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FUTOJNLS 2016 VOLUME-2, ISSUE-1, PP- 70 - 78

Futo Journal Series (FUTOJNLS)


e-ISSN :2476-8456 p-ISSN : 2467-8325
Volume-2, Issue-1, pp- 70 - 78
www.futojnls.org

Research Paper July 2016

Physiological Crop Adaptation Mechanisms to Moisture Stress:


Basis for Exploiting Crop Genetic Yield Potentials
Adikuru, N.C.*, Obiefuna, J.C., Okoli, N.A., Alagba, R.A., and Ibeawuchi, I.I

Department of Crop Science and Technology, Federal University of Technology, Owerri, Nigeria
*Corresponding author: adikuru@yahoo.com

Abstract
Plants are subjected to a variety of environmental stresses which reduce plant growth and yields. Abnormal
temperatures, moisture stress, poor soil condition, and pest attack are common and regular. Moisture stress
is most critical in tropical agriculture. The extreme weather events due to climate change, creates additional
drought that threaten crop farming even in the rainforest agroecology. Today, increased global water
demands negate conventional irrigation strategy. The alternative is the development of drought tolerant
crop varieties and improved agronomic packages based on physiological knowledge of crop adaptations
to moisture stress. This paper reports the natural crop adaptations to moisture stress across Nigeria
agroecological zones with a view to exploit the knowledge in breeding programme to mitigate the challenges
of moisture stress and climate change.

Key words: Crop adaptation, mechanisms, moisture stress, yield potentials.

1.0 Introduction

Water is significant to the life of crops and other higher plants because of its functions. It is a major
constituent of protoplasm and reduction of water below some critical level is accompanied by changes in
structure and ultimately in death (Kramer, 1983). As a solvent, water provides the medium for the movement
of molecules (gases, minerals and other solutes) within and between cells and to a large extent influences the
structure of proteins, nucleic acids, polysaccharides and other cell constituents (Taiz & Zeiger, 2002).

Large internal hydrostatic pressures (turgor pressure) built up in plant cells as a result of the normal
water balance in the cells, is essential for many physiological processes including cell enlargement, gas
exchange in the leaves, transport in the phloem and various transport processes across membranes (Taiz &
Zeiger, 2002). Irrespective of its importance, water is frequently in short supply and this is responsible for
moisture stress during crop production. Among several environmental stresses including abnormal
temperatures, poor soil condition, and pest attack, moisture stress has been observed as the major abiotic
stress factor limiting crop productivity worldwide (Sharp et al., 2004; Barnabas et al., 2008). This limiting
ability of water is the reason for the practice of crop irrigation. While irrigation has been effective in
addressing the problem of moisture stress, it is becoming less so as global water demand increases (Boyer &
Westgate, 2004) and extreme weather events create additional drought that threatens crop farming even in
the rainforest agroecology. The development of drought tolerant crops should provide an inexpensive
alternative towards addressing the problem of moisture stress. In this regard efforts should be based on
physiological knowledge of crop adaptation to moisture stress. Therefore, this paper reports the natural crop

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adaptations to moisture stress across Nigeria agroecological zones with a view to exploit the knowledge in
breeding programme to mitigate the challenges of moisture stress and climate change.

2.1 Concept of Moisture Stress

Water stress in crops occur when crop water potential and turgor deficits interfere with normal
physiological function of the crop. Thus, crops experience water deficits during periods when water loss in
transpiration exceeds absorption (Kramer, 1983). The exact cell water potential at which water deficit occurs
depends on the crop, the stage of development and the metabolic process under consideration. For example,
cell enlargement usually decreases at a water potential of only -0.2 to -0.4 MPa (megapascal), stomata
closure does not begin until the water potential falls to -0.8 to - 1.0 MPa or even lower in some crops. Major
causes of water stress in crops include rate of transpiration, availability of soil moisture and competition for
water among organs.

2.2 Rate of Transpiration

The rate of transpiration which depends on irradiance determines the degree of water stress development in
crops. Water deficits caused by excessive midday transpiration often reduce daytime growth of plants. The
interrelationship between soil water and crop growth is affected by atmospheric factors that influence the
rate of transpiration, mainly because of the high tropical temperature (35 - 42 oC) and high vapour pressure
deficit accompanying a high level of irradiance (Kramer, 1983).

