Response of Plants to Drought Stress

Plants growing in drought stress may have the ability to control / avoid stress by escaping (Enduring Drought) or
tolerating stress (by developing succulent or Non-succulent habit). These two capabilities are collectively termed as
Drought Tolerance.

i. Drought Evading Plants

These plants remain under dormant / perennation to avoid stress period by

seeds and shoots. Such plants complete their life-cycle in few weeks within the
rainy season (eg. CO 16 variety of sorghum). They are called as Ephemerals.
These plants also prolong their life cycle for some time based on the necessity.
They reduce water loss by certain mechanism.

ii. Succulents

These (CAM) plants store enough water in their tissues. Their stomata open at
night. They have thick leaves and possess modifications (such as phyllodes
and phylloclades) under water stress conditions. They fix carbon during day
time with the help of malic acid and CO2, which is released internally during
respiration.

iii. Non-succulents

These plants endure drought with the following adaptive features:

1. Smaller leaves with thick cuticle

2. Sunken stomata with hair (pubescence) eg. Nerium
3. Shedding their leaves during summer to avoid excess water loss
4. Dehydration of protoplasm
5. Reducing enzyme activity
6. Favouring the syntheses of ABA (stress hormone) and Ethylene (senescence hormone)
7. Closing stomata due to increase ABA concentration, thereby reducing
water loss

Thus, because of these special features, succulents and non-succulents grow

well under drought conditions. They are not or least affected by stress.
Similarly, the arid zone plants also develop mechanisms to tolerate water
stress, hence they are not adversely affected in terms of growth and yield. But,
the non-arid zone plants suffer heavy loss in growth and yield because they do
not have above said mechanisms to tolerate the stress.

iv. Drought Resistant Plants

These plants resist the water stress situations due to the following adaptive
features / mechanisms. Therefore, these plants can be grown in drought facing / arid-zone areas.

1. Higher rate of photosynthesis because of efficient carboxylating systems (increased activities of RuBPCase,
PEPCase, Malic Enzyme etc.)
2. Store much water for proper hydration of protoplasm
3. Fix carbon by C4 pathway rather than usual C3
4. Producing “Aquaporins” – an intrinsic membrane protein in water-stressed plants, which enhance the water
flow by 10 – 20 folds (Chrispeels and Maurel, 1994).

Drought Resistance Mechanism

The ability of a crop species or variety to grow and yield satisfactorily in areas subjected to periodic water deficits is
termed as drought resistance

Types of drought resistance

1. Drought escape: The ability of a plant to complete the lifecycle before serious soil and plant water deficits
develop.
2. Drought tolerance with high tissue water potential: The ability of the plant to endure periods of drought
whilst maintaining a high plant water stress. This is also referred to as drought avoidance (Levitt, 1972).
3. Drought tolerance with low tissue water potential: The ability of the plant to endure periods without
significant rainfall and to endure low tissue water potential.

I. Drought Escape

Two features of desert ephemerals that are important in drought resistance are

1. Rapid phonological development

2. Developmental plasticity.

1. Rapid phonological development

Ability to produce flowers with a minimum of vegetative structure enables them

to produce seeds on a limited water supply.

2. Developmental plasticity

 This feature enable the plants to produce an abundance of vegetative growth,

flowers and seeds in seasons of abundant rain, enables the desert ephemerals
to both escape drought and survive long periods without rain.  

In crop plants, the greatest advance in breeding for water limited environment is
achieved by a shortening of life cycle, thereby allowing the crops to escape
drought. Therefore, there is a strong consistent negative correlation between grain yield and days to first ear
emergence and 40-90% variation in wheat yield under drought condition was accounted for by earliness. In wheat it
was observed that drought resistance is greater in early lines than late ones even at the same intensity of drought.
However, under adequate water supply, yield is often positively correlated with maturity date in determinate annual
crops such as maize, sorghum and sunflower.

An important aspect of developmental plasticity is the ability of plants to transfer assimilates accumulated prior to
seed-filling to the grain during the seed filling stage. It was also suggested that when water supply is adequate only a
small proportion of grain dry weight comes from the store of prior assimilate in the stems and roots, but when stress
occurs in the seed filling stage, an increased proportion of the prior assimilate is transferred to the seed.     

To achieve the developmental plasticity, plants frequently have an indeterminate habit. This is an important survival
mechanism in that it enables the large amounts of seed produced in wet years to carry the species through prolonged
drought periods.   

Selection of rapid phonological development is the most rewarding approach in breeding for drought resistance in
crops. In cereals, drought resistance varieties of wheat and barley flowered early than the others. However earliness
is often negatively correlated with yield in year of adequate rainfall.

II. Drought Tolerance at High Tissue Water Potential

Ability of the plant to endure periods of drought by maintaining high tissue water potential. This mechanism is also
called as drought avoidance.
To maintain a high water status during a period of high evaporative demand / or increasing soil water deficit, the plant
has two options. It must either reduce the water loss or maintain its supply of water.

A. Reducing Water Loss

i) Increased pubescence and ii) Increased leaf waxiness

Leaf pubescence Leaf waxiness

B. Maintenance of water uptake

i) Deeper root system 

ii) Hydraulic conductance of plants (increasing either the diameter of xylem vessels or their numbers).

