Unit 16 Plant Stress

UNIT 16
PLANT STRESS

Structure
16.1 Introduction 16.5 Plant Responses to
Specific Stress Conditions
Objectives
Water and Osmotic Stress
16.2 What is stress?
Salinity
16.3 Nature of Stress
Pollutant Stress
Physical stress
Temperature Stress
Chemical stress
Stress by Infection and
Biological stress
Wounding
16.4 Ways to Adapt to Stress
16.6 Future Prospects
Changes at Cellular and
16.7 Summary
Molecular Level
16.8 Terminal Questions
Biochemical Alterations
16.9 Answers
Changes in Plant Morphology
and Behaviour

Alternate Metabolic Pathways

16.1 INTRODUCTION
The previous units have provided you information about the various essential
physiological processes in plants. These include photosynthesis, respiration,
transpiration, translocation of solutes and assimilation of nutrients such as
carbon, nitrogen, sulphur. Plants get exposed to environmental variations.
These changes bring alterations in the growth and development of plants by
affecting various biochemical and physiological processes. Exposure to abiotic
and biotic stresses such as alteration in temperature, water availability, soil
pH, salinity, alkalinity, heavy metals, pathogen infestation, infection and injury
brings a change in the metabolic status of the plants. Exposure of plants to
environmental conditions or stress triggers various metabolic responses in
plants. The present unit introduces you to the concept of abiotic and biotic
stress in plants and provides detailed information about plant response to
stress. 213
Block 4 Nitrogen Metabolism and Plant Growth Regulators

Objectives
After studying this unit you would be able to:

define stress and differentiate between abiotic and biotic stress to

which plants gets exposed;

describe changes in morphological, physiological and metabolic

behaviour of plants in response to stress;

explain plant response to water and osmotic stress, temperature

stress;

discuss plant response to pollution; and

explain about plant response to infection and wounding.

16.2 What is stress?

Any change in the surrounding environment that affects the functioning of
plants and alters the homeostasis is defined as biological stress. A sudden
transition in the optimal environmental conditions which disturbs the
homeostasis in plants and induces adverse effects on their physiology is
referred as plant stress. Plant species get exposed to fluctuations in
temperature (chilling, freezing and high temperatures), water availability,
changes in the soil conditions such as salinity, heavy metals etc. Plants have
an optimum requirement for various environmental variables and show
susceptibility to any alteration. These changes impose stress in plants which
include delay or alteration in growth and development, effects on productivity
and in extreme cases, death.

Let us recall what happens within the natural communities occupying the same
habitat. The relative location of two plants may place them under differing
conditions with respect to a given environmental factor such as light. The top
cover of a rainforest, for example, consists of relatively tall trees and receives
maximal irradiance while the floor dwellers manage with sunflecks.

214 Fig. 16.1: Graphic representation of yield gap.

Unit 16 Plant Stress
What will happen if we artificially shadow the outer cover plants and illuminate
the forest floor? Being a deviation from the natural situation, this is likely to
have adverse effects on growth of plants. Fig. 16.1 shows the yield gap due to
biological and environmental constraints. Thus, any deviations from optimal
environmental conditions usually with adverse effects are considered as
stress. This in turn affects the yield of useful agricultural products

We must, however, realise that in no habitat, all conditions - temperature, light,

availability of water, nutrient supply and soil characteristics can be controlled
and fixed to optimal level for any species. But since the species must survive,
it must adapt itself to the deviations in the environmental condition(s). This
could be achieved either by breeding crop varieties tolerant to stress or by
offering conditions that help the plants to withstand the stress. For example, if
there is a deficit of some nutrient one could try to supplement the same. Plants
capable of adapting themselves to changes in environmental conditions
perform the best under stressful conditions.

16.3 NATURE OF STRESS

As mentioned in the preceding section, stress can be considered as any
deviation in environmental condition from optimal for the overall performance'
of the plant species under reference. If you recall the environmental factors,
you can tell the various kinds of stress that plants may be subjected to. A
broad outline on the nature of possible environmental stress is illustrated in
Fig. 16.2.

Fig.16.2: Nature of Environmental Stress Conditions.

16.3.1 Physical Stress

Physical stress originates from environmental conditions such as:

i) Temperature : We are familiar with the plants and other organisms that
live at temperatures close to the temperature range in which we are 215
Block 4 Nitrogen Metabolism and Plant Growth Regulators
adapted to live i.e., 15° to 45° C. However, we know that there is life
below 0° C in the arctic and above 90° C in the sulphur springs. In the
subtropical zones, plants face stress -when they get exposed to freezing
temperatures during the winters while in the deserts of the tropics the
native plants withstand over 55° C during summer. High temperature can
be inhibitory for photosynthesis (Fig. 16.3).

Fig. 16.3: Effect of light intensity and temperature on the rate of photosynthesis.

ii) Osmotic strength of the fluids in immediate surrounding: The

availability of soluble mineral salts varies widely from habitat to habitat.
In fact, a big chunk of land in our country has been classified as
'wasteland' because of high salinity of the soil cover or the water
leachates. Common crop cultivars cannot be grown in such
environments.

iii) Photosynthetically active radiation: This is usually in direct proportion

with the incident solar radiation. You know that tropics are probably the
best illuminated part of the globe. Also, variation in day length is minimal
in this zone. No wonder, light-dependent life has attained maximal
density in the tropics. Within a dense population of plants, different
organisms may have different degrees of exposure. However, if a plant
is exposed to increasing light intensity it shows a corresponding increase
in the rate of photosynthesis upto a certain point, beyond which further
increase in light causes either no change in the rate or causes an
inhibition (Fig. 16.3). Thus, each plant species shows a characteristic
photosaturation and/or photoinhibition of photosynthesis. In other words,
each species has its own optimum with respect to light intensity. Plants
get stressed if they get exposed to intensity above or below the
optimum.

16.3.2 Chemical Stress

Survival of cells is dependent on carrying out of a set of chemical reactions
(metabolic reactions) in a particular order. This results in a net gain in mass as
216 well as energy in a growing cell. Even for dormant cells as in dormant buds
Unit 16 Plant Stress
and seeds, where there may not be any net gain in mass, a certain amount of
energy must be spent to maintain them in a viable state. Various catabolic and
anabolic pathways operate according to the availability of chemical
constituents present in the cytoplasm. This may in turn be influenced by the
composition of the extracellular environment with respect to the following:

i) Nature of the solutes: The acidic or basic reaction of soil and water of a
particular habitat reflects its geochemical history beginning with its
formation and subsequent interactions with other constituents of the
earth up to its current chemical activities. Mineral deposits in their oxide
form ('Bhashma') are usually basic in their reaction. The reaction of
chloride, sulphate and nitrate is acidic or neutral depending upon the
nature of the conjugate ions. You can visualise the reaction of a salt
through its acidic and basic radicals. On a natural course, one would
expect the neutralisation reactions and consequently change in the
character of the habitat towards neutral. However, there are soils which
are very high and others very low in pH and certain plant species survive
in such soils.

ii) Mineral composition: The living systems make use of several mineral
ions that they might have encountered at the very origin or during
evolution, particularly for transformation of matter involving proteins and
nucleic acids. These elements continue to remain essential requirements
for life. You have already learnt in Unit 12 that nitrogen, phosphorus,
calcium, potassium and magnesium are familiar major requirements for
plant growth besides carbon, hydrogen and oxygen. Apart from these,
many other micronutrients such as manganese, iron, zinc, cobalt and
molybdenum are required for healthy plant growth in much smaller
quantities. Availability of these elements in the environment in quantities
smaller than required cause’s deficiency symptoms while a surplus may
cause toxicity leading to stunted growth, necrosis, abnormal
development of vegetative and reproductive parts.

iii) Atmospheric imbalances: At least two components of the environment,

oxygen and carbon dioxide that plants require are predominantly in the
gaseous form. As you have learnt, atmospheric nitrogen can be used by
nitrogen-fixing bacteria, some prokaryotic algae and plant-bacterial
symbiont. The biological cycling of these gases keeps their overall
availability buffered within a reasonably stable range. Yet, intensive
industrial activities that involve emission of one of these gases or high
levels of the oxides of carbon, nitrogen and sulphur have visible effects
on the performance of most plant species.