2.3 Competition among Organs

As a result of differences in stage of growth and exposure, different parts of crops lose water at different
rates. The internal resistances to water flow and differences in osmotic potential, results in differences in
water potential and the eventual redistribution of water within crop. For example, upper leaves are more
exposed than lower leaves and transpire more rapidly, thereby lowering their water potential and causing
water to move to them from the shaded lower leaves. Both young leaves and growing fruits compete for
water at the expense of older leaves which often die prematurely from desiccation on water stressed crops.
Leaves compete successfully with older fruits which often show reduced growth or even shrinkage at
midday, when transpiration is rapid.

2.4 Availability of Soil Moisture

The amount of soil water available to a given species depends primarily upon the size distribution of the soil
pores. Size distribution, in turn, depends on both soil texture and structure (degree and type of aggregation).
Generally, medium to fine textured soils hold more water for plant use than coarse textured soils. As
depletion of the nearest pores proceeds, a gradient of water potential is set up between the root and the bulk
soil causing water to flow towards the root from at least a few millimeters (mm) from the root surface.
However, the withdrawal of water from the larger pores reduces the volume of soil through which flow can
take place and increases the tortuosity of the pathway between the moist bulk soil and root. These combined
effects stimulate a drastic reduction in the hydraulic conductivity of a soil as it dries (Fitter & Hay, 1981).
Generally, the supply of water hardly meets crop water demand during rapid transpiration, such that
temporary wilting and leaf water stress usually occur long before the soil permanent wilting point is reached.

3.0 Effects of Moisture Stress in Crops

The effects of water stress on crops may be considered at the cellular level, the whole plant level or in terms
of the processes affected. The most obvious effect however is to reduce growth, with cell enlargement being

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particularly sensitive to water deficits. Cell division is normally less sensitive compared to cell expansion
(Jones, 1992).

In crop response to drought stress process, numerous physiological and biochemical changes occur
(MohammadKhani & Heidari, 2008) including the increased expression and synthesis of proteins (Riccardi
et al., 1998) and retarded seedling growth and dry mass production of corn cultivars (Rahman et al., 2004).

Moisture stress may adversely affect assimilate translocation from the source to the sink in crops for
example, when the source (leaves) is small and less photosynthate is available for translocation to sink
(fruits) is a case in point (Hale & Orcutt, 1987). The effects of moisture deficits on vegetative crop growth
phases include the reduction of tillering in grasses and the early termination or formation of dormant buds of
growth in perennials. During reproductive phase, moisture stress stimulates the emergence of ready-
differentiated floral buds which advances flowering in annuals and delays flowering in perennials (Jones,
1992). In maize, moisture stress increased tassel anthesis (1 to 5days), delayed silking (3 days) and increased
anthesis silking interval by 2 days (Munyiri et al, 2010). These apparently small physiological variations
resulted in 17 - 81 % yield reduction. Thus, crop yield loss depends on the crop growth stage at the time and
severity of moisture stress incidence.

3.1 Crop Adaptation Mechanisms to Moisture Stress

Moisture stress resistance refers to the ability of a crop plant to produce its economic product with minimum
loss in a water-deficit environment relative to the water-constraint-free management (Mitra, 2001). In
rainfed crop agriculture and horticulture in particular, a tolerant cultivar has a high yield of marketable
product in drought conditions (Jones, 1992).

Naturally, plants may adapt to dry conditions in several morphological or physiological ways. However, the
crop plants adapt to moisture stress in rainfed agriculture through any or combinations of the three
mechanisms of drought management: escape, avoidance and tolerance (Turner et al., 2001).

