The Mitigation of Drought Stress

By Dr. Abraham Blum
PlantStress.com

 

 

 BY CROP MANAGEMENT

Drought management by irrigation

Drought management in dryland

 

 BY CROP DROUGHT RESISTANCE

Nature of drought resistance

Breeding for resistance

NOTE: Numbers in parenthesis throughout text are ID number for searching that reference

in the Reference Database on this site. Sometimes it is also a live link to the report.

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

MITIGATION OF DROUGHT STRESS BY CROP MANAGEMENT

 

Managing Drought Stress by Supplemental Irrigation


Irrigation, where economically available, is the major means for combating drought conditions. It is a prime approach to the intensification of agriculture and the generation of stable income. The development of irrigation depends on various environmental, economical and social factors on both the macro and micro scales.

There are hazards in irrigation if practiced indiscriminately, such as soil erosion, soil salination, soil leaching and soil disease infection. Irrigation as such is not an important topic in this site. Links to irrigation sites throughout this section should provide additional information.

 

General Crop Irrigation Guidelines

 

The key to planning irrigation system and scheduling is knowledge of the crop, the soil properties and the potential evapotranspiration

General Irrigation Links

Irrigation Manuals

WWW Virtual Irrigation Library

 (PET) of the specific crop at the site. This information can also be used to estimate dryland crop water use and deficit at any given time during the crop cycle, which is actually an index of crop drought stress.

The Penman-Monteith potential evapotranspiration equation is recommended by the FAO as the standard method for estimating reference and crop evapotranspiration. The new method has been proved to have a global validity as a standardized reference for grass evapotranspiration and it has been recognized by both the International Commission for Irrigation and Drainage and by the World Meteorological Organization.

The (FAO) Penman-Monteith method was developed by defining the reference crop as a hypothetical crop with an assumed height of 0.12 m having a surface resistance of 70 s m-1 and an albedo of 0.23, closely resembling the evaporation of an extensive surface of green grass of uniform height, actively growing and adequately watered.

Further educational information and guidelines on the applications of the Penman-Monteith method and the general approaches to prediction of crop water requirements and the supply of requirement is provided by:

The FAO Methodology for Crop Water Requirements.

FAO CROPWAT, downloadable software to carry out standard calculations for design and management of irrigation.

 

Irrigation to Control Drought in Various Crops

 

Supplemental irrigation is the more common irrigation practice for crops not designated a priori for fully irrigated conditions. It is also recently being labeled as "deficit irrigation" since it does not supply the full seasonal crop water requirement. Supplementary irrigation is a practice dictated by constraints, which can be derived from the limited availability of water, irrigation equipment, the cost of water, or other economical and technical constraints. With supplemental irrigation the amount of water applied to the crop in irrigation is well below the full requirement of the crop. The water-use efficiency of supplemental irrigation is generally high if applied logically. It can be applied to save the crop in case of un-expected drought or as a planned practice to supplement the expected total seasonal rainfall. The practice may vary extensively with crop and region. In many environments, and especially the Mediterranean region, if only a single supplementary irrigation is given it is usually more effective if applied pre-planting. As such the crop enters the season with a stored supply, which can insure growth despite unexpected transient rainfall fluctuations.

There are numerous computer models based on simulations of crop growth combined with field water balance computations (see above), which allow the grower to input his environmental data set in order to develop recommendations for irrigation at different scenarios.

 


Managing the Dryland Crop Environment

Modern dryland farming is a system of low inputs combined with soil and water conservation practices and risk reducing strategies. The system can be sustainable if practiced properly.  Water shortage is the main limiting factor, but successful dryland systems also maintain reasonable practices to eliminate other limiting factors (poor nutrient status, weeds, biotic stresses, etc'), which can reduce the efficiency by which the crop uses the limited moisture. However, as water shortage a priori dictates a limit on yield, all other inputs must be carefully adjusted downwards to fit the expected low economic return. Except for subsistence agriculture, economical dryland farming depends on a large area per farmer.

The most advanced systems have been developed in the Great Plains of the USA, Canada and Australia, even though traditional systems employed in Asia and the Middle East also offer important insights. In the USA, the lesson learned during the "dustbowl" years in the early 1900's prompted extensive legistration and investments in developing sustainable dryland farming systems. These systems and the associated technological progress such as plant breeding, brought about an increase in mean winter wheat yield from 0.5 ton ha-2 in 1930 to about 2 ton ha-2 in 1980. In Southern Australia the "ley farmimg" system was developed in the 1920's and adopted widely in the late 1940's. The system involves a rotation between a self-seeding legume grown for several years and wheat. The farmer grows wheat and raises sheep while the legume serves to sustain soil fertility (mainly nitrogen). This system has become less popular in recent years with the increase in economic pressures and other considerations.