16.3.3 Biological Stress

Since in nature, the various organisms do not live in complete isolation from
others, stress to a plant species might also be caused by what other
organisms in the community require and consume. The following situations
exemplify biological stress.

i) Population density: You are aware of what might follow an uncontrolled

growth of human population. There will be competition for common 217
Block 4 Nitrogen Metabolism and Plant Growth Regulators
consumables and for space but this is not merely a human problem. Too
thick population densities of plants can resist the competition for
common nutrients, water and photosynthetically active radiation (PAR).
The effect can be a self inhibition or inhibition of other plant species
growing in the same environment. That is one of the reasons why weeds
are removed from croplands.

ii) Parasites: Many insects and microorganisms can feed upon tissues and
saps of living plants. Hence plants must be protected against such
parasites either by expressing evasive devices such as inhibitors and
toxins against the enzymes of parasites or by developing preventive
morphological and biochemical alterations that keep the parasite away
from aggregating near the plant.

iii) Symbiotic interaction: Presence of symbiotic microorganism can result

in differential growth stimulation of those plants which can recognise the
beneficial symbionts and utilise their presence in the environment. This
interaction is a mutual one. For example in Rhizobium-legume
symbiosis, the rhizobium causes a drain on the carbon fixed by the plant
(refer Unit 13) while the plant gains access to the nitrogen fixed by the
bacteria.

SAQ 1
What is the nature of stress faced by the following species if the change
mentioned below is brought about in their environment?

a) A plant species growing in Manali is brought to Jaipur during summer.

b) A sun-loving plant gets shaded by other plants.

c) A wheat crop gets flooded due to heavy rains for several days.

d) Trees growing near the roadway are subjected to heavy traffic.

e) Plants acclimatised to grow near fresh water are grown in coastal areas
of Goa.

16.4 WAYS TO ADAPT TO STRESS

Plant stress responses are dynamic and involve various regulatory pathways.
These include physiological, morphological adaptations and adjustment of
metabolism. Alterations in the physiology or morphology help the plants to
survive the changing environmental conditions (Fig 16.4). Plants respond to
stress by undergoing change in metabolic functions, morphological features
and physiological features. These responses are referred as stress
responses. In salt sensitive plants (glycophytes), a number of physiological
responses get stimulated. These response help plants to cope salinity stress.
This includes SOS pathway (known as salt overlay signaling) that leads to
enhanced efflux of sodium (Na) ions and reduction in toxicity induced by
218 salinity.
Unit 16 Plant Stress

Environmental stress

Abiotic Biotic

Response

Resistance Susceptibility Avoidance

Survival
Survival Death

Fig. 16.4: Effect of stress on plants.

Stress responses require no genetic modification, hence are referred to as

phenotypic plasticity. In phenotypic plasticity, the plants respond to changes
in environmental conditions via changes in morphology and physiology. The
changes are reversible. The ability of the plant to survive and establish in a
given environment is related to balance between the genetic adaptation and
phenotypic plasticity

Abiotic stress produces primary and secondary alterations in plants. Stress

primarily causes reduction in hydrostatic pressure (turgor pressure) which
leads to cellular dehydration. The change in cell hydration potential affects the
physical and biochemical properties of cell. The secondary effects of stress
include reduction in metabolic activities, production of reactive oxygen species
(oxidative stress) and disruption of cellular integrity, ultimately death. Stress
factors such as increase in temperature and exposure to ionizing radiation, UV
light and environmental pollutants leads to generation of reactive oxygen
species (ROS) which causes oxidative stress. The most common ROS formed
include superoxide radical (O2.-), singlet oxygen (1O2), hydrogen peroxide
(H2O2) and hydroxyl radicals (OH.).

Oxidative stress is caused when the ROS are produced and accumulated in
high amounts in a cell or tissues than those that can tackled by the biological
system. Oxidative stress leads to inhibition of major metabolic functions in
plants. These include inhibition of photosynthesis and respiration. This results
from down regulation or disruption of electron transport chains (ETC) in
chloroplasts and mitochondria. Abiotic stresses such as water deficit, salinity
and freezing disrupt cellular structures and impair physiological processes in
plants. The damage to cellular structures leads to reduction in growth, fertility
and cause premature senescence. The cell membranes become disorganized
as the proteins get deactivated or denatured. 219
Block 4 Nitrogen Metabolism and Plant Growth Regulators
Box16.1: Tolerant plants

Two major strategies of combating stress have been recognized in plants. These are
avoidance and tolerance strategies. Stress avoidance includes a various protective
mechanisms that delay or prevent the negative effect of stress on a plant. Stress
tolerance is the potential of a plant to acclimate to a stressful condition. Many plants
have the capacity to tolerate stress. These are referred to as stress
resistant/tolerant plants (Fig. 16.5). These plants exhibit the capacity to adjust or
acclimatize to stress conditions. The plants ability to resist or tolerate stress depends
on their genetic capacity to adjust/overcome stress and establish a new homeostatic
state over a period of time. Some plants escape the stress. These include
ephemerals or short lived plants. They germinate fast, grow and flower quickly. They
complete their life cycle during a period of adequate moisture and form dormant
seeds before the onset of dry conditions. Many arctic plants complete their life cycle
during the short arctic summer and survive during winter in the form of seeds.
Ephemerals never get exposed to stress because they survive the stress by
avoidance. Plants of alfa alfa (Medicago sativa) survive dry habitats by sending their
roots deep into the soil near the water table. In this way the plant ensures survival
under conditions of drought. Other plants develop fleshy leaves that store water, thick
cuticles and pubescence (leaf hairs) to reduce transpiration such as Bryophyllum,
Opuntia etc.

Fig 16.5: Medicago sativa and Opuntia.

16.4.1 Changes at Cellular and Molecular Level

Responses to environmental stresses occur at all levels of organization. The
changes can be at cellular or genetic level.

Cellular level - At this level, plants tackle stress by bringing modifications in

the structure and architecture of the cell wall and membrane, cell cycle and
cell division. Plants bring alteration in the metabolic events. These include
production of compatible solutes such as proline, raffinose, glycine betaine.
These solutes are the organic compounds that are osmotically active in the
cell and play a role in stabilization of proteins, membranes and organelles.
These compounds also contribute to maintenance of cell osmotic status by
removing ROS and re-establishment of redox balance.

Molecular level - At this level, gene expression gets modified due to stress.
Stress-inducible genes i.e. genes that are involved in protection of cell from
stress via synthesis of osmoprotectants, detoxifying enzymes, transporters
and those which encode regulatory proteins such as transcription factors,
220 protein kinases, and phosphatases get induced.
Unit 16 Plant Stress

SAQ 2
Define the following terms:

a) Stress

b) Tolerant plants

c) Ephemerals

d) Phenotypic plasticity

16.4.2 Biochemical Alterations

The plants exposed to stress depict alterations in their metabolic processes.
Drought (water deficit), salinity and low temperature stress leads to change in
osmotic/water potential resulting in loss of turgor in plant cells. The plants
adjust to the condition of loss of turgor by accumulating solutes. The osmotic
adjustment refers to the capacity of the plant cell to accumulate solutes which
help in maintaining the osmotic potential during stress conditions.