3.2 Drought Escape Mechanism

In rainfed crop agriculture, drought escape plants complete their life cycle during the period of sufficient
water supply (rainy season) before the onset of drought. This mechanism involves rapid development, early
flowering and early maturity, developmental plasticity, variations in duration of growth period depending on
the extent of water deficit and remobilization of pre-anthesis assimilates to grain (Mitra, 2001). Phenological
adjustment to escape from the moisture stress duration is most successful in C3 cereals such as wheat or
barley in Sahel agroecology of Nigeria which experiences early terminal drought (Lopes et al., 2011). This
mechanism is also typical of the desert ephemerals that can complete their life cycles from germination to
seed maturation in as little as four to six weeks (Jones, 1992). In many crop plants, the most drought-tolerant
cultivars usually flower and mature early thus avoiding the worst of the dry season, e.g. early maturing
soybean, okra and maize. The early maturing cultivars are planted (March-April) with the early rain to
mature in July-August (Manavalan et al., 2009). Better partitioning of assimilates to developing fruits is
also a critical attribute in improved reproductive success usually associated with the ability of the plant to
store reserves in stems and or roots and to mobilize same for harvest production. (Chaves et al., 2003). This
ability to mobilize reserves is increased in moisture stressed plant, most likely due to stimulation of
sink capacity of seeds and pods (Chaves et al., 2002). Thus, a large photosynthetic accumulation prior to
flowering is an important factor for plant production and survival during a drought enhancing flowering
process in tropical fruits. Therefore, differences among genotypes in the ability to store and utilize stem
reserves, as well as in photosynthetic capacity could be selection criteria in crop breeding for Sahel

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agroecology of Nigeria. However, biochemical changes in stems and the process regulating storage of
reserves in plant subjected to drought are poorly understood (Chaves et al., 2002).

3.3 Drought Avoidance Mechanism

Drought avoidance or dehydration avoidance is the ability of plants to maintain relatively high tissue
water potential despite a shortage of soil moisture. Performance of drought avoidance involves
maintenance of turgor through increased rooting depth, and efficient root system e.g. tree crops and food
legumes, increased hydraulic conductance and reduction of water loss through reduced epidermal
(stomatal and lenticular) conductance, reduced absorption of radiation by leaf orientation, rolling or
folding and reduced evaporation surface or leaf area (Mitra, 2001).

Each of these mechanisms may either minimize water loss (stomata closure, leaf rolling, increased
reflectance or steep leaf angles, and reduced canopy leaf area or shedding of older leaves) or maximize water
uptake through increased investment in the roots (Chaves et al., 2003). Changes in turgor of guard cells
relative to epidermal cells, determine the opening and closing of stomata. At any given moment, stomata
may be responding to a complex set of factors such as light intensity and CO2 concentration, in addition to
the usual leaf water status. Thus, the detailed mechanisms of stomatal response to drought remain complex
and difficult to explain. However, stomatal closure is one of the earliest known crop responses to shoot or
root dehydration (Chaves, 1991; Chaves et al., 2003) or drying soil even when shoot water status is
maintained at high turgor (Gowing et al., 1990; Schurr et al., 1992). Stomatal closure is usually mediated by
chemical signals from plant growth regulators, e.g. abscisic acid (ABA) which travels from the dehydrating
roots to the shoots. Jones (1992) reported that externally applied ABA closed stomata, levels of endogenous
ABA increased rapidly during stress and that tomato and potato mutants deficient in the capacity for ABA
synthesis were not able to close their stomata.

Easlon & Richards (2009) later noted that root drought stress or afternoon leaf dehydration may
have contributed chemical signals such as ABA and apoplast pH to induce stomatal closure before complete
loss of turgor in five self-compatible species of wild tomatoes. Chemical signal induced stomata closure
indicates a drought avoidance strategy. Manavalan et al., (2009) found that in soybean, decreased stomatal
conductance coincided with an increase in xylem ABA and occurred before any significant change in leaf
turgor was detected.

Thus, chemical signals (root-originated ABA) control stomatal behaviour at moderate soil water
deficit. Root-to-shoot signaling requires that chemical compounds or physical signals travel through the
plant in response to stress sensed in the roots. These signals may be either positive or negative. If cells
respond to changes in hydraulic pressure, then xylem cavitation itself may act as a hydraulic signal (Chaves
et al., 2003).

