Effects of Drought on Crop Physiology

By
Idris Zakariyya Kiri
E-mail: idriskiri@yahoo.com
G.S.M: 08025593505
Department of Biology
Jigawa State College of Education, P.M.B. 1002, Gumel, Nigeria
Abstract
The importance of water to plant growth has been recognized for over
300 years (Fageria et al., 2006). Drought stress is a major constraint to
crop production and yield stability in many regions of the world (Boyer,
1982). It is now understood that shortages of water at any stage in the
crop life cycle will have consequences on yield (Fisher, 1973). This paper
discusses the effects of water stress on crop physiology and water
drought resistance mechanisms as strategies to reduce effects of drought
on crop yield.
Key words: Drought, water use efficiency, water stress, drought
resistance.
Introduction
Drought is a meteorological term that means lack of precipitation over
prolonged periods of time (Hale and Orcutt, 1987). According to Mc-
William (1986), drought is a meteorological and hydrological event
involving precipitation, evaporation and soil water storage. It is an
environmental situation where decreases in soil moisture occur in rooting
zones of crops (Kobata, 1995). Droughts denote periods of time without
appreciable precipitation during which water contents of soil are reduced
to such extents that plants suffer from lack of water (Larcher 1995).
    Drought stress is a major constraint to crop production and yield
stability in many regions of the world. Soil water deficits are estimated to
depress agricultural crop yields world wide (Fageria et al., 2006).
Droughts are inevitable and recur frequently throughout the world,
despite improved abilities to predict their onset and modify their impact.
Drought remains the single most important factor affecting world security

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and stability of land resources from which food is derived (McWilliam,
1986).
Dent (1980) observed that reduction in soil water potential decreased
water potential of whole plants or parts of plant organs. Drought
influences biological and economical yield by inhibiting various
physiological functions because of water deficiency. Drought is the most
prevalent environmental stress and limits crop production on about 28
percent of the world land area.
Plants and Water Stress
       Higher plants are exposed to varying degrees of water stress at some
stage of their ontogeny. The type of water stress may vary from small
fluctuations in atmospheric humidity and net radiation in more mesic
habitats to extreme soil water deficits and low humidity in arid
environments (Morgan, 1984). Dunphy (1985) described the impact of
water stress as a function of duration, crop growth stage, type of crop
species or cultivars, soil type, and management practices. Drought is
sporadic in nature, resulting in drastic crop losses even in humid climates.
    Water deficits have major effects on plant photosynthetic capacity.
Reductions in photosynthetic activity have been found to adversely affect
crop yield (Gerik et al., 1996). Other adverse effects of water deficit on
plant photosynthetic capacity include reduction in cell growth and
enlargement in leaf expansion; assimilate translocation, and transpiration
(Hsio, 1973).
     Water deficiencies inhibited the enzyme-nitrate reductase. It has
been reported that even brief period of negative water balance causes
nitrate reductase activities to decrease by 20 percent or even by 50
percent after long period of water deficit (Hsio, 1973). The nitrate
contents of plants that have been supplied with fertilizer were found to
be increased as a result of water stress condition as observed (Larcher,
1995).

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      A sudden decrease in plant water status might induce decrease in
plant water status might induce stomatal closure, but adjustment by
leaves to stress may eventually result in stomata opening again
(Passioura, 1994).
Water Use Efficiencies of Crops
     The amount of biomass accumulated per unit of water used is refered
to as water use efficiency (WUE). In physiological terms; WUE has been
defined as the ratio of C assimilated to water transpired (Farquhar, Eh-
leringer, and Hubic, 1989; Dingkuhn et al., 1991). Passioura (1983)
expressed the role of WUE in determining yield per unit land area as:
Yield (g DM/m2) =WUE (g DM/g H2O) x transpiration
                  (g H2O/m2) x H1
Where DM is dry matter and H1 is harvest index (H1=grain yield/grain
plus straw yield).
      Water use efficiency, according to Mian et al. (1998), is a
physiological trait associated with drought tolerance of plants. It is the
reciprocal of the transpiration coefficients; it is the weight unit of plant
dry matter produce per weight unit water consumed (Mengel et al., 2006).
      Water use efficiency can be improved by increasing efficiency of
water delivery and timing of water application. This approach, according
to Gleick (1993), is widely adopted in many developed and developing
countries and in other countries where water is becoming increasingly
scarce. The share of water to agriculture will decline at even faster rate
because of increasing competition for available water from urban and
industrial sectors (Tuong and Bhuiyan, 1994). Under these situations, the
approach of adapting plants to less available water may be an important
strategy for crop production (Fageria, Baligar, Clark, 2006).
      Although genetic variations of WUE for many field crop plants have
been reported in recent years, nevertheless, improvement of WUE of field
crops through conventional breeding methods may not be practical
because of the complexity and difficulty of measuring WUE of large
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numbers of breeding lines grown on field conditions (Ismail and Hall,
1992). Indirect selection for improved WUE through molecular markers
linked to quantitative trait loci (QTL) conditioning WUE of crop plants
may prove useful (Mian, Ashely, and Boerma, 1998). Ludlow and
Muchow (1990) reported that differences among crop species for WUE
are related to carboxylation pathways (e.g., C4 species commonly have
twice higher values than C3 species) and to energy requirements for
production of biomass containing different proportions of proteins, lipids
and carbohydrates.
Effects of Water stress on Yield
      Four main aspects of plant behavior relative to drought that can
readily be linked with yield are modification of leaf area, root growth,
efficiency by which leave exchange water for CO 2, and processes
involved in setting and filling of seeds (Passioura, 1994).
      Effects of water deficits on crop growth and yield depend upon
degrees of stress and developmental stages at which stress occurs (Hsio
and Acevedo, 1974; Sullivan and Eastin, 1974). Westgate and Boyer
(1985) observed low water potentials during anthesis and early grain
filling can decrease yields in grain crops. Photosynthesis is inhibited
under these conditions and carbohydrate reserves may become limited for
grain growth (Westgate and Boyer, 1985).
  Effects of water stress on yield of the following crops have been
reported:
Cowpea:
(i) Two-week drought stress during vegetative or flowering growth
stages had no significant effect on seed yield of cowpea (Wien,
Littleton, and Ayanaba, 1979).
(ii) Flowering and pod filling, with yield reduction 35 to 69 percent
depending on timing and length of drought treatments, were
reported as the most sensitive growth stage to drought treatment
in cowpea (Shouse, Dasbery, Jury and Stolzy, 1981). No
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 significant reductions in seed yield were noted when water stress
occurred during vegetative stages (Labanauskas et al., 1981).
(iii) Moisture stress has been reported to increase efficiency of water
use by cowpea during vegetative flowering and pod-filling
growth stages (Turk and Hall, 1980).
Rice:
(i) Internal water deficits in rice were found to affect various
processes of photosynthesis such as inhibition of leaf blade
elongation (Turner et al., 1986), reduction of photosynthesis
rates per unit leaf area (Ishihara and Saito, 1983) and
acceleration of wilting (O'Toole and Moya, 1976).
(ii) Reduction in water potentials of leaf blade beyond certain units
led to delays in young panicle development (Tsuda, 1986), pollen
development was inhibited by reduction of water potential in flag
leaves (Namuco and O' Toole, 1986), exertion rate of panicles
during heading decreased, and numbers of sterile grains in-
creased (Cruz and O' Toole, 1984).
(iii) According to Fukai and Cooper (1996) rice grain yield decreased
at rates of 2 percent per day when 15-day stress periods (when
early morning leaf water potentials were less then -1.0 Mpa)
occurred during panicle development.
(iv) Rice reacted to drought stress with reductions in height, leaf area
biomass production, tiller abortion, changes in dry matter and
rooting depth and a delay in reproductive development (Asch,
Dingkuhn, Sow, and Audebert, 2004).
   Soybean:
(i) Water stress during any stage of development has been found to
reduce yield of soybean, although depending on stage of
development. Negative effects of stress are particularly important
during flowering, seed set, and seed filling (Souza, Egli, and
Bruening, 1997).