High levels of sodium (Na) or chloride (Cl) ions exert detrimental effects in
plants. Accumulation of ions (Na and Cl) in vacuoles (vacuolar
compartmentalization) facilitates osmotic adjustment in halophytes growing in
saline conditions.

Compatible solutes (osmolytes) help in tackling stress. These include a group

of chemically diverse uncharged, polar, and soluble organic compounds and
do not interfere with the cellular metabolism even at high concentrations. They
mainly include proline, glycine betaine, sugar, and polyols. They act as source
of carbon and nitrogen to the cell under normal conditions. Some plants show
accumulation of amino acids on exposure to abiotic stress. This results from
increase in amino acid synthesis or stress-induced protein breakdown.

Carbohydrates such as starch and fructans show accumulation under stress

conditions. They act as storage substances and are used as energy source
during stress. Fructans have high water solubility and possess resistance to
crystallization at freezing temperatures. They stabilize membranes and
indirectly contribute to osmotic adjustment by the release of hexose sugars.
The roles played by these carbohydrates in stress mitigation are
osmoprotection, carbon storage and scavenging of reactive oxygen species.
Sugars function as osmolytes and help in maintaining turgor of cell and protect
membranes and proteins from damage. Salt and drought stress generally
leads to a depletion of starch content but accumulation of soluble sugars in
leaves.

16.4.3 Changes in Plant Morphology and Behaviour

The plants adapt to extreme environmental conditions through modification in
their life cycles. Desert plants have short life cycles and complete their life
cycle when the water is available. The deciduous trees of the temperate zone
shed their leaves before winter so that leaf tissue does not get damaged by
low temperature. Besides these the growth habits of plants also contribute to 221
Block 4 Nitrogen Metabolism and Plant Growth Regulators
tolerance in plants. Studies have shown that plants which flower for long
period of time are more tolerant to changing environmental conditions than
those flowering for short periods.

The leaves of some plants develop certain characters that help them to
escape extreme environmental changes. These include changes in leaf area,
leaf orientation, thickness of leaf, presence of trichomes and cuticle. Large leaf
area provides conditions for production of more photosynthates under stress
conditions. Large surface area proves advantageous for leaf cooling as more
water can be lost via evaporation, though excessive loss of water results in
dehydration.

Plants growing under stress conditions generally reduce their leaf area by
restricting cell division and expansion, altering leaf shapes and initiating
senescence or abscission of leaves. The cell division and expansion of leaf
gets restricted under conditions of water scarcity and salinity. Several
signaling mechanisms slow down or stop cell cycle thereby limiting growth.
Water deficit reduces turgor which affects cell expansion and reduces leaf
expansion. The smaller leaf area reduces transpiration resulting in effective
conservation of water. The thin film of air at the surface of the leaf (boundary
layer) permits the transfer of heat from the leaf to the air. It helps the leaves to
maintain surface temperature close to that of air so that transpiration is
reduced and overheating can be prevented.

Alteration in leaf shape is another means by which plants overcome stress.

Under conditions of water, heat, salinity stress, leaves become narrower or
develop deeper lobes. This leads to reduced leaf area and prevents loss of
water caused due to excessive heating. In some plants, leaf abscission occurs
under conditions of water deficit and reduced leaf area prevents loss of water
effectively. The plants adapted to drought show leaf abscission. Abscission
results from the enhanced synthesis and responsiveness of the plant hormone
ethylene. Some plants particularly crops such as corn, sorghum are able to
maintain green leaf area during grain maturation stage under stress
conditions. The retention of photosynthetically active leaves under stress is
known as stay green. Some varieties maintain green stem and leaves even
during drought conditions. The delay in senescence has been noted in these
varieties.

The change in orientation also allows greater light absorption by leaves. Under
conditions of high temperature and/or soil water deficits, plants can alter their
leaves to avoid excessive heating. Leaves of some plants orient themselves
away from sunlight to protect themselves from overheating. These leaves are
called paraheliotropic. Some leaves gain energy by orienting themselves
perpendicular to the sunlight is called as diaheliotropic. Wilting and leaf rolling
also alters the absorption of light. Wilting changes the angle of the leaf while
leaf rolling minimizes area exposed to sun.

The presence of trichomes on the surface of leaves help in keeping the leaf
surface cool. The presence of densely packed hairs on the leaf surface
reflects light. These hairs give leaf a silvery appearance. The cuticles (a layer
of waxes and hydrocarbons present on the cell wall of epidermis) present on
the leaf surface also reflect light thereby reducing heat. This layer also restricts
diffusion of water, gases and entry of pathogens. Plants exposed to water
222 stress develop thick cuticle to prevent water loss through transpiration.
Unit 16 Plant Stress
Water stress affects the development of roots and shoots. It has been
speculated that the shoot tends to grow until water uptake by the roots
becomes limiting for further growth. In contrast the roots tend to grow until the
demand for photosynthate from the shoot exceeds the supply. The functional
balanced gets disturbed during water stress. When water supply becomes
limiting, leaf expansion is reduced and a greater proportion of the plant
assimilates can be allocated to the root system where they can support root
growth. As the water deficit progresses the upper layers of the soil dry but the
roots proliferate into deeper, moist soil. This change in root architecture is
considered as defensive strategy against drought. Enhanced root growth into
the deeper soil requires allocation of photosynthates to the growing root tips.

SAQ 3
a) Match the statements from the column A with the correct options from
column B.

Column A Column B

i) salt sensitive plants a) water deficit or salinity

ii) dehydration b) osmolyte

iii) results in loss of cell turgor c) trichomes

iv) stabilizes proteins and d) ephemerals

membranes

v) present on the surface of leaves e) glycophytes

vi) short lived plants f) Reduction in water content

b) Differentiate between:

i) abiotic and biotic stress

ii) paraheliotropic and diaheliotropic leaves

16.4.4 Alternate Metabolic Pathways

The metabolic pathways change in response to stress conditions. Water deficit
decreases photosynthesis and consumption of assimilates in the expanding
leaves. The decreased water potential inhibits movement of assimilates and
less amount of photosynthates get exported from leaves. The ability of plants
to continue translocating assimilates under drought conditions is considered
as a key factor for plant resistance.

Some plants such as C4 and CAM plants change their mode of

photosynthesis. In CAM plants (succulents such as cacti), stomata open at
night and close during the day. This prevents loss of water through
transpiration and increases water use efficiency. The change in CAM mode of
photosynthesis has been noted in plants exposed to water deficit or salinity.
This adaptation allows plant to acclimate to arid environments. 223
Block 4 Nitrogen Metabolism and Plant Growth Regulators
During flooding, oxygen levels are low (hypoxia) and roots, stem of the plants
develop interconnected gas filled channels with in cells (aerenchyma) that
provide resistance pathway for the movement of oxygen and other gases. The
gases enter stomata or lenticels in woody stem and roots. They move by
molecular diffusion or by convection driven by small pressure gradients. In
many plants growing in wet habitats, the root cells develop into aerenchyma
cells. When the oxygen supply is insufficient for respiration, roots begin to
ferment pyruvate to lactate through the action of lactate dehydrogenase. The
production of lactate lowers the intracellular pH inhibiting the lactate
dehydrogenase and activating pyruvate decarboxylase. The change in the
enzyme activity leads to ethanol production. Hence during fermentation, 2
moles of ATP are formed per mole of hexose sugar catabolized in comparison
to 36 moles of ATP produced from each hexose catabolized in aerobic
respiration. Thus injury to the root metabolism by oxygen deficiency leads to
lack of ATP which affects essential metabolic processes.