The lesson learned from the American and Australian experience is that the development of a sustainable dryland farming system involves the following principles, not necessarily in their order of importance:

1.     Improved soil and water conservation practices and the associated tillage systems.

2.     Optimization of the fit between crop growth cycle and the available moisture.

3.     Weed control

4.     Soil fertility management.

5.     Optimized plant population density and spatial arrangement of plants with respect to the expected soil moisture regime.

6.     Control of soil biotic stress factors that reduce root development.

7.     Improved forage/livestock/grains integration and rotation.

8.     Avoidance of mono cropping and the diversification of farming.

9.     The increase of precipitation by cloud seeding, as an ongoing experiment.

 

Some of the above principles of the dryland farming system constitute general agronomic knowledge, which can be explored in our Web Resources page as well as in standard agronomy textbooks and other publications (e.g. Drought Management of Farmland - by Joan Sydney Whitmore, 2000, 360 pp., Springer, SBN 0792359984). Here only several topics will be touched upon.

 

Soil and water conservation

 

Fallow and conservation tillage

 

The fallow system is designed to conserve soil moisture from one season to another or from one year to the other, depending on climate and crop. Increasing storage of soil moisture by the fallow system with or without conservation tillage is standard agricultural practice in dryland farming. The benefit of fallow and conservation tillage in terms of increasing available soil moisture to the crop depends on soil water-holding capacity, climate, topography and management practices. Fallow efficiency, in terms of percent increase in soil moisture availability to the crop measured at planting date normally ranges from about 5% to 30%. While these amounts are not impressive they can make a difference between crop failure and success. The fallow carries additional benefits such as improved soil nutrients availability and the eradication of certain soil-born pests, such as nematodes.

Conservation tillage involves the principle of minimized tillage operations to conserve soil structure and to maintain ground cover by mulch, such as stubble. These practices reduce water runoff and increase soil infiltration. Conservation tillage has become the cornerstone of dryland systems in certain regions of the USA, Canada and other regions. It has been re-demonstrated in dryland wheat experiments carried out in Southern Israel (#4407). While the benefits of conservation tillage are well-documented it has also been noted that crop residues under this system may promote certain crop diseases.

In certain soils deep tillage was found very useful to improve soil moisture storage, especially when hard soils or hardpans are a problem. This is an expensive operation that cannot be deployed regularly in dryland farming.

 

For further online information on dryland farming and its research visit the following sites and the links therein:

§       Soil and Water conservation in Semi Arid Areas – (FAO Manual, 1987)

§       USDA-ARS Bushland Texas Experiment Station – dedicated to conservation research

§       Soil, water and reclamation publications --From Alberta, Canada

 

Furrow dikes and Soil pitting

 

These techniques constitute a field surface tillage manipulation to minimize runoff away from the field.

Furrow dikes are furrows, which are divided into short basins by small dikes (see right side photograph). This is achieved by special equipment. The system is very amenable to row crops such as cotton, corn and sorghum and it can be integrated with or without furrow irrigation. It is generally considered effective for increasing rainfall capture and raising dryland yield where annual rainfall ranges between 500 and 800 mm.

Soil pitting (left side photograph) involves the formation of small depressions at close proximity to reduce runoff from rainstorms. The crop is planted over this modified surface. Experiments performed with wheat in nine farmer demonstration plots in Southern Israel during 1988 showed that pitting increased yield by an average of 7.5% at a mean yield of 3.25 ton/ha. Unlike furrow dikes these system is not limited to row crops.

 

Water harvesting/spreading

 

This is a broad term to describe various methods to collect runoff from large contributing areas and concentrate it for use in smaller crop area. This is an ancient practice already adopted by Nabatian desert settlements in the Middle East several centuries A.D. Presently, the basic water harvesting systems involve an external contributing area to induce runoff. This area is physically or chemically treated for maximizing runoff. The water is diverted into a receiving area comprising of cultivated plots, individual trees or small terraces. The contributing area may lie in the agricultural field (a system sometimes referred to as "conservation bench terrace") or outside the field in the natural watershed system. The size ratio between the contributing and the receiving areas is determined by the expected rainfall events, crop water requirements, soil characteristics and topography. The resulting yield increase in the receiving (crop) area is proportional to the amount of water gained in that area.

For more detailed information see ‘Water Harvesting, and Runoff Farming.