III. Drought Tolerance at low tissue water potential

It is the ability of the plant to endure periods of drought and endure low tissue water potentials. This tolerance can be
achieved by

1. Maintenance of Turgor
2. Desiccation Tolerance

Desiccation Tolerance

Based on the desiccation tolerance of the protoplasm, plants can be classified as poikilohydric or homohydric plants.

1) Poikilohydric (resurrection plants)

The protoplasm of poikilohydric plants can withstand almost complete dehydration and can also withstand
dehydration and rehydration in concert with available water without damage.

2) Homoiohydric plants

Majority of the plants are homoiohydric plants. During growth and development, the protoplasm of homoiohydric
plants cannot withstand low water potential without injury. Dehydration caused mechanical injury to the protoplast by
physical tearing and destruction during water extraction and shrinkage. Small cells with no vacuoles and also the
cells that lose their vacuoles and also the cells that lose their vacuoles during dehydration can withstand the most
severe desiccation without mechanical injury. The changes in viscosity o the protoplasm and permeability of the
membrane play a role in desiccation  tolerance. It was also observed that cytoplasmic proteins are more stable to
denaturation, coagulation or hydrolysis in desiccation resistant plants and that enzymes are less susceptible to
inactivation by stress. RNA-DNA complex through which enzymes are manufactured is generally susceptible to
desiccation and sugars play a role in protecting this mechanism in desiccation resistant species and varieties. Sugars
may also provide protection against desiccation.
Biochemical effect of drought tolerance

1) Accumulation of Proline, Glycinebetaines etc.

2) Synthisis of Abscisic acid (ABA) etc

Mitigation of Water Stress

The adverse effects of water stress on crop growth can be mitigated by the application of chemicals such as nutrients, anti-transpirants a
Regulators (PGRs), which induce the plants to become adaptive to water stress situations for a specified period and the water requireme
minimized or saved.

1. Nutritional Management

Among the major nutrients, potassium and magnesium are found to be highly deficient due to water deficit conditions. Therefore, applica
the water uptake as well as the water relations in the plant tissues by osmoregulation processes, by acting as a potent osmoregulator (os
potential is reduced. Besides, potassium nutrition also helps in the favourable stomatal regulatory mechanisms, which regulate the water
has also resulted in the increased WUE of the plants. Similarly, magnesium is component of chlorophyll, its content and uptake is drastic
water stress effect. This is most prominent in Mg-loving crops like cotton.

 Besides macronutrients, deficiencies of micronutrients also appear under water deficit situations due to the following reasons:

1. Depletion due to erosion and leaching. In India, annual soil loss is estimated to be about 6000 Metric tons and obviously due to
and soils
2. Continuous use of micronutrients free NPK fertilizers in dryland agriculture and diminishing the use of organic matter, FYM, com
manures.
3. Use of high-yielding varieties (HYVs), adoption of intensive systems of farming and cropping and use of heavy doses of fertilize
proportionately the mining of micronutrients from the soil
4. Since increased crop production arising from the heavy demand of the nutrients in rapid depletion of macro and micro-nutrients
replenished. Consequently, the deficiencies of micro-nutrients in general and that of Zn, Fe and B in particular are widely sprea

     
Therefore, foliar application of the following nutrients depending upon the occurrence of their efficiencies will mitigate the water-stress ind
in crops.

a. 2 % DAP
b. 0.5 to 1 % potassium chloride (KCl)
c. 0.5 % Zinc sulphate
d. 0.5 – 1.0 % Ferrous sulphate + 1 % urea
e. 0.3 % Boric acid

Use of Antitranspirants

In India, about 90% of the land is under rainfed farming; therefore, it is very essential to manage every drop of water received through ra
measures are adopted to conserve he rain water, yet rainfed farming is often subjected to drought. Transpiration is said to be unavoidab
functions to attend in the crop cycle. For producing one tone of food, the crop plant requires varied amount of water as furnished below:

Cereals and legumes: 400 – 500 litres of water / kg of grains

Fruits and vegetable: 1000 litres / kg of food
Water transpired by crops (season / plant):

Maize                                 : 200 litres

Sunhemp                           :  27 litres
Cotton                               : 8 – 10 litres / day
Citrus                                :  100 – 200 litres / day
Trees (9 – 10 m height)      : 300 – 800litres / day
Forest trees of 400 – 600 trees:  20,000 barrels / day (1 barrel = 500 litres)

Similarly, the WUE of crops is also different and ranges from 0.24 to 1.75 kg / mm of water / ha. The WUE of sorghum is higher but that
difference lies with the maturity period and nutritive value of the crop. Cotton grows for six to seven months while sorghum grows for four

Drought reduces the yield by 0 – 100% depending upon the severity. Prolonged drought can drastically reduce the yield to zero level. Bu
– 15 days at early or late stage is common under rainfed conditions. Drought during the critical phenological phase like flowering and gra
detrimental. However, the crop productivity is dependant on how fast a plant can recover after a stress of 6-10 days.

