of environment
stimuli. In few non wet land plants both monocots & dicots O2 deficiency
induces the formation of aerenchyma in the base of stem of newly developed
roots. In root tips of maize hypoxia stimulate action of ACC synthase & ACC
oxidase. Thus causing quicker formation of ethylene. Ethylene causes death &
disintegration of cells in root cortex. The space these cells formly occupied
provides gas filled gaps that facilitates movement of O2. Ethylene signal, death
in selective, cell not destined to die in roots remains unaffected or increase in
cytosolic Ca conc. promotes cell death under non inducing conditions.
Alternately conditions that lower Ca conc. block cell death in hypoxic roots that
normally form aerenchyma. This ethylene dependent cell death in response to
hypoxia is an example of programmed cell death. Some plants can tolerate
hypoxia several weeks / months before developing aerenchyma. These include
embryo & coleoptiles of rice plants. Deep water rice can response to partial
submergence with an increase in internodal growth. Its survival depends upon
its ability to keep parts of its foliage above water. Ethylene have been shown to
be responsible for enhanced internodal growth of tip of watered rice, in rice a
low O2 tension in roots provide a stimulation of ACC synthase. It has also been
suggested that ethylene sensitizes to cell gibberellins which promotes rapid
growth of cells.

TEMPERATURE

Temperature along with light & water is one of the most critical factors in physical
environment of plants because plants unlike homoeothermic animals are not able to
maintain their tissue at constant temperature. Environmental Temperature exerts
profound influence on cellular metabolism & also plant growth & there geographical
distribution. The temp. at which biological process occur is generally limited by
freezing point of water at lower side & irreversible denaturation of proteins on higher
side. Temp. curve for growth of an organism represents composite of temp. curve for
photosynthesis, respiration and other critical metabolic processes. Growth curve
exhibits 3 cardinal points maximum, minimum & optimum. Plants & related organism
may be classified according to their ability to with stand temperature.

1. Psychrophiles: optimal range of temp. is between 0-10 °C and primarily includes

algae , fungi and bacteria.
 2. Mesophiles: optimum range of temp lies between 10-30 °C and most higher plants
fall in this category.
3. Thermophiles: optimum growth is between 35-65°C. many of cyanobacteria fall in
category.

Dehydrated organisms & organs such as resurrection plants & dry seeds with moisture
content as low as 5% are able to with stand much broader range of temperature for
extended period of time. Plants in nature are subjected to a complex mosaic of
fluctuating air & soil temperature regimes. Air temp. fluctuates widely depending upon
time of day, cloud cover, season & other factors. Soil temp. varies with soil structure,
organic content and other physiological characters such as slope & direction it faces
with respect to sun.

IN NATURE HOW DO PLANTS ADAPT TO TEMPERATURE

Many CAM & succulent higher plants are adapted to high temperature and can tolerate
tissue temperature of 60-65°C. CAM plants keep their stomata close during the day
therefore they cannot cool by transpiration. Instead they dissipate the heat from
incident solar radiation by reemission of long wavelength radiations ( infra red) & lose
heat by conduction and convection. On the other hand typical non irrigated C3 & C4
plants rely on transpirational cooling to lower leaf temperature. In these plants the leaf
temperature can readily lie above 4-5 above ambient temp. in bright sunlight, near
midday when soil water deficit causes partial stomatal closure or when high relative
humidity reduces the potential for evaporative cooling. Increase in leaf temperature
during day can be pronounced in plants forms in arid & semi arid regions experiencing
drought & high irradiance from sunshine. Heat stress is also a potential danger to green
houses where low air speed & high humidity decrease the rate of leaf cooling. A
moderate degree of heat stress slows growth of whole plants. Some irrigated crops such
as cotton use transpirational cooling to desiccate heat. In irrigated cotton enhanced
transpiration cooling is associate with higher agronomic yields.

PHYSIOLOGICAL EFFECTS OF HIGH TEMPERATURE STRESS

1. On photosynthesis & respiration: Both photosynthesis & respiration are

inhibited by high temperature, but as the temperature increase the
photosynthetic rate drops before the respiratory rates. The temperature at which
amount of CO2 is fixed by photosynthesis is equal to amount of CO2 released by
respiration in a given time interval is called temp. compensation point. At temp.
 above temp. comp. point , photosynthesis cannot replace the carbon used as
substrate for respiration as a result carbohydrate reserve decline and fruits &
vegetables loose sweetness. This imbalance between photosynthesis &
respiration is one of the main reasons for deleterious effect of high temp. In a
same plant temp. compensation point is usually lower for shade leaves and for
sun leaves. Enhanced respiration rate relative to photosynthesis rate at high
temperature are more detrimental in C3 plants than C4 and CAM because the
rate of both dark respiration and photorespiration increases in C3 plants at
higher temperature.