 

Diversification of farming

 

Diversification of farming is an ancient but an effective approach to reduce the risk associated with farming in unpredictable environments. Reduced diversification to the extent of mono-cropping is possible only if a high level of control is possible over the crop environmental conditions. Such control method (irrigation, chemical pest control, etc’) are among the main reasons for the more recent environmental quality problems found to be associated with mono-cropping.

Diversification of cropping to reduce risk is especially important under dryland conditions. It is achieved on several levels, as described by Pandey et al (#4194) for the case of traditional rain-fed rice in Eastern India.

1.     Spatial diversification of fields. The farmer’s land is divided into several fields or plots which may differ in their topography, soil and hydraulic properties. Some fields may be prone to flooding while others do not hold water. Certain fields may be on a warmer slope while others on a cooler one. The different field conditions allow to achieve a better fit between the crop and the environment and to reduce the general probability of stress affecting the farmer.

2.     Crop diversification is an important feature of traditional farming. It takes an advantage of the generally low correlation between crops in performance when grown in a single stress environment. Crops differ in their response to a given environment and this difference is used to reduce the risk associated with growing one crop. “Mixed cropping” or “intercropping” is an example of a traditional and a successful approach to crop diversification on a single parcel of land, where two or more crops are grown together in various possible configurations. If for some reason only one crop is grown, a certain (though lower) level of risk reduction can be achieved by varietal diversification. Planting of several crop varieties offer a better probability for reducing loss due to environmental stress, as compared with growing one variety only. For environmental stress conditions varietal diversification is based mainly on differential phenology, primarily flowering date. A typical example is a transient frost or heat wave that is likely to occur around flowering time of the specific crop. Damage reduction can be achieved when the crop is sown to several varieties of different flowering dates.

3.     Temporal diversification may achieve the same result as varietal diversification, when phenology is concerned. The purpose of setting a distinct planting date is to optimize crop development with respect to seasonal climate, mainly rainfall in rain-fed agriculture. Ideally the crop is planted at the beginning of the rainy season, rainfall peaks when crop evapotranspiration peaks and it terminates just before harvest time. When such conditions are reasonable predictable, planting date can be set to optimize production. Where the timing of rainfall is very unpredictable, adopting more than one planting date for the given crop can reduce the risk involved with untimely rainfall and a given planting date.

 

Cloud seeding

 

Cloud seeding is a form of weather modification attempt. The process of cloud seeding involves deposition of cloud condensation nuclei (CCN) into a specific region of the cloud. Seeding may be achieved from above or through the clouds by aircraft, and from below where CCN are carried into the cloud by updrafts. With either method, the CCN must reach the super cooled cloud region, where water molecules remain unfrozen at temperatures below 0C.

Experiments in cloud seeding have been performed for the last 50 years. The results and benefits of this practice are still under debate.

Information is available in the report on 'Weather Modification by Cloud Seeding-A Status Report 1989-1997 by William R. Cotton, Colorado State University; and at the Oklahoma Weather Modification Demonstration Program.



MITIGATION OF DROUGHT STRESS BY CROP PLANT RESISTANCE

 

The nature of Drought Resistance

 

Drought Resistance and Crop Yield

Crop plant breeding for drought resistance has long been part of the breeding process in most crops that have been or are being grown under dryland conditions. During the period of the pre-scientific agriculture the genetic improvement of plant adaptation to dry conditions was simply attained by repeatedly selecting plants that appeared to do well when drought stress occurred. As a result of many generations of selection by generations of farmers we now encounter such materials, which are defined as “landraces” of the crop. Such landraces were shown to posses distinct drought resistance traits. Later, as scientific agriculture developed and following the emergence of Mendelian genetics, elaborate biometrical and statistical methods for quantitative genetics analysis were developed to enable selection for yield stability. An important factor of yield stability is coping with drought and other abiotic plant stresses. Subsequently, yield-based selection programs were augmented by observing plants under carefully managed stress environments, followed by the development of physiological selection criteria for stress resistance. More recently, molecular methods, such as marker-assisted selection are being adopted to facilitate more efficient selection for distinct components of stress resistance. Finally, biotechnology is experimenting with genetic transformation, which has not yet been applied as a solution to breeding for drought resistance. Some applications may be forthcoming in the private seed business sector.