Reduction in leaf area and plant growth is another early response to moisture deficit for reduced
transpiration and increased water use efficiencies (Xu & Zhou, 2008). Short term reduction in leaf area as
practiced for most seedling transplants, allows plants to avoid damaging leaf water potential by reducing the
water flow through the leaf surface (Tardieu, 2005). The high sensitivity of leaf growth to water deficits is
considered to be a stress avoidance mechanism because it enables plants to tolerate severe drought
conditions by saving soil water (Lopes et al., 2011). Leaf rolling can also reduce effective leaf area and
consequently reduce radiation interception and hence reduce transpiration under water stress (Izanloo et al.,
2008). Leaf rolling or epinasty is a short-term, reversible mechanism that decreases the active leaf area. This
phenomenon is common in maize and Australian bread wheat cultivars (Izanloo et al., 2008).

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The adverse effects of drought can also be avoided by changing carbon allocation patterns to allow
the formation of a deep root system before the onset of a growth-limiting water shortage (Campos et al.,
2004). Plants can adapt to drought by developing a longer taproot to reach the lower and available water
table while an extensive fibrous root system forages the subsoil surface moisture and nutrients especially
phosphorus (Manavalan et al., 2009).

Under moderate levels of water stress, root growth - in opposition to inhibition of shoot growth
observed in the early phases of drought results from rapid osmotic adjustment that allows for partial turgor
recovery and maintenance of the ability to loosen cell walls. This ability of roots to maintain growth under
mild water stress is correlated with ABA accumulation. Thus, at low water potentials, mutants deficient in
ABA production maintained better shoot growth than root growth as compared with wild-type plants (Sharp
et al., 2004). Recent work with maize primary root suggests that endogenous ABA accumulation under
drought restricts ethylene production preventing the ethylene induced growth inhibition (Chaves et al.,
2003).

The shedding of leaves (leaf fall) is another water saving mechanism viewed as nutrient recycling in
crops through biodegradation and leaf litter mulch for soil microclimate moderation. During drought-
induced senescence, some drought–specific proteases are produced, for example cysteine proteases (Chaves
et al., 2003). Tolerant and susceptible species appear different in the degree of drought stimulation of
endoproteolytic activity (aspartic protease). The stimulation is higher in susceptible than in tolerant species
(Cruz de Carvalho et al., 2001). Ability to manipulate these processes may permit the development of plant
breeding or genetic engineering techniques for leaf senescence control under drought. Leaf senescence,
leading to short supply of sugars, is an important factor in fruit abortion during drought stress. Therefore,
delayed-senescence phenotypes may be desirable in crops where yield is source-limited and stem reserve
storage mobilization and use are insufficient to support fruit (sink) growth under stress e.g. poor filling in
plantains and bananas.

4.0 Drought Tolerance Mechanism

Drought tolerance involves maintenance of turgor through osmotic adjustment (a process which induces
solute accumulation in cell), increase in elasticity in cell and decrease in cell size and desiccation tolerance
by protoplasmic resistance (Mitra, 2001).

Osmotic adjustment (OA) is defined as the active accumulation of solutes that occur in plant tissues
in response to an increasing water deficit. It is a useful attribute because it provides a means for maintaining
cell turgor when tissue water potential declines (Manavalan et al., 2009). Osmotic adjustment is a net
increase in solute content per cell that is independent of the volume changes that result from loss of water.
Osmotic adjustment usually develops slowly (at least in shoots) and is triggered above the economic
threshold of cell water deficit.

The decrease in solute potential is typically limited to about 0.2 to 0.8 MPa, except in plants adapted
to extremely dry conditions (Taiz & Zeiger, 2002). Manavalan et al., (2009) reported that capacity for OA in
soybean ranges from 0.3 to 1.0 MPa while in chickpea (Cicer arietinum) and pigeon pea (Cajanus cajan) it
ranges from 0.1 to 1.3 MPa. Most of the adjustments can usually be accounted for by increase in
concentration of variety of common solutes including sugars, organic acids, amino acids and inorganic ions
(especially K+).

However, the accumulation of ions during osmotic adjustment is restricted to the vacuoles, where
the ions are kept out of contact with enzymes in the cytosol or sub cellular organelles. Cytosolic enzymes of

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plant cells can be severely inhibited by high concentrations of ions. As a result of this compartmentalization
of ions, other solutes accumulate in the cytoplasm to maintain water potential equilibrium within the cell.
These compatible solutes (or compatible osmolytes) which are organic compounds scarcely interfere with
enzyme functions.