2. Membrane stability: The stability of various cellular membranes is important

during high temperature stress, excessive fluidity of membrane at high
temperature is correlated with loss of physiological functions. In oleander
(Nerium) acclimation to higher temperature is associated with greater degree of
saturation of fatty acids in membrane lipids which makes membrane less fluid.
At high temp. there is a decrease in strength of hydrogen bond, electrostatic
interaction between polar groups of proteins with in aqueous phase of
membrane. High temp. thus modify membrane composition & structure and
cause linkage of ions. Membrane disruption also causes inhibition of processes
photosynthesis & respiration that depends upon the activity of membrane
associated as electron carries & enzymes. Photosynthetic enzymes Rubisco,
NADP, G3P, PEP carboxylase are less stable at high temp. however the temp. at
which these enzymes began to denature & lose activity is distinctly higher than
the temp at which photosynthesis begins to decline so this suggests that early
stages of heat injury to photosynthesis are more directly related to change in
membrane properties & to uncoupling of energy transfer mechanism in
chloroplast than to a general denaturation of proteins.

ADAPTATIONS TO HIGH TEMPERATURE:

In environment with intense solar radiation & high temp. plants avoid excessive
heating of their leaves by decreasing the absorption of solar radiations. Both drought
resistance & heat resistance depends upon some adaptation i.e. presence of reflective
hair on leaf surface, leaf waxes, leaf rolling, vertical leaf orientation & growth of small
highly dissected leaves to minimize the boundary layer thickness and maximize heat
loss by convection & conductance.

PRODUCTION OF HEAT SHOCK PROTEINS (HSP) : In response to rise in temp.

plants produce unique set of proteins referred to as heat shock proteins. Most of heat
shock proteins function to help cell withstand heat stress by acting as molecular
chaperone (escort). Heat stress may cause many cell proteins that function as enzymes
of structure components to become unfolded or misfolded there by leading to loss of
proper enzyme structure & activity. Such misfolded proteins often precipitate therefore
creating serious problems in cell. HSP act as molecule chaperon & serves to attain
proper folding of misfolded aggregated proteins & to prevent misfolding of proteins.
Thus facilitate proper cell functioning at stressful temp. Plants & most other organism
make HSP of different sizes in response to temp. increase. The molecular mass of HSP
ranges from 15 to 104 KDa and they can be grouped in 5 classes on the basis of size.
Different HSP are located in nuclei, mitochondria, endoplasmic reticulum, cytosole,
chloroplast. Member of HSP 60, HSP 70, HSP 90 & 100 groups acts as molecular
chaperone involving ATP dependent stabilization & folding proteins & assembly of
oligomeric proteins. Some HSP assist in polypeptide transportation across the
membrane into cellular compartments. Low molecular weight i.e. 15-30 KDa is more
abundant in higher plants than in any other organism. The different classes of 15-30 mol
wt. of HSP in plants are distributed in the cytosole, chloroplast, endoplasmic reticulum,
mitochondria, but the function of these small HSP is not under stood. Cells that have
been induced to synthesize HSP shows improved thermal tolerance & can tolerate
exposure to temp that are otherwise lethal. Some of HSP’s are not unique to high
temperature stress, they are also induced by environmental stress conditions such as
water deficit, ABA treatment, wounding, low temp. & salinity. Thus cells previously
expose to one stress may gain cross protection against another stress for e.g. in tomato
in which heat shock (48hrs ,38° C) has been observed to promote HSP accumulation &
protect cells from chilling at 2 °C for 21 days.
 LOW TEMPERATURE STRESS(CHILLING AND FREEZING)
Chilling temperature are too low for normal growth but not low enough for ice to form.
Typically tropical & subtropical species a susceptible to chilling injury, among the crops
maize, beans, rice, tomato, cucumber, sweet potato & cotton are chilling sensitive. When
the plants growing at relative warm temperature 25-35 °C are cooled to 10-15°C, chilling
occurs. Freezing injury on the other hand occurs at temperature below freezing point of
water.
Physiological changes in respiration due to chilling injury: Leaves from plant injured by
chilling show inhibition of photosynthesis, slower carbohydrates metabolism, lower
respiration rate, inhibition of protein synthesis & increased degradation of existing
proteins. All these responses depends upon a common primary mechanism involving
loss of membrane function during chilling for e.g. solutes leak from the leaves of
chilling sensitive plants when floated in water at 0 °C, but not from those of chilling
resistant plants. Plant membrane consist of lipid bi layer which is interspersed with
protein & sterols. The physical properties of lipids influence the activity of integral
membrane protein including H+-ATPases carrier & channel forming proteins that
regulates transport of ions & other solutes as well as transportation of enzyme on which
metabolism depends
In chilling sensitive plants lipid bi layer have a high %age of saturated fatty acid chain
& membranes with this composition tend to solidify in semi crystalline state at
temperature above 0°C. As membrane becomes less fluid these protein components can
no longer function normally. The result is inhibition of H+-ATPases activity or solutes
transport into & out of cell or energy transduction & of enzyme dependent metabolism.
In addition chilling sensitive leaves exposed to high photon fluxes & at chilling
temperature photosynthesis is inhibited leading to cause damage to photosynthetic
machinery. On the other hand membrane lipids from chilling resistant plants have
greater proportion of unsaturated fatty acids than from chilling sensitive plants &
during acclimation to cool temperature the activity of desaturated enzymes increases
and the proportion of unsaturated lipids rises. This modification lowers the
temperature at which membrane lipids begin a gradual change from fluid to semi
crystalline state & allows the membrane to remain fluid at low temperature. Thus
desaturation of fatty acids provides some protection against damage from chilling. The
ability to tolerate freezing temperature under natural condition varies among tissues,
seeds other partly dehydrated tissue & fungal spores can be kept indefinitely at
temperature near absolute zero, indicating that very low temperatures are not
intrinsically harmful. Fully hydrated vegetative cell also retain viability if they are
cooled very quickly to avoid formation of large slow growing ice crystals that will
puncture & destroy sub cellular structure. Ice crystals that are formed during very rapid
freezing are too small to cause mechanical damage. Conversely rapid warming of
frozen tissues is required to prevent growth of small ice crystals into crystals of
damaging size or to prevent the loss of water vapour by sublimation both of which to
be placed at intermediate temperature of -100 to -10 °C. under natural conditions
cooling of intact multicellular plant organ is never fast enough to limit crystal formation
fully hydrated cells to only small harmless ice crystal. Ice usually forms Ist with in
intercellular spaces & in xylem vessel. This ice formation is not lethal to hardy plants &
tissues recover fully when warmed. However when plants are exposed to freezing
temperature for an extended period the growth of extra cellular ice crystal result in
movement of water from protoplast to the extracellular ice causing excessive
dehydration. During rapid freezing the protoplast including vacuole super cools i.e
cellular water remain liquid even at temperature several degree below at its freezing
point. Several hundred molecules are needed to form ice crystals. The process where by
these hundred of water molecules start to form stable ice crystal is called nucleation & it
strongly depends on properties of involved surface. Some large polysaccharides &
proteins facilitates ice crystal formation & are called nucleators. Several specialized
plant proteins may help to limit growth of ice crystals, these are called anti freeze
proteins. The anti freeze proteins are induced by cold temperature & they bind to
surface of ice crystal to prevent or slow further crystal growth. In Rye leaves anti freeze
proteins are located in epidermal cells & the cells surrounding intercellular spaces
where they can inhibit growth of extra cellular ice. Sugars & some cold induced
proteins have cryo-protective effects. They stabilize protein & membrane during
rehydration induced by low temperature in winter wheat. The greater the sucrose conc.
greater is the freezing tolerance. Sucrose predominately among soluble sugars
associated with freezing tolerance. During cold acclimation of winter cereals, soluble
sugars accumulate in cell wall where they may help to restrict growth of ice. Cryo-
protective lipoproteins have been isolated from cold acclimated cabbage. In invitro
studies the proteins protect thylakoid isolated from non acclimated spinach against
damage from freezing and thawing.