The severity of intermittent drought of 6-10 days during critical stages of the crop can reasonably be avoided by the use of antitranspiran
saved. Antitranspirants can effectively be used to the crop under water stress with adverse rainfall.

Classification of Antitiranspirants (ATs) and field responses

The ATs are categorically classified on mode of action in the following four types:

I. Materials causing stomatal closure

1. Herbicides like 2, 4 – D, Phosphon D and Atrazine

2. Fungicides like Phenyl Mercuric Acetate (PMA)
3. Metabolic inhibitors like hydroxy sulfonates, potassium metabisulphite etc.
4. Growth hormones like ABA, Ethrel, TIBA, succinic acid, ascorbic acid and 
   Cycocel (CCC)

II. Reflectant Types

1. Kaoline
2. China Clay
3. Calcium bicarbonate
4. Lime water

III. Thin-forming chemicals

1. Hexadecanol (Higher alcohols)

2. Cetyl alcohol
3. Methanol
4. Paclobutrazol
5. Brassinolide
6. Resorcinol

IV. Polyethylene materials forming thick films

1. Mobileaf
2. Folicot
3. Waxol
4. S- 800
5. Hico-110R
(All the above chemicals are trade names given by the companies)

The purpose of ATs is to maintain the growth and productivity under stress conditions and it is never recommended for high productivity
and helps to get marginal yield when the expectations are zero.

Role of ATs in Irrigation Water Saving

Some of the ATs can also be used through drip (as Fertigation) to save the frequency of irrigation. In this context, the crop productivity c
26.2, 23.6 and 15.4 % over unsprayed control with the sprays of Hico-100 R, paclobutrazol and 8– Hydro Quinine respectively under 6 li
9 irrigations and thus considerably saved irrigation water.

Thus, assured benefits of ATs to the crops can be summarized as below:

1. Optimized yield levels under infrequent rainfall situations

2. Assured better crop growth and yield when no yields are expected using severe drought
3. Getting normal sized grains
4. Improved seed quality (so that produce can be used for seed purpose)
5. Saving of crops with marginal crop productivity under drought
6. Reducing irrigation especially in post-rainy long duration crops like cotton and pigeon pea
7. Minimizing irrigation frequency and saving water through drip irrigation (eg. Cetyl alcohol and / or Hexadecanol)
8. Monitoring crop loss with limited inputs
9. Monitoring / managing drought
10. Arresting fast receding soil moisture for better growth and yield of rabi crops
11. Very useful for farmers with minimum irrigation facilities
12. Saving large nurseries when water is scarce in summer months

USE OF PLANT GROWTH REGULATORS (PGRS)

The plants possessing moderate canopy development (moderate values for LAI), less reduction in photosynthesis, deeper root system, h
delayed senescence will perform better under water stress conditions. 

Toward this, application of some of the PGRs will prove beneficial for better crop growth and development when grown under water defic
PGRs and their effects on crops in order to suit to the water stress conditions are:

Cycocel & Mepiquat chloride: 

For promoting root growth (for more water absorption) and suppressing leaf area development (for reducing transpiration loss of water) a
senescence.

Cytokinins and Salicylic acid:  

They delay the leaf senescence processes and also favour stem reserve utilization by the developing grains especially during the water

Brassinolides: 

These PGRs increase the photosynthetic activity of the plants

Ascorbic acid: 

Ascorbic acid acts as an anti-oxidant agent for scavenging Reactive Oxygen Species (ROS) accumulating under stress and thus avoidin

Pre-sowing Hardening of Seeds / Plants:

Hardening of seeds / plants to required temperature / chemicals enables the plants to overcome the specific stresses. This process actu
(by osmoregulation), which enables the seeds to absorb more water under favourable situations to maintain its viability under unfavourab

Chemicals used for seed hardening process especially under rainfed conditions:
           

1. 1% KCl      2. 1% KH2PO4    3. 100 ppm Succinic acid        4. 0.5% NaCl        5. 100 ppm ZnSO4 
                 

6. 100 MnSO4         7. 100 ppm Ascorbic acid        8. 250 ppm Cycocel               9. 0.5% MgSO4                

Thus, these chemicals / PGRs could serve as boon to the frustrated farmers of rainfed areas, if rightly adopted with perspective vision to
Adoption of the agrotechniques is the only solution for farmers of dryland and water stressed scenario to save millions of world populatio
the developing countries, like India.

Agro-techniques for mitigating Water Stress

ource: 

Dept. of Crop Physiology, TNAU, Coimbatore

Related links:

http://www.plantstress.com/Articles/drought_m/drought_m.htm
http://www.plantstress.com/Articles/index.asp 
http://www.drought.unl.edu/
http://www.drought.noaa.gov/
http://www.aces.edu/issues/drought//

University drought sites:

Alabama Cooperative Extension Service 

Clemson University (South Carolina) 
Colorado State University
Iowa State University 
North Carolina State University 
North Dakota State University 
Ohio State University 
Oklahoma State University 
Pennsylvania State University 
Purdue University
Texas A&M University 
University of Georgia 
University of Kentucky 
University of Maryland 
University of Nebraska