 

Looking at crop drought resistance from a botanical perspective it must be realized at the onset that there is a vast difference between drought resistance in natural vegetation and in crop plants. Natural vegetation has evolved to conserve the species. Henceforth, plant survival and the capacity to produce at least one seed per life cycle despite stress is the most powerful component of natural selection. On the other hand, drought resistance in modern agriculture requires sustaining economically viable plant production despite stress. Crop survival is of a lesser consequence to economical farming. On the other hand, plant survival can be a critical factor in subsistence agriculture, where the ability of a crop to survive drought and produce some yield at all may translate into a difference between famine and livelihood. Breeding for drought resistance is therefore very tightly linked to the target environment of the crop, not only with respect to its physical and chemical features but also its social grounds. The recent developments in GIS technology are extremely important as a tool for defining a target environment for the breeding program. GIS has been discussed under impact of stress above.

 

John B. Passioura’s (#6030) general description of yield and water-use is widely accepted by agronomists. It is based on the initial model of C.T. de Wit and in its simple form it can be expressed as:

 

Yield = T x WUE x HI,

 

Where, T= total seasonal crop transpiration, WUE= crop water-use efficiency and HI= crop harvest index (the ratio of economic yield to total aboveground biomass). T is a component of ET (crop evapotranspiration), where E denotes direct evaporation from the soil.

It can be immediately seen that yield is proportional to crop water-use. The older concept of formulating drought resistance by designing a plant that is “water-saving” will not work for most crop environments. Moderated water-use by design can be important in systems where the crop grows mostly on pre-seasonal stored soil moisture. Here, fitting crop-water use to the limited available seasonal water supply is crucial.

Text Box: Fig. 7. The association between yield and total seasonal precipitation for 3 different wheat cultivars.

 Generally, crops must use water under stress in order to sustain yield. Drought resistance is a finite trait. Life cannot be sustained without water. Drought resistance in modern farming is based on the plant ability to obtain water or to use water efficiently, when water is limited by drought.

 

A most preoccupying issue for the dryland crop breeder, and not only the beginner, is the formulation of a plant ideotype for the target environment addressed by his program. The ideotype is a detailed morphological, physiological, developmental and conceptual description of the desirable plant type. The construction of an ideotype can be based on accumulated knowledge, consultation, biased opinion and even intuition. Lately, serious attempts are made at adopting crop growth simulation models to assist in developing a breeding ideotype. The formulation of an appropriate and relevant ideotype for a drought target environment is probably the most crucial component of planning a breeding effort for improved drought resistance. It is hoped that this contribution will help to understand what is involved in formulating a drought resistant ideotype and hopefully assist in applying such knowledge to practice.

In order to further understand what constitutes drought resistance in terms of crop production, one should closely examine how different cultivars may respond to changing moisture availability (Fig.7).

Wheat cultivar C is different from A and B in that it has a lower yield potential (yield at high moisture conditions) but as moisture becoming deficient C turns out to be superior to A and B. In terms of yield, C may be defined as drought resistant while cultivars A and B are of high yield potential but are relatively drought susceptible. The “crossover” where the advantage of C over A and B under stress begins to be expressed is at about 300 mm or at a yield level of about 300 g m-2. Hence, drought resistance of C is expressed only when stress is sever (<300 mm). Still, it is extremely important to realize that the high yielding cultivars A and B are superior to the drought resistant C when drought stress is moderate (e.g. at 400 to 500 mm). A high yield potential therefore ascribes an advantage under moderate stress conditions. On the other hand Fig.7 seems to indicate that drought resistant wheat cultivars have lower yield potential, which in a sense is a penalty for drought resistance. This point will be discussed later.


 

Constitutive plant traits affecting drought response


As discussed above under the “Impact of Stress”, moisture stress signals certain stress responsive genes, which are responsible for a chain of events, expressed at various levels of plant organization. It has been assumed almost axiomatically that stress responsive genes are involved in adaptation; henceforth that they are ‘stress adaptive’. It was later realized that not every stress responsive gene is necessarily adaptive in terms of drought resistance or survival.

Irrespective of the role and function of stress adaptive genes in plant drought resistance, it should be recognized that not only stress adaptive genes condition plant performance under drought stress. Genes that are expressed irrespective of the environment also condition plant function and performance under stress. These genes are expressed constitutively. For example, genes conditioning early flowering are basically expressed in any environment. Their expression is not conditioned by stress. Early flowering may have a critical role in plant response to and performance under stress (see below). Constitutive plant traits can be modified by stress, and the rate of modification can be considered to involve stress responsive elements. Using the above example, stress may cause one early variety to flower a little later than normal while another similarly early variety will not be affected by stress and will maintain its normal flowering time. The delay in flowering may be generally considered as un-desirable and an expression of drought susceptibility.

