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Stress Physiology in Plants

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 REFORMS IN AGRICULTURE AND RURAL DEVELOPMENT UNDER COVID-19 PANDEMIC

Stress Physiology in Plants

Pooja, Asha Sharma and Jyoti Sharma

Maharshi Dayanand University, Rohtak, Haryana, India

Stress in plants is a condition in which the plant growing in non-optimal or poor

state that negatively influences the plant growth, crop productivity, reproductive
capacity or death if the stress exceeds the plant tolerance limits. It comprises a large
range of factors which can be generally divided into two major groups: biotic or
biological stress factors and abiotic or environmental stress factors. Abiotic stress
factor includes the various environmental issues that disturb plant growth (such as
light, water-logging, temperature, salinity, drought and heavy metal toxicity), whereas
biotic stress factor is a biological harm (pathogens and pests), which a plant faces
during its period. Due to constant climatic change and worsening of situation, a high
risk is developed for food security and this state is imbalanced by human action. This
chapter continues with the actual knowledge of general views of the various types of
plant stresses, their effects and how plants respond to these different types of stress.
Responses to stress generally involves the various defense mechanism shown by
plants to cope up all these stresses. In this chapter we will also make available the
different defence mechanism adopted by plant to defend themselves from the
environmental stress includes hypersensitive response, accumulation of plant
hormones such as salicylic acid, jasmonic acid, and systemic acquired resistance
(SAR).
Introduction
Plants stress can be well-defined as any external factor that affects
the plant development, production, and all the processes of its life forms. Any changed
physiological condition caused by factors in surrounding environment may disrupt
homeostasis. Environmental changes in this equilibrium condition defined as
biological stress. It is an adverse effect on the plant which induced upon a rapid
conversion from some normal environmental condition and it disrupts this initial
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homeostasis state. Strain is somewhat physical or chemical alteration formed by a

stress, i.e. a well-known condition, which forces a system away from its normal state.
These environmental stresses signify various aspects which diminish the agricultural
production. Their detrimental effect is not only on present crop, but they also develops
barricades in the introduction of new species to the environment, so that the new
species are not used for cultivation.
Plant stress can be divided into two main groups. Abiotic stress is a physical (e.g.,
water, light, salt) or chemical way of damaging the plant and its surrounding. Biotic
stress is a biological harm, (e.g., insects, disease) which a plant faced during its
lifeforms. Many plants affected by a stress, shows abnormality in their growth and
biochemical functions. Plants may recover soon, when the stress is low or for short
time, the damage induced is temporary. But when the stress is more or for a long time,
plants shows inhibition in their development processes such as flowering,
germination, and reproduction that leads to death of the plant. These types of plants
are called susceptible. Those plants which escape the stress overall are stress escapers,
For example; desert plants, ephemeral or short-lived.
These short-lived plants germinate quickly following seasonal rains, and complete
their life cycle during a period of suitable moistness and leads to the formation of
dormant seeds before the beginning of the dry season. Similarly, many arctic plants
rapidly complete their life cycle during summer and survive in the form of seeds in
winter. These plants avoid stress condition in the environment by the avoidance
mechanism and never come in contact of it. If stress is present in the environment,
avoidance mechanism reduce the influence of any stress in plants. Some plants have
the ability to tolerate a wide range of stress which is called stress tolerance or
resistance by adapting or acclimating to it.
Two main types of stress
1. Biotic Stress
2. Abiotic Stress
Biotic Stress
It is the most important environmental stress factor which occur by the
communication among the plant and any living organisms that results in either
incomplete injury that the plant can overcome or complete injury that the plant can
die. Almost all kinds of pathogens, viruses, bacteria and fungi contribute to the biotic

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stress in plants. But microorganisms are most important in the form of diseases (viral,
bacterial and fungal diseases) inducing biotic stress. As we can say that, biotic stress
is an external biological stress that disturbs the plant growth and affects almost all the
group of plants, as well as cells, tissue, molecules, organelles, entire plants or even
entire plant residents. In this chapter we will examine about the biotic stresses and the
response shown by plants to the different biotic stresses.
Abiotic Stress
Abiotic stresses are external stress factors that can affect the plant growth for a longer
duration. In normal condition each plant have the ability to completing its life
processes and reaching to the maturity stage. Any abiotic stress or any change in the
normal biological factor in the environment alters the different physiological,
biochemical and metabolic function of plants and affect the plant growth. To cope up
with these abiotic stresses in the surrounding, plants protect themselves by acquiring
various defense mechanism to prevent their negative effect on growth and production.
Drought or water-logging condition, high or low temperature, metal toxicity,
excessive soil salinity, too much or too little light and nutrient deficiency in the soil
are some causes of abiotic stresses. Plant shows various antioxidant defense
mechanism to deal with all these severe stresses. In this chapter, we will discuss in
more detail about the abiotic stresses and response of plants for these abiotic stresses.
The concept of stress is closely associated with the two terms i.e. adaptation and
acclimation in plants. When tolerance increases in plant as a result of prior stress
exposure, it is said to be acclimated (or hardening). It is a temporary process while
adaptation is permanent. Adaptation is an evolutionary process and continues over
many generations in plants and animals. Both adaptation and phenotypic plasticity
provide tolerance to the plants in highly stress condition.
Strategy to face the stress condition
 Escaper: Those plants or organism who completing their life cycle before the
occurrence of a stress.
 Stress Avoider: By altering their morphology in accordance with the stress,
such as reduced leaf area, stomatal conductance.
 Stress Tolerant: Tolerate the effect of stress without dying or suffering injury.
Attained by specific physiological, biological and molecular mechanism at