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 (ii) Carbon and Nitrogen are redistributed from leaves and other
vegetative plant parts developing seeds during leaf senescence
(Zeiher et al., 1982).
  Wheat:
(i) Wheat was sensitive to drought stress during anthesis. Drought
stress at anthesis lead to a reduction in the number of ears and
number of grains per ear and is thus detrimental to grain yield
(Sionit, Teare, and Kramer, 1980).
(ii) Drought stress is most sensitive in depressing grain yield in
spring wheat (Chiristen et al., 1995).
   Maize:
(i) Early kernel development was highly dependent upon current
supplies of assimilate in maize plants because reserves were not
sufficient to maintain kernel growth during low water potentials
(Wesgate and Boyer, 1985).
(ii) Wilson and Allison (1987) reported that mild water stress during
grain-filling reduced total dry matter accumulation but not grain
yield because of remobilization of assimilates from Stover to
grain.
(iii) Water stress during vegetation development of maize decreased
dry matter yield at harvest by reducing leaf area development,
stalk dry matter accumulation, and potential grain-filling capacity
of plants (Wilson and Allison, 1978; Eck, 1984; Lorens, Beanett,
and Loggale, 1987).
(iv) Maize is sensitive to drought stress at the tasseling stage (Mengel
et al., 2006).

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 Drought Resistance Mechanisms
Several physiological traits are taken into consideration when designing
programme for improving drought resistance in crops. Drought resistance
mechanisms have been divided into drought avoidance and resistance.
a) Drought avoidance: this refers to mechanisms or adoption abilities of
plants for reducing effect of soil and air dryness and plant water
deficits. Drought avoidance, sometimes, called dehydration
postponement or desiccation avoidance, may be achieved by
improved plant uptake of water from soil, reduced loss of water by
plants, high plant water-conducting capacity, or high plant storage of
water (Fageria et al., 2006).
b) Drought resistance: this is often related to physiological traits and
include moderating water use through reduced leaf area and shorter
growth duration, ability of roots to exploit deep soil moisture to
provide for evapotranspirational demands; capacity for osmotic
adjustment in leaf cells so that they can retain tugor and protect
meristems from extreme desiccation, and; control of non stomatal
water losses from leaves (Nguyen, Babu, and Blum, 1992).
Conclusion
Crop water use and WUE vary with different crops. The fact that
high temperature, poor irrigation practices, over exploitation of land
desertification, etc. along with drought reduce food production;
makes the impact of drought on agricultural production difficult to
quantify. Water productivity can be increased; increasing yield per
unit land area, either by using better crop cultivars, improved organic
practices or growing crops during most suitable period. Crop
physiology can provide many contributions to plant breeding
programme designed for improving drought resistance (Fageria et
al., 2006).

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 Recommendations
The following recommendations are put forward:
i) More research is needed for understanding physiology of
drought resistance in plants.
ii) Appropriate and correct phenology based on the
understanding of crop growth and development, to water
environments is highly essential.
iii) Advanced agronomic research towards defining the
various moisture environments, with particular references
to the amount, frequency, and probability of rainfall and of
expected soil moisture regime in average seasons (Fageria
et al., 2009).
iv) To optimize crop yields in irrigated environments,
irrigation should be timed in a way that nonproductive soil
water evaporation and drainage losses are minimized
(Fageria et al., 2006).
v) In rain fed environmental condition, management practice
must be employed in a way that water availability matches
crop water requirements.
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