16.5 PLANT REPONSES TO STRESS

CONDITIONS
16.5.1 Water and Osmotic Stress
The condition that arises due to scarcity or excess of water is known as water
stress. Water scarcity results in drought while excess of water leads to
flooding. Water makes a large proportion of the cell volume and is used for
expansion and metabolic processes. Flooding stress leads to anaerobic
conditions i.e. oxygen stress. The decreased supply of oxygen limits
respiration, nutrient uptake and other root functions. During drought
conditions, the plant cells lose water; they shrink because of collapse of cell
wall. The damage to root hairs affects the absorption capacity. The outer
layers of root cortex get covered with suberin, a water impermeable lipid that
increases resistance to water flow in the root. The resistance to water flow
also results from breakage of water columns within xylem under tension.

Plant growth becomes limited under conditions of water deficit. This is

because water deficit leads to cell dehydration. Water deficit causes reduction
in cell turgor and dehydration which results in reduction in cell size and/or
volume. The cell wall extensibility decreases due to increase in cell wall pH. In
long term, water deficit leads to reduction in vegetative growth. Shoot growth
and leaf production gets severely affected. There is reduction in leaf
expansion and growth which is an adaptation in plants to reduce rate of
transpiration. The loss in turgor of cells affects the cell enlargement. Reduction
in low water potential enhances root growth gets thereby changing the root
shoot ratio. Improvement in root shoot ratio improves water supply as the roots
can extract more water by exploring larger volumes of soil.

Water deficit leads to closure of stomata. This has been considered as a

strategy adapted by plants to conserve water. The closure of stomata leads to
low internal CO2 concentration in plants and this limits the photosynthetic
capacity of the plant. Water stress affects the photosynthesis in two ways.
First by blocking supply of carbon dioxide to the chloroplasts due to closure of
224 stomata and second by changing the structural integrity of the photosynthetic
Unit 16 Plant Stress
machinery due to low water potential. The change in the photosynthetic
machinery affects electron transport and photophosphorylation.

The metabolic processes such as accumulation of solutes get triggered by

stress. The solutes which take part in osmotic adjustment include inorganic
ions (such as K+), sugars and amino acids. The solutes maintain hydration of
the protoplasm by decreasing osmotic potential under stress. They prevent
loss of membrane integrity and maintain protein stability. The amino acid
proline, sorbitol (sugar alcohol) and glycine betaine are the other solutes that
help in the recovery of turgor under conditions of water stress.

Anatomical and physiological adaptive mechanisms limit transpiration in

conditions of water deficit. The presence of thick cuticle, trichomes, sunken
stomata limit the loss of water through transpiration.

16.5.2 Salinity
Excessive accumulation of salt in the soil is referred as salinity. High
concentrations of Na+, Cl-, Ca2+, Mg2+ and SO42- are present in saline soils.
High concentration of salts replaces the essential ions present in the soil
resulting in toxicity. Salinity results in accumulation of Na and Cl ions in the
cytosol. The soils having high sodium (Na) content are called sodic soils. They
degrade soil structure by decreasing porosity and water permeability. High
concentrations of salts cause denaturation of proteins and membrane
destabilization by reducing hydration of molecules. The nutrient acquisition
gets disturbed from accumulation of toxic ions in soil.

Some plants growing in saline soils are not or less adapted. They are called
as glycophytes. These plants get exposed to stress referred as salinity
stress. In contrast, some plants are well adapted to saline conditions. These
are called as tolerant plants. For example-halophytes. These plants develop
various physiological and biochemical mechanisms to survive in highly saline
soils. The mechanisms involved in salt tolerance mainly include ion
homeostasis, compartmentalization, biosynthesis of
osmoprotectants/compatible solutes, activation of antioxidant systems
(enzymes and compounds), synthesis of polyamines, generation of nitric oxide
(NO) and hormone modulation.

Halophytes possess specialized salt glands on the surface of leaves that

excrete salt. Some plants accumulate toxic ions in the older leavers which
senesce and abscise to allow younger, more photosynthetically productive
leaves to survive.

Compatible solutes function as a protector or stabilizer of enzymes or

membrane structures that are sensitive to dehydration. Proline provides
tolerance to stress and serves as a nitrogen reserve. It functions as a
quencher of superoxide radical and hence exhibits antioxidant potential. It
provides tolerance to salt stress by increasing the activity of enzymes involved
in antioxidant defense system. Straight-chain polyols such as mannitol,
sorbitol and cyclic polyols such as myo-inositol or its methylated derivatives
increase in response to conditions of salinity. Trehalose accumulation caused
due to salinity stress protects plants against several physical and chemical
damages. Polyamines play an important role inducing tolerance in plants. 225
Block 4 Nitrogen Metabolism and Plant Growth Regulators
Although, abscisic acid (ABA) is a phytohormone which inhibits growth but its
application ameliorates the effect of stress conditions in plants. This hormone
gets upregulated in conditions of water deficit. Increased production of ABA
has been observed in shoots and roots of plants exposed to salinity stress and
water deficit. ABA acts as a vital cellular signal that modulates the expression
of many salt and water deficit-responsive genes. The genes related to
accumulation of K+, Ca2+ and compatible solutes (such as proline and sugars)
get upregulated by ABA. Hormones such as salicylic acid (SA) and
brassinosteroids (BR) are also known to participate in plant responses to
abiotic stresses.

A large number of genes and transcription factors are upregulated in response

to salinity in different plant species. These are called salt-responsive genes.
These genes are mainly involved in ion transport or homeostasis (e.g., SOS
genes, AtNHX1, and H+-ATPase), senescence [senescence associated gene
(SAG)], molecular chaperones (such as heat shock proteins), and
dehydration-related transcription factors (DREB). Upregulation of
metallothionein and water channel proteins has also been noted in plants
exposed to stress.

16.5.3 Pollutant Stress

Plants have adapted various ways to tackle stress caused by exposure of
various gaseous pollutants and toxic trace elements present in the
environment.

Exclusion - The process/means by which the plants maintain level of

contaminants below a toxic threshold. It mainly involves removing the element
out of the cell/body.

Tolerance - The process/means by which plants develop various biochemical

adaptations which allow the plant to survive stress conditions. These mainly
include compartmentalization and chelation.

Plants get exposed to high levels of heavy metals present in the soil. Some
plants show high accumulation of heavy metals such as copper (Cu), nickel
(Ni), zinc (Zn), chromium (Cr), lead (Pb), cadmium (Cd), cobalt (Co), iron (Fe)
in their tissues. These are called hyperaccumulators. High concentration of
heavy metals induces oxidative stress in plants. Antioxidant defensive
mechanisms play an important role in tackling oxidative stress caused by
heavy metal accumulation.