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cell level which include specific gene expression and accumulation of specific
proteins.
Stress Factors:
As we discussed earlier that only at normal condition, plant grows and reproduce in a
better way and produce good yield quality. But when plants are subjected to a different
level of stresses, (biotic and abiotic) changes the growth condition from the normal
state. These stress factors also known as stressors. When plants faces these kinds of
stressors, they trigger certain mechanisms for their adaptation and defences which we
will discuss later in this chapter. This section deals with some important biotic and
abiotic stresses and their effect on plant growth.
Some Major Types of Abiotic Stress
1. Drought or water stress
2. Salt stress
3. Temperature stress
4. Light stress
5. Heavy metal stress
Drought or water stress
Water is the main component of life forms containing 80-90% of the inner mass of
non-woody plants, and the major intermediate for conveying various nutrients and
metabolites by which plants complete their different processes of development. The
availability of excessive (flooding) or inadequate (drought) supply of water is called
the water stress. It is one of the major abiotic stress that affect the overall status of
plant and its development and happens for numerous reasons, such as excessive
salinity, flodding, insufficient rainfall, excess of light and fluctuations in temperature.
Plants faces water-deficit condition either when the uptake of water through roots
becomes restrictive, or when the rate of transpiration turn into high. The first response
of drought stress is wilting because the function of turgor pressure is lost, which
expands plant cells and retains them rigid. In plant leaves, cells start to collapse
without this turgor pressure and forming a floppy appearance. Plant cells start
collapsing due to increase in wilting and leads to deaths. Drought stress occurs when
plant demands more water at a given time or its ability of uptake is reduced.
Changes in plant growth
 Under prolonged drought, plants will dehydrate and die.

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 Reduce the plant-cell’s water potential.

 Affects translocation indirectly by altering the source to sink relationships for
assimilates.
 Changes in structure of macromolecules by removal of water.
 Rate of cell expansion is inhibited due to loss in turgor pressure.
 Dehydration of mesophyll cells inhibits photosynthesis.
 Decrease in turgor causes stomatal closure.
 Overproduction of reactive oxygen species.
 ABA inhibits shoot growth appears to be promoted.
 Alters gene expression in the plant leaves decline during water deficit
condition by supressed synthesis.
 Disturb the membrane stability by destabilization between proteins and
membrane lipids. Alteration in fatty acid arrangement causes drought
condition.
 Osmotic adjustment- certain organic compounds such as sucrose, amino acids
(especially proline and betaine), inorganic ions (especially K+ ) and several
others that lower the osmotic potential and thus, maintain water potential of
cells without limiting enzyme function.
Plant Responses to Drought Stress for Prevention
 Primary responses of plants to drought stress generally support the plant
to relive for a while.
 The main aspects of plant responses to water involve the maintenance of
homeostasis by ionic balance and osmotic adjustment.
 Detoxification of plant cells through antioxidant enzyme such as SOD and
Catalase are produced by the activation of defense responsive mechanism.
 Reduced leaf area, stomatal conductance and highly developed root
system.
 Osmolytes is the accumulation of certain compatible solutes like proline
and betaine in the plants during water stress condition and provide
resistance to the plants by the osmotic adjustment in cells. These osmolytes
helps in various stress conditions.
 Abscisic acid, a plant hormone which is produced mostly in stress
condition, also known as stress hormone. Accumulation of ABA under
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drought conditions helps the plant to survive from dehydration and provide
tolerance.
 ABA induces the expression of various genes and closing of stomata as a
defensive response during drought stress. Increase in concentration of
ABA near the guard cell results in closure of stomata and supports the plant
to save water from transpiration.
 During drought tolerance plants perform functions normally even at low
water potential whereas in drought avoidance, plants maintains its ability
of high water potential under water-deficit conditions.
Salt Stress
Salt stress can be defined as the accumulation of excess content of salt in the soil
which cause inhibition of plant growth and sometime also leads to plant death. It is
one of the major abiotic factors which decrease the crop yield in the arid region.
Salinity leads to a number of inhibitory effects on the plant like limiting the crop
productivity, seed germination, and crop yield. Salinity, in the higher concentration,
also cause damage to the plant by ion toxicity, membrane disorganization, oxidative
stress, water potential and decrease in cell expansion and its division. All these effects
cause an adverse effect on the plant development and crop yield. Salt stress caused
numerous damage to plants which includes symptoms like accelerated growth, growth
inhibition and senescence. It may also lead to death due to longer exposure to the
salinity. Under the salt stresses, different plant characteristics like physiology,
morphology, anatomy, chemical composition and water content of plant tissues are
affected.
Affects plant adversely in two ways:
 High solute content in rooting medium creates water stress by decreasing
osmotic potential and
 Direct toxic effect of higher concentration of ions.
Different plants response differently to the salt stress based on their salt tolerance
ability. So, based on the tolerance ability, plants generally classified into two
categories:
1. Halophytes
2. Glycophytes
Halophytes
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 These are not usually affected at higher salinity.