ROLE OF COLD RESISTANCE IN WOODY PLANTS

In dormant state, the woody plants are extremely resistant to low temperature.
Resistance is determined partly by previous acclimation to cold but the genetic play
important role in determining the degree of tolerance to low temperature. Native
species of Prunus from Northern America are hardier after acclimation than those from
milder climatic. Under natural conditions woody species acclimate to cold in 2 stages
1. In the 1st stage hardening is induced in the early autumn by exposure to short
days and non-freezing chilling temperature both of which combine to stop
growth. A diffuseable factor ABA which promotes acclimation moves in phloem
from leaves to over wintering stems & may be responsible for change. During
these period woody species also with draw water from xylem vessels thereby
preventing the stem from splitting in response to expansion from water during
freezing. Cells from 1st stage of acclimation can survive temperature well below
0° C but they are not fully hardened.
2. In 2nd stage direct exposure to freezing is stimulated but no known translocatable
factor can confer the hardening resulting from exposure to freezing. When fully
harden, the cells control exposure to temperature of -50 to -100 °C.

Role of ABA in acclimation

In seedling from alpha alpha tolerate to freezing at -10 °C is improved by previous

exposure to cold at 4 °C or by treatment with exogenous ABA without exposure to
cold. These treatments cause changes in pattern of newly synthesized protein. Some
of changes are unique to particular treatment i.e. cold or ABA but some of newly
synthesized proteins induced by cold appearance to be the same as those induced by
ABA. Plants develop freeze tolerance at non acclimated temperatures when treated
with exogenous ABA. Many of the genes or proteins expressed at low temperature
or under water deficit are also induced by ABA under non acclimated conditions.
All these findings support role of ABA in freezing tolerance.