The plants exposed to air pollutants show alterations in the physiological and
biochemical features in plants. These features help in determining
susceptibility or tolerance to stress caused by pollutants. Changes in stomatal
behavior, carbon and nitrogen assimilation have been noted in plants exposed
to air pollutant stress. Air pollution stress leads to stomatal closure which
reduces availability of carbon dioxide to leaves and inhibits carbon fixation. As
a result the net phostosynthetic rate decreases thereby limiting the plant
226 productivity. Absorption of air pollutants such as sulphur dioxide (SO2), oxides
Unit 16 Plant Stress

of nitrogen (NOX) and carbon dioxide (CO2), suspended particulate matter

(SPM) by leaves causes reduction in the concentration of photosynthetic
pigments viz. chlorophyll and carotenoids. This affects the plant productivity.
The plants respond to pollutant stress by closing their stomata. This depicts
the avoidance mechanism adapted by plants. In contrast, synthesis of
antioxidant compounds reflects strategy adapted by tolerant plants.

SAQ 4
Answer in one word:

a) The plant hormone that accumulates in the water stressed leaves.

b) Soils having high concentrations of Na+, Cl-, Ca2+, Mg2+ and SO42-.

c) Solutes that protect or stabilize enzymes or membrane structures during

stress.

d) The process by which plant maintains the level of elements below a toxic
threshold.

e) Some plants show high accumulation of heavy metals in their tissues.

f) Some plants possess specialized salt glands on the surface of leaves

that excrete salt.

g) The condition in which roots and stem of the plants develop

interconnected gas filled channels that provide low resistance pathway
for the movement of oxygen and other gases.

16.5.4 Temperature Stress

Plants exhibit sensitivity to variations in temperature. Each plant requires an
optimum temperature for growth and development. Exposure to extremely
high and low temperatures adversely affects growth in plants.

Low temperature stress

An exposure to moderate chilling temperature may kill or cause injury to

plants. Maize, tomato, soybean, cotton are susceptible to injury when exposed
to temperatures between 10 to 15o C. The seedlings show wilting and
chlorosis in conditions of chilling stress. The low temperature causes
reversible changes in the physical state of the membranes. This occurs due to
change in the proportion of saturated and unsaturated fatty acids present in
the membrane lipids. The temperature at which the transition from liquid to
solid state occurs is called as transition temperature. At temperature above
the transition temperature, the membrane remains fluid but becomes solid or
gel like at temperatures below the transition temperature. With the drop in
temperature, the membranes show a transition from a flexible liquid crystalline
structure to a solid gel state. The rate of transition varies with the species and
depends upon the lipid composition of the membrane. Sensitive plants have 227
Block 4 Nitrogen Metabolism and Plant Growth Regulators
high proportion of saturated fatty acids while chilling resistant species possess
low proportion of saturated fatty acids but more of unsaturated fatty acids. The
membranes with this composition tend to solidify into a semicrystalline state at
temperature above 0º C. For functioning and stability, membrane needs to be
maintained in a liquid gel like state but under stress they are changed to solid
state which affects the membrane structure and activity. Freezing leads to
intra and intercellular crystal formation. Intercellular crystal formation damages
membranes and organelles. Extracellular ice crystal formation cause cellular
dehydration. This is because ice formation lowers the water potential in the
apoplast. It forms a gradient from high water potential gradient in the symplast
to low water potential gradient in the apoplast. Water moves from the symplast
to apoplast resulting in cellular dehydration which affects the membranes. The
formation of a stable ice crystal involves participation of several hundred water
molecules. This is called ice nucleation.

The damage to the plant cell membranes i.e. structure and function affects the
metabolism. Enzymes and enzymatic reactions are sensitive to temperature.
The rate of reaction doubles for each 10oC rise in temperature until an
optimum is reached beyond which the rate of reaction declines. The decline in
enzymatic activity results from unfolding of protein and is referred as thermal
denaturation. The Q10 increases linearly with short term rise in temperature.
Q10 measures change in rate of respiration. An increase in ambient
temperature causes change in the rate of respiration. .

The herbaceous species grown at low temperature exhibit a short, compact

growth habit, thicker leaves due to an increase in leaf mesophyll cell size
and/or an increase in the number of palisade layers.

Deciduous trees, conifers and shrubs such as birch, willow, survive cold
stress because they are able to adapt to low temperature conditions. The
adaptation in woody tissues begins in autumn when the growth and
photosynthesis ceases and plant enters dormancy. Hence the plants enter the
dormant phase prior to the onset of frost to prevent damage caused due to
freezing. During frost the respiratory activities sufficiently provide the energy
required for the numerous metabolic changes to attain maximum cold
acclimated stage. During this period the conversion of starch to sugars and
level of organic phosphate increases. The glycoproteins accumulate and the
protoplasm becomes more resistant to dehydration.

During adaptation to cold conditions, the temperate woody trees withdraw

water from the xylem vessels thereby preventing the stem from splitting in
response to the expansion of water during freezing. The physical properties of
lipids get altered at low temperatures resulting in increased membrane rigidity.

High temperature stress

High temperature (HT) stress is injurious to growth and development of plants.

High temperature stress causes loss of water content from the cell as a result
cell size gets reduced and ultimately the plant growth gets affected. Reduction
228 in seed germination potential, seedling growth, reduced radicle and plumule
Unit 16 Plant Stress
growth has been noted as the major impact of heat stress in plants (Fig.16.6).
Reduction in relative growth rate (RGR) due to decrease in net assimilation
rate (NAR) has been noted in conditions of HT stress. The negative effect of
heat on leaf includes reduction in water potential, leaf area and pre-mature
leaf senescence.

Exposure to high temperature alters the total phenological duration of the plant
life. Increases in temperatures affect the grain filling periods. A short period of
heat stress can cause significant decrease in production of floral buds. The
increase in sterility is caused due to impairment of meiosis in both male and
female organs, negative effects on pollen germination, reduced pollen tube
growth, reduced ovule viability, abnormality in stigmatic and style positions,
disturbance in fertilization process, hindrance in growth of endosperm and
proembryo. High temperature treatment reduces anther dehiscence and pollen
fertility. The reduced fertilization results from decrease in number of pollens on
the stigma.

Oxidative Alteration in
stress phenology
Water loss

High temperature

Reduction in
Reduction in Yield
metabolic events
growth reduction

Fig. 16.6: Major effects of high temperature stress on plants.

The plants that can adjust to high temperature conditions are called
thermotolerant plants. Plants exposed to cold temperature exhibit a lower
optimum temperature for photosynthesis while those growing at high
temperatures exhibit high temperature optima for photosynthesis. The tissues
of the plants are not able to survive temperatures above 45oC but some of the
tolerant species show survival at these temperatures because of their ability to
show of evaporative cooling. Leaf temperatures can raise to 4 to 5oC above
ambient air temperature in bright sunlight when soil water deficit causes
stomatal closure to high relative humidity reduces gradient driving evaporative
cooling.

Plants exhibit various mechanisms to ensure survival under high temperature

conditions. These include phenological changes, morphological adaptations
and short-term avoidance or acclimation (adaptation) mechanisms such as
changing leaf orientation, transpirational cooling, or alteration of membrane
lipid compositions. Closure of stomata and reduced water loss, increased 229
Block 4 Nitrogen Metabolism and Plant Growth Regulators
stomatal and trichome densities and larger xylem vessels are common heat
induced features in plant.

Early maturation noted in many crop plants has been considered as a

mechanism to escape heat stress. Plants growing in a hot climate avoid heat
stress by reducing the absorption of solar radiation. This ability is supported by
the presence of small hairs (tomentose) that form a thick coat on the surface
of the leaf as well as cuticles, protective waxy covering. In such plants, leaf
blades often turn away from light and orient themselves parallel to sun rays
(paraheliotropism). Solar radiation may also be reduced by rolling leaf blades.