 Known for their ability to grow comfortably in saline soil with high salt
concentration.
 Have a specific mechanism of salinity tolerance.
 Some of the halophytes excrete extra salt content with the help of salt glands
present in their leaf cells while some plants excrete via salt hairs present on
their stem.
Glycophytes
The plant which cannot grow in the presence of high concentration of salts. But
manage to grow in saline soil by adopting some mechanisms as
 Accumulation of sugars in leaves
 Compartmentalization (checked from reaching to photosynthetic parts)
Mechanism of Tolerance
 Presence of salt excreting glands that reduce the concentration of salt in the
plant e.g. Frankenia, Spartina.
 Development of small leaves, water storage hairs, and aerenchyma.
 Succulence that may lead to dilution of intracellular salt such as occurs in
Salicorma sp.
 Sometimes there is a synthesis of organic solutes that aids in the maintenance
of turgor.
There are two ways by which plant adapt themselves from the severe consequence of
salt stress:
I. Avoidance: A state of avoiding excessive salt from the parts of the plant where they
are detrimental through-
• Salt exclusion: Some plants exclude salts from their parts are known as salt
excluders. In this, movement of salt ions are restricted from the root part whereas
permitting the water to pass. For example; mangrove plants.

• Salt excretion: Those plants which eliminate the salt through cuticle or bladders or
glands present on each leaf called salt excreters. For example; Salt bladders – e.g.
Atriplex, Mesembryanthemum crystallinum L.

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Salt glands – These are the trash sites for the absorption of excess salt from the soil
water; support the plants by adapting in saline soil. For example; Excretion by cuticle
– e.g. Tamarix.

• Salt Dilution: Plants having dilution capacity in their parts and they dilute the ions
in them by balancing succulence. Plants attain this ability by maintaining their storage
volume high and thick like a succulent shape. And it is acquire through vacuoles of
mesophyll cells filling with water as well as large in proportions. This adaptation is
inadequate in the plant tissues.

II. Tolerance
• Osmotic adjustment- Some plants can tolerate salt stress by dropping the cellular
osmotic potential by increasing the solute accumulation (sucrose, amino acid, proline,
betaine).

• Hormone synthesis - ABA stress hormone stabilizes plants against excess salt
stress.
Temperature Stress
Temperature stress is one of the severe environment factor which affect
plant development, by showing low germination rate, growth retardation, reduced
photosynthesis, and often die.
It is divided into three main parts:
1. Chilling Injury
2. Freezing Injury
3. Heat Stress or High-Temperature Stress
Chilling Injury
The injury which causes due to low but above zero-degree centigrade temperature is
called chilling injury.
 Temperature 0oC to 15oC causes chilling stress.
 Less than 0oC will cause freezing stress.
 This condition is shown by many tropical area plants.
 This chilling temperature is very low for optimal growth but not sufficient for
ice formation.
Effect of Cold Temperature on Plants
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 Discoloration of leaves
 Delayed transition of flowering and plant growth.
 Shrinkage of protoplast due to extracellular ice formation.
 Destruction of chlorophyll.
 Photosynthesis inhbition, degradation of proteins, lower transpiration rates,
reduced carbohydrate consumption.
 Water potential inside the cell is affected due to low temperature. Because the
water inside the cell starts to convert in ice at low temperatures and will result
in high solute potential and low water potential.
 This will affect the enzyme and ion channel of the plant and as a result, plants
will die due to ice crystal formation.
 This is shown by plants- tomato, bean, rice, etc.
Freeing Injury
Freezing injury occurs at a temperature below the freezing point of water (below 0oC)
and primarily the liquid phase in the cell wall freezes.
 At 0oC there is a phase transition in water from liquid to solid. Development
of ice crystals in the cell walls and intercellular places.
 Bulky polysaccharides enable ice crystal development, which is known as ice
nucleators.
 Damage occurs when ice crystals grow and puncture into the cytoplasm.
Adaptation of Plants for Prevention
1. Needle-like leaf structure which prevents freezing and transpiration of water
due to low surface area. They also have terpenes and alcohol which are
antifreezing factors.
2. Termination of growth activity during winter.
3. Cold area plant has anti-freezing proteins. It is also found in the winter Rye
plant.
4. Production of dehydrins (stress proteins) which prevent dehydration in plants
in response to drought and cold fluidity of membrane lipids stress.
5. Some trees become leafless so to avoid the accumulation of ice on their
surface.
Cold Stress Signalling