Commonly accumulated compatible solutes include the amino acid proline, sugar alcohols (e.g.
sorbitol and mannitol), and a quaternary amine called glycine betaine (Taiz & Zeiger, 2002). Some
hydrophilic proteins and carbohydrates (fructans and sucrose) are also synthesized.

In addition to decreasing cell osmotic potential, these solutes provide protection to the cell
membrane and the metabolic machinery under dehydration, by interacting with water molecules rather than
by interacting with proteins. Osmoprotection mechanisms are probably not functional until severe
dehydration occurs with the implication that osmotic adjustment may be critical to survival rather than to
increase plant growth and crop yield under drought (Chaves et al., 2003). In rice, for instance, drought
tolerance correlates with proline and total soluble sugar (TSS) accumulation ability. Drought-tolerant rice
lines accumulated up to fifteen fold of proline content after five weeks of dehydration stress, while the
original line, RD23 increased approximately five folds (Vajrabhaya et al., 2001). Similarly approximately 4
fold increases in TSS content was detected in the tolerant lines while only 2.5 fold accumulation was
observed in the original line. Thus, the proline, besides playing a role in osmotic adjustment, is also involved
in the protection of the enzymes and cellular structure and acts as a free radical scavenger.

In Sahel regions, many of the evergreen shrubs and trees combine the high solute concentration in
living cells, low photosynthetic capacity and stomatal conductance. Such small leaves are well adapted to
high light and temperature that prevail in most Sahel regions. The small size of leaf permits greater sensible
heat dissipation and an efficacious control of water loss by stomatal closure (Chaves et al., 2003).

4.1 Other Mechanisms of Crop Adaptation to Moisture Stress

Partial dormancy is another means through which some plants survive drought, as observed in Retama
raetam, (a stem-assimilating, C3, evergreen desert legume common to arid ecosystems). From acclimation
studies Mittler et al., (2001) observed that Retama raetam contain two uniquely different populations of
stems: those of the upper canopy, exposed to direct sunlight, and those of the lower canopy, protected from
direct sunlight. During the dry season, stems of the upper canopy contain very low level of a number of
essential proteins. However, RNA encoding these proteins was present.. Upon water application, as well as
following the first rainfall of the season, these ‘photosynthetically suppressed’ stems recover and accumulate
essential proteins within 6 to 24 hours. In contrast, stems of the lower canopy contain the essential proteins
throughout the dry season. Thus, R. raetam uses an acclimation strategy of partial plant dormancy to survive
the dry season.

In C4 plants possessing crassulacean acid metabolism (CAM) photosynthesis, metabolic adaptations


to drought occurs in a most intriguing manner. The CO2-fixation process in C4 plants is separated in
mesophyll cells and the bundle sheath cells. Although reduced CO2 concentration may have been the major
reason for the evolution of C4 plants, this specialized photosynthesis brought about greater water use
efficiency and ecological success in arid environments (Chaves et al., 2003).

In CAM plants, CO2 is concentrated during the night into C4 acids re-fixed to carbohydrates during
the day, in a process similar to C4 plants. By switching the pattern of CO2 uptake through the stomata to the
night period and by fixing CO2 into carbohydrates during the day (with closed stomata), these plants greatly
increase their water use efficiency, thereby improving their chances of survival in extremely arid

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environments. This mode of adaptation of photosynthesis to drought is more interesting in inducible CAM
plants. Generally, these plants operate under a C3 mode of photosynthesis, but once exposed to dehydration
stresses, switch from the C3 mode to the more water-use-efficient CAM photosynthesis (Chaves et al.,
2003).

5.0 Conclusion

Stress due to moisture deficit is a common limitation to crop production especially in Sahel regions.
Consequently, most crop species and varieties grown in the sub Saharan Africa have developed adaptations
to moisture stress. However, more areas including humid regions are becoming more prone to moisture
stress as a result of the current change in global climate which has diverse implications across the
agroecologies in Nigeria. The use of drought tolerant species and varieties is an inexpensive means to
overcome this limitation. This may require development of new drought tolerant varieties for specific areas
or improvement of existing drought tolerant varieties. For this purpose, physiological understanding of the
mechanisms underlying crop adaptation to moisture stress is critical to facilitate the crop improvement
process.

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