Plants with small leaves are also more likely to avoid heat stress. They
evacuate heat more quickly due to smaller resistance of the air boundary layer
in comparison with large leaves. In well-hydrated plants, intensive transpiration
prevents leaves from heat stress, and leaf temperature may be 6°C or even
10-15° C lower than ambient temperature. High temperature can affect the
degree of leaf rolling in many plants

When plantlets or tissues of plants are shifted to 42° C and above, the
synthesis of normal proteins rapidly declines, instead synthesis of new
proteins is induced. These proteins are known as heat-shock proteins (HSP).
These proteins are self-regulatory i.e. their synthesis is switched off after 6 to
8 h of exposure to elevated temperature while synthesis of the normal proteins
resumes. The molecular weight of heat-shock proteins ranges from 15 to 102
kDa. HSPs are known to be induced also by heavy metals and arsenites. The
HSPs occur in representatives of all the major groups of organisms. A pre-
treatment at elevated temperature (at 45°C for 2 h) eliminates the heat-shock
response. It is believed that heat shock protein 2 protects essential enzymes
and nucleic acids from denaturation.

Induction of synthesis of heat-shock proteins has also been observed under

field conditions. In dry fields during summer when the leaf temperature
reaches or exceeds the ambient temperature (>40° C), synthesis of HSPs is
induced as under experimental conditions. The synthesis of HSP shows a
transcriptional as well as translational control.

How do HSPs help in heat-shock avoidance? They probably help important

cellular proteins to acquire conformations that would be safe and functional
under high temperature and the protein will remain in soluble state in the
cytoplasm.

Molecular approaches are also being followed to improve plant tolerance.

Plants tolerate HT stresses by modulating multiple genes involved in various
pathways. Heat stress up-regulates several heat inducible genes referred as
“heat shock genes” (HSGs) which encode HSPs. These proteins protect
intracellular proteins from being denaturation and preserve their stability and
function through protein folding. HSPs act as chaperones. High temperature
also induces synthesis of low molecular mass proteins known as heat shock
proteins (HSPs). The synthesis of these proteins in cells improves thermal
tolerance. HSPs are also induced by water deficit, ABA treatment, wounding,
230 low temperature and salinity. Three distinct classes of these proteins have
Unit 16 Plant Stress
been recognized on the basis of their molecular mass in plants. These include
HSP90, HSP70 and heterogenous group of proteins in the range of 17 to 28
kDa.

16.5.5 Stress by Infection and Wounding

Plants respond to various pathogens through an intricate and dynamic

defense system. The mechanism of defense has been classified as innate and
systemic plant response. An innate defense is exhibited by plant in two ways,
viz., specific (cultivar/pathogen race specific) and non-specific (non-host or
general resistance). A large array of proteins and other organic molecules are
produced prior to infection or during pathogen attack. Constitutive defenses
include morphological and structural barriers (cell walls, epidermis layer,
trichomes, thorns, etc.), chemical compounds (metabolites, phenolics, nitrogen
compounds, saponins, terpenoids, steroids and glucosinolates), proteins and
enzymes. These compounds confer tolerance or resistance to biotic stresses
by not only protecting the plant from invasion, but also giving the plant strength
and rigidity. The production of toxic chemicals, pathogen-degrading enzymes
(chitinases and glucanases) is used by plants. These compounds may be
present in their biologically active forms or stored as inactive precursors that
are converted to their active forms by host enzymes in response to pathogen
attack or tissue damage. Plant defense strategies involving these compounds
can fall in the category of either innate or systemic acquired resistance (SAR).
Innate immunity is of greater efficiency and is the most common form of plant
resistance to microbes.

Imbalances in biotic factors reduce cell proliferation, photosynthesis,

membrane integrity, protein stability and induce production of ROS, oxidative
damage and death. The plant exposed to insects or pathogens responds with
changes in the composition and properties of the cell walls and synthesis of
secondary metabolites that limit the infection.

Secondary metabolites associated with the hypersensitive reaction constitute

signal transduction pathways that prepare cells and tissues to resist secondary
infection. Because of this effect the plant reacts to infection by slowly
developing a general immune capacity. The phenomenon is called as
systemic acquired resistance (SAR). One important component of this
signaling pathway is salicyclic acid (SA). It is a naturally occurring secondary
metabolite with analgesic properties.

Methyl jasmonate is the principle constituent of the essential oil of Jasminium

and high concentrations of Jasmonic acid (JA) has been isolated from fungal
culture filtrates. Both JA and methyl ester of jasmonate mediate insect
resistance. Besides the role in insect and pathogen resistance, jasmonic acid
is supposed to be involved in many physiological processes such as seed and
pollen germination, protein storage, root development and coiling of tendrils.

Plants show specific responses to abiotic or biotic stress. These responses

mainly include production/synthesis of specific hormones, proteins and
accumulation of solutes that helps in maintenance of the physiological and
metabolic processes (Fig. 16.7). 231
Block 4 Nitrogen Metabolism and Plant Growth Regulators

Fig.16.7: Plants responses in various kinds of stress.

SAQ 5
Complete the statements given below:

a) The low temperature causes reversible changes in the ……………… .

b) The chilling injury can result in many ……………………. .

c) The measure of Q10 depicts change in ……………………….. .

d) The negative effect of heat on leaf mainly include ……………………… .

e) High temperature has a greater influence on the photosynthetic capacity
of …………………….. .

f) The plants that can adjust to high temperature conditions are called
……………………….. .

g) Synthesis of low molecular mass proteins in cells that improve thermal

tolerance are …………………….. .

h) …………….. acts as important component of the signaling pathway in

systemic acquired resistance.

i) Hormone that mediates insect resistance in plants is ………………….. .

232
Unit 16 Plant Stress

16.6 FUTURE PROSPECTS

Plants have evolved a variety of mechanisms to withstand stress conditions. In
many cases we can precisely define the way plants enable themselves to
survive and perform well under stress conditions. Based on such knowledge, it
should be possible to construct and breed plants tolerant to many
environmental extremes found in our country and elsewhere on earth while
trying to cultivate plants. There are so many salt affected areas in our country.
Salt tolerant varieties can be planted in large areas affected by salinity.

One of the ways to manipulate stress tolerant varieties is through genetic

engineering technique. With this technique it is possible to isolate a gene and
introduce it in a desired organism. This results in the expression of transferred
gene in the new organism which then starts behaving like the organism from
which this gene was originally isolated. The gene products related to a
particular phenomenon also get synthesised in the organism. The synthesis of
one or more proteins can be achieved easily through genetic engineering
technique. Thus, for instance, if genes for betaine synthesis can be transferred
to a plant sensitive to osmotic stress, it might confer tolerance against such
stress or an elicitor like phytoalexin can be produced by plants by artificial
gene transfer, it would be possible to have disease resistant plant.

These programmes can be directed towards incorporating the following traits

among common crop plants.

1) Non-photoinhibition of photosynthesis particularly in the tropics would

mean several fold increase in biomass produced over the same period of
time as water and carbon dioxide supply are non-limiting.

2) Resistance to high temperature beyond mesophilic ranges can allow

agriculture in several areas that are left uncultivated because of
prohibitive temperatures. To a large extent any strategy to achieve this
would depend on the water status of the environment.

3) Cold-hardiness in cultivars can help curb the losses often incurred

because of extremely low temperatures reached during winters in some
parts of the world.