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 Calcium level increases inside the plant cell on cold stress conditions and this
will lead to the further expression of genes that will help in tolerance from
cold.
 Low temperature will increase the level of ABA and Jasmonic acid. Their
signaling pathway will also increase.
 Low temperature will decrease the level of Gibberellic acid, ethylene, and
Cytokinin. So, all the signaling will get inhibited.
High Temperature Stress or Heat Stress
High temperature (HT) or Heat stress is also a main abiotic stress that affects the
development of plant, their different processes, and their overall production. Almost
every physiological and biochemical processes are sensitive to high
temperature. Different plants have different ability to respond to the high temperature.
 Normal temperature ranges from 0oC to 40oC.
 Stress temperature is above 40oC.
Effect on Plant Growth
 Change the plant growth pattern, reduction in seed germination, changes in
leaf pattern, and decrease in plant production.
 ROS generation that will lead to apoptosis of plant cell.
 High temperature may cause injury to the chloroplast by-
1. Disturbing thylakoids structure
2. Swelling of grana
3. Affect the grana stacking ability
 Inhibition of photosynthesis and respiration. Chloroplast enzymes become
unstable and performs abnormal function.
 Fluidity of membrane lipids, destabilize the bond between polar proteins
within membrane.
 Change membrane composition and structure and can cause leakage of ions.
 Denaturation of proteins occur by losing its three-dimensional structure which
is required for the activity and function of proteins. Formation of unfolding
proteins, destabilize the cell functions.
Some plants avoid high-temperature stress by:
 Angle and arrangement of the leaves.

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 Thick cuticle and leaf hairs (improves conductive cooling) reduce the
absorption of solar radiation.
 Closure of stomata to reduce water loss at high temperatures.
 Large xylem vessels.
 Rolling of leaf blades.
Tolerance to High Temperature
 High expression of antioxidant enzymes like Catalase, Peroxidase, SOD,
Glutathione reductase.
 Plant produce antioxidant metabolites like ascorbate, carotene etc.
 Expression of Heat Shock Proteins (HSPs).
Light Stress
Light stress is also a stress factor that damages plants and its development very badly.
Light is one of the most important components for photosynthesis as an energy source
and a main environmental factor for plant growth and development. But the changed
light quality and quantity can also be harmful to plant processes and lead to photo-
destruction and photo-inhibition. Due to changed light intensity either low or high,
plants unable to protect them by being stabilized and loss their normal metabolic
functions.
 When light intensity is low below the compensation point, carbohydrate level
in plants declines by using it as a substrate first for respiration and then for the
other purpose.
 In high-intensity, light photosynthesis performance of plants, is depends upon
the relationship between photo-inhibition of photosynthesis and the damaged
chlorophyll ratio.
Ultraviolet radiation
At sea level, electromagnetic radiation contains around 7% ultraviolet rays coming
from the sun. Due to these harmful ultraviolet rays, membrane permeability of plant
cells changes by changing in lipid and protein ratio. These are in the result of reduced
photosynthesis and respiration.
Avoidance of Light Stress by Plants
 To avoid the absorption of excess of light, various plants moves their
chloroplasts in a way that they receive minimum light. On the other hand,

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during low light, chloroplasts may be moved to high-intensity light to make

the best use of it.
 To protect themselves from the excess of light, plant leaves make an angle to
diminish light interception.
 Plants of shady region grow horizontally to take full advantage of light
interception.
References
 Loon LC (1985). "Pathogenesis-related proteins". Plant Molecular
Biology. 4 (2–3): 111–116.
 Lutz, Diana (2012). Key part of plants rapid response system revealed.
Washington University in St. Louis.
 Matthews B. "The Hypersensitive Response". Agricultural Research Service:
Plant Science Institute. The United States Department of Agriculture.
Archived from the original on 2007-02-22. Retrieved 2007-01-12.

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