4) Drought resistance in crop plants will be particularly helpful to our

primarily rain-fed agriculture.

5) Salt tolerance in plants will bring a lot more of territory under green
cover.

6) Disease resistance has always been a trait sought after in plants

adopted for cultivation. This can improve the present yield by 20 to 50%
(depending on the plant in question and the climatic zone).

7) Pest tolerance in the crop plants can be achieved by producing

proteinase inhibitors or by producing bacterial gene coding for pesticidal
protein. This limits their nutritional value but in cases like cotton where
our prime interest is fibre instead of food, this can still be a way to
achieve upto 50% improvement in yield. 233
Block 4 Nitrogen Metabolism and Plant Growth Regulators
16.7 SUMMARY
• Transition in the optimal environmental conditions which disturbs the
homeostasis and induces adverse effects on their physiology in plants is
referred as stress. Stress is of two types: abiotic and biotic. Plant
species get exposed to various stresses such as fluctuations in
temperature (chilling, freezing, high temperatures), water scarcity,
salinity, infections and pathogen infestations etc.
• Plants respond to stress in various ways. These mainly include
physiological, morphological and metabolic adaptations. These changes
help the plants to survive the changing environmental conditions. These
responses require no genetic modification and hence are referred as
phenotypic plasticity.
• Plants combat stress by two major ways i.e. avoidance and tolerance.
Stress avoidance includes a variety of protective mechanisms that delay
or prevent the negative effect of stress on plant. Stress tolerance is the
potential of a plant to acclimate to a stressful condition.
• Abiotic stress produces primary and secondary effects in plants. Primary
effects include reduction in water potential, and cellular dehydration.
These effects alter the physical and biochemical properties of cell. The
secondary effects include reduced metabolic activity, production of
reactive oxygen species (oxidative stress) and disruption of cellular
integrity.
• Plants respond to these stresses at various levels of organization. The
changes can be at cellular or genetic level. At cellular level, plants tackle
stress by alteration in the metabolic events such as production of
compatible solutes.
• Studies on plant responses to stress provide useful information. In
future, it would be possible to take measures by genetic engineering and
other means to prevent losses in yields of crops, fruits, vegetables and
other useful products due to stressful environmental conditions. It would
also be possible to bring into use chunks of land that at present cannot
be used for cultivation.

16.8 TERMINAL QUESTIONS

1. What are the two different strategies adapted by plants to overcome
stress?

2. What is oxidative stress and how does it affects plants?

3. What compatible solutes are synthesized by plants under conditions of

abiotic stress?

4. What are the main components of the antioxidant defense system in

plants?

5. Explain the reason for closure of stomata under conditions of water

stress.

6. What are heat shock proteins? What role do they play in plants exposed
234 to stress?
Unit 16 Plant Stress

16.9 ANSWERS
Self-Assessment Questions
1. a) physical stress i.e., heat/temperature stress
b) physical stress i.e., photosynthetically active radiation
c) physical stress i.e., flooding
d) chemical stress i.e., Pollution
e) physical stress

2. a) Stress-Any change in the surrounding environment that affects the

functioning of plants and disturbs homeostasis in plants is referred
as stress.

b) Tolerant plants – The plants that have the capacity to tolerate

stress by adjusting or acclimatizing to stress conditions are
referred as resistant/tolerant plants. The plants tolerance capacity
depends on their genetic capacity to adjust/overcome stress and
establish a new homeostatic state over a period of time.

c) Ephemerals- Some plants escape the stress. These are called

ephemerals or short lived plants. They germinate fast, grow and
flower quickly. They complete their life cycle during a period of
adequate moisture and form dormant seeds before the onset of
dry conditions. They survive the stress by avoidance. Many arctic
plants complete their life cycle during the short arctic summer and
survive during winter in the form of seeds.

d) phenotypic plasticity- Stress response in which plants respond to

changes in environmental conditions via changes in morphology
and physiology without any genetic modification. The changes are
reversible.

3. a) i) glycophytes
ii) water deficit or salinity
iii) reduction in water content
iv) osmolytes
v) trichomes
vi) ephemerals

b) i) Stress imposed on plants due to change in physical or

chemical environment is called as abiotic stress while the
stress caused by living organisms, specially viruses,
bacteria, fungi, nematodes, insects, arachnids and weeds is
called as biotic stress.

ii) Leaves of some plants orient themselves away from sunlight

to protect themselves from overheating. These leaves are
called paraheliotropic. Some leaves gain energy by orienting
themselves perpendicular to the sunlight is called as
diaheliotropic. 235
Block 4 Nitrogen Metabolism and Plant Growth Regulators
4. a) Abscisic acid (ABA)

b) Saline soils

c) Compatible

d) Exclusion

e) Hyper accumulators

f) Halophytes

h) Hypoxia

5. a) physical state of the membranes

b) metabolic dysfunctions in plants

c) rate of respiration

d) reduction in water potential, leaf area and pre-mature leaf

senescence.

e) C3 plants

f) thermotolerant plants

g) heat shock proteins (HSPs)

h) Salicyclic acid (SA)

i) Jasmonic acid (JA)

Terminal Questions
1. Two major strategies of combating stress recognized in plants are
avoidance and tolerance. Stress avoidance includes a variety of
protective mechanisms that delay, avoid or prevent the negative effect of
stress on plant. The plants which are able to tolerate stress are called as
stress resistant/tolerant plants. The plants ability to resist or tolerate
stress depends on their genetic capacity to adjust/overcome stress and
establish a new homeostatic state over time.

2. Oxidative stress occurs due to generation of reactive oxygen species

(ROS). During oxidative stress, the ROS are produced and accumulated
in high amounts in a cell or tissue than those that can tackled by the
biological system. The most common ROS include superoxide radical
(O2.-), singlet oxygen (1O2), hydrogen peroxide (H2O2) and hydroxyl
radicals (OH.). ROS possess strongly oxidizing capacity and potentially
affect metabolic activities such as photosynthesis, damage cellular
structures leading to reduction in growth, reduction in fertility and
premature senescence.

3. The plants respond to abiotic stress such as drought, salinity by

undergoing metabolic alterations. One of the strategies adapted by
plants to overcome stress is synthesis of compatible solutes. Compatible
236 solutes are the organic compounds which help in curtailing stress. They
Unit 16 Plant Stress
mainly include amino acids such as proline and sugars. Proline acts as a
ROS scavenger, molecular chaperone and helps in stabilization of
protein structure thereby protecting cells from damage caused due to
stress. The non-protein amino acid, γ-aminobutyric acid (GABA) helps in
maintaining carbon–nitrogen balance and ROS scavenging.
Carbohydrates such as starch and fructans act as storage substances
and are used as energy source during stress conditions. They stabilize
membranes and indirectly contribute to osmotic adjustment upon
freezing and dehydration by the release of hexose sugars. Sugars
function as osmolytes to maintain cell turgor and protect membranes and
proteins from damage caused due to stress. Trehalose (non-reducing
disaccharide) functions as an osmolyte and stabilizes proteins and
membranes in desiccation-tolerant plants. The level of trehalose
increases under stress. Polyols play a role in stabilizing macromolecules
and scavenging hydroxyl radicals. Accumulation of polyols such as
mannitol and sorbitol has been noted in several plants species exposed
to stress.

Glycine betaine (GB) is a quaternary ammonium compound produced in

plants exposed to abiotic stress such as cold, drought, and salinity. The
compound protects photosystem II, stabilizes membranes, and mitigates
oxidative damage. Various stresses such as drought, salinity and cold
induce synthesis of polyamines (PA). High PA levels have been
positively correlated with stress tolerance. PAs have been implicated in
protecting membranes and alleviating oxidative stress.

4. Antioxidant system includes antioxidant enzymes and non-enzymatic

compounds. These play critical role in detoxifying ROS. The activity of
antioxidant enzymes, such as superoxide dismutase (SOD), catalase
(CAT), glutathione peroxidase (GPX), ascorbate peroxidase (APX), and
glutathione reductase (GR) and accumulation of nonenzymatic
antioxidant compounds increases under stress conditions. Antioxidant
compounds such as ascorbate, glutathione also help in mitigation of
stress. They react with superoxide radical, hydroxyl radical, and
hydrogen peroxide thereby functioning as a free radical scavenger.

5. Abscisic acid (ABA) accumulates in the water stressed leaves and

mediates the loss of solute from guard cells. The plant regulates ABA
metabolism by modulating ABA synthesis in response to dehydration.
ABA accumulation in the chloroplast lowers the pH. The pH increases in
the region surrounding the mesophyll cells stimulates release of ABA
from the mesophyll to the apoplast. This supports efflux of K ions from
the guard cells. The guard cells lose turgor resulting in closure of
stomata.

6. Heat stress leads to up-regulation of genes referred as “heat shock

genes” (HSGs). These genes encode proteins called heat shock proteins
(HSPs). These proteins protect denaturation of intracellular proteins from
being and preserve their stability, function through protein folding. HSPs
act as chaperones. The synthesis of these proteins improves thermal
tolerance in cells.
237
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Volume 2 Plant Physiology and Metabolism

FURTHER READING
• Appling, D.R., Anthony-Cahill, S.J. and Mathews, C.K .2015.
Biochemistry, Concepts and Connections. Pearson Education
Limited. New Delhi.

• Buchanan, B.B., Gruissem, W., and Jones, R.L. (eds), 2000.

Biochemistry & Molecular Biology of Plants 2nd ed.. Wiley, Blackwell,
U.K.

• Hopkins, W.G. and Hüner, N.P.A. 2008. Introduction to Plant

Physiology.4th ed. Wiley, New York.

• Mohr, Hans and Schopfer, Peter (eds). 1995. Plant Physiology,

Springer-Verlag, Berlin.

• Nelson, D.L. and Cox, M.M. 2017. Lehninger Principles of

Biochemistry.7th ed. W.H. Freeman, U.S.A.

• Ochs. R.S. 2014. Biochemistry. Jones & Barlett Learning, USA.

• Smith, R.J., Lea, P.J. and Galton, J.R. 1999. Nitrogen Fixation. In Plant
Biochemistry and Molecular Biology (P.J. Lea and R.C. Leegood eds.)
2nd Ed. John Wiley and Sons. Chichester, New York.

• Taiz, L., Zeiger, E., Møller, I.M. and Murphy, A. 2015. Plant Physiology
and Development, 6th ed. Sinauer Associates, Inc. Publishers
Sunderland, Massachusets U.S.A.

• Salisbury, F.B. and Ross, C.W. 1969. Plant Physiology, Wadsworth

Publishing Company.

• Verma, V. 2016. Plant Physiology. 2nd Ed. Athena Academic, U.K.

• Woo H.R., Kim H.J., Nam H.G., Lim P.O. 2013. Plant leaf senescence
and death – regulation by multiple layers of control and implications for
aging in general. J. Cell Sci.126: 4823-4833.

• Woo H.R., Masclaux-Daubresse C., Lim P.O. 2018Plant senescence:

how plants know when and how to die. J. Exp. Bot. Volume 69: 715–
718.

• Matos F.S., Borges L.P., Müller C. (2020) Ecophysiology of Leaf

Senescence. Journal of Agronomy & Agricultural Science 3, 22.

• Plaxton. W.C. 1996. The organization and regulation of plant glycolysis.

Annu. Rev. Plant Physiol. Plant Mol. Biol. 47: 185-214.

239
Volume 2 Plant Physiology and Metabolism
GLOSSARY
Antioxidant : A substance that reduces damage caused by
free radicals.

Actinorhizal : Pertaining to several woody plant species,

such as alder trees, in which symbiosis occurs
with soil bacteria of the nitrogen fixing genus
Frankia.

Apoplast : Space in between the cells creating a pathway

through which materials may diffuse freely.

Arbuscles : Branched structures of mycorrhizal fungi that

form within penetrated cells; the sites of
nutrient transfer between the fungus and the
host plant.

Arbuscular Mycorrhizal : Symbiotic fungi that form hyphae growing in a

fungi loose arrangement, both within the root itself
and extending outward from the root into the
surrounding soil. The hyphae form unique
structures, such as the arbuscules that
enhance the exchange of nutrients between
the fungus and its host.

Bacteroids : Nitrogen-fixing organelles that develop from

endosymbiotic bacteria upon a signal from the
host plant.

Cessation : The ending of a process.

Chelates : Substances such as EDTA that from a

complex with divalent cations eliminating their
biological activity.

Chelator : A carbon compound that can form a non-

covalent complex with certain cations
facilitating their uptake (e.g., malic acid, citric
acid).

Coleoptile : A sheath protecting a young shoot tip in a

grass or cereal.

Denaturation : Breaking of weak linkages or bonds within a

molecule.

Eccentrically : Deviation from conventional or usual pattern or

style.

Endogenous : Anything that originates internally.

Heterogenous : Consisting of different, distinguishable parts or

240
238 elements.
Volume 2 Plant Physiology and Metabolism
Homeostatic : A stage by which an organism tends to
maintain stability or adjusting to conditions
required for its survival.

Hydroactive : Activated by water

Indeterminate : Not exactly known or defined.

Leghemoglobin : An oxygen-binding heme protein found in the

cytoplasm of infected nodule cells that
facilitates the diffusion of oxygen to the
respiring symbiotic bacteria.

Mesophilic : growing or thriving best in an intermediate

environment (moderate temperature)

Nitrate reductase : Enzyme that reduces nitrate NO3- to nitrite

NO2-. Catalyzes the first step by which nitrate
absorbed by roots is assimilated into organic
form.

Nitrite reductase : The enzyme that reduces nitrite NO2- to

ammonium (NO4+.).

Nitrogenase enzyme : The two-component protein complex that

complex conducts biological nitrogen fixation in which
ammonia is produced from molecular nitrogen.

Nod factors : Lipochitin oligosaccharide signal molecules

active in gene expression during nitrogen
fixing in which ammonia is produced from
molecular nitrogen.

Nodules : Specialized organs of a plant host containing

symbiotic nitrogen-fixing prokaryotes.

Nodulation genes (nod) : Rhizobial genes, the products of which

participate in nodule formation.

Nodulin (Nod) genes : Plant genes specific to nodule formation.

Photomorphogenesis : Light mediated development of plant.

Phytoalexin : substances produced by plants that inhibit the

growth of pathogen (such as a fungus)

Rhizobia : Collective term for the genera of soil bacteria

that form symbiotic (mutualistic) relationships
with members of the plant family
Leguminosae.

Skotomorphogenesis : The development of a seedling in the dark.

Stimulus : Thing that triggers a specific functional

241
reaction in an organ or tissue.
Volume 2 Plant Physiology and Metabolism
Susceptibility : The state of being influenced or harmed by
something.

Symplast : Inner side of the plasma membrane in which

water and low-molecular-weight solutes can
freely diffuse.

Synchronization : Two activities/events happening at a same

time.

242