 

Plant Phenology

 

 

Short growth duration (generally defined by early flowering) constitutes an important attribute of ‘drought escape’, especially for conditions of a late-season drought stress (Fig.8). On the other hand, longer growth duration is often associated with high yield potential. Consequently, using drought escape as a solution may involve a cost in terms of reduced yield potential. This is serious, especially when the moisture environment is unpredictable and may vary to a large extent between years. The more predictable the environment is, the easier it becomes to optimise phenology. The unpredictability of the environment may reach a state where short growth duration is a drawback, especially in indeterminate plants that offer a potential for regrowth and productivity upon recovery. A longer growth duration in both determinate and indeterminate plants would improve the probability for regrowth upon recovery simply because, on the same calendar day, late-maturing genotypes are younger than early ones and younger plants recover better. The final decision on the optimum growth duration has, of course, to consider additional factors, such as late-season disease and insect pressure or periods of frost.

Early maturity lead to reduced, total seasonal evapotranspiration simply because of the shorter time in the field. However, as growth duration is genetically linked with leaf number, early genotypes have a small transpiring leaf-area index. Thus, early genotypes show reduced evapotranspiration during most growth stages, up to the point where a full ground cover is achieved. At most growth stages, root-length density and total root length per plant is generally greater in a late than in an early cultivar. This should be reflected in an advantage for the late genotype under conditions where extensive rooting is required.

A phenological trait specific to maize is the timing of anthesis with respect to silking, defined as anthesis-to-silking interval (ASI). Evidently a short interval is desirable whereas a large interval results in poor pollination. The maize program at CIMMYT dedicated many years of work to research the trait and explore its significance in tropical maize breeding for stress environments (#3347). Maize germplasm can vary for ASI irrespective of the effect of stress; a short ASI is a universally important trait for maize production. However, stress, and especially drought during the reproductive stage may extend ASI and thus reduce yield. Maize genotypes may vary in ASI under drought stress from few days up to around 60 days. The effect on yield of change in ASI between null and 10 days is exponential. Selection for short ASI under drought stress proved to be the most effective approach to improve drought resistance of tropical maize. QTLs (quantitative trait loci) controlling ASI were identified and marker-assisted selection is possible.

Different crop plants may advance (e.g. wheat) or delay (e.g. rice) their flowering when stress occurs before flowering. The rate of delay is a function of plant water deficit and probably also ABA signalling. The rate of change in flowering time under stress can be taken as an index of genotypic rate of stress in the field.

 

Plant development and size

 

Plant size as expressed mainly in terms of single plant leaf area or leaf area index (LAI) has a major control over water-use, as explain under Impact of Stress. Small plants and reduced leaf area are generally conducive to low productivity while they limit water use. Botanists have long recognized small plants bearing small leaves as typical ecotypes of xeric environments. While such plants withstand drought very well their growth rate and biomass are relatively low.

In the domain of plant breeding, cultivars developed for dryland conditions by selecting mainly for yield under such conditions often result in plants of moderate size and productivity. For example this can be seen in dryland temperate cereals, which tend to have moderate tillering. On the other hand researchers in the CSIRO Australia have concluded (#3903) that early plant (and seedling) vigour are important traits for dry conditions. The reason is in the rapid ground cover achieved and the subsequent decrease in water loss by direct soil evaporation at this stage. However, other benefits for seedling vigour were also noted, such as the nitrogen status of the plant (#7059).

As for other constitutive traits, irrespective of the genetic control of plant size, drought stress signals a reduction in leaf area to effectively reduce water use. Drought stress reduces the transpiring leaf area by leaf expansion inhibition, leaf wilting (such as leaf rolling in the cereals) and leaf death and abscission. While leaf desiccation is caused by dehydration, leaf growth inhibition and abscission can result at least partly from stress induced ABA accumulation.

 

The Root

 

The most important control of plant water status is with the root, whereas roots are the main engine for meeting transpirational demand. Two major dimensions describe the root: root depth and root-length density (Fig.9). The more practically important dimension for most breeding scenarios is root depth, which facilitate deep soil moisture extraction where such moisture is available. It is a primary component of drought resistance. The development of lateral roots at very shallow soil depth may have a role in capturing small amount of intermittent rainfall.

Text Box: Fig. 9. Left panel: hydropnically grown roots of two wheat cultivars differing in root length. Right panel: roots of two sorghum cultivars in soil in a root observation box, differing in root-length density. See “sorghum root growth”

Root depth in the cereals is generally associated with a small number of main thick axes. Such fibrous root system is often seen in upland rice, which has a deeper root system, in contrast to lowland rice with the shallower roots. The control of root growth is not only in the root. In the cereals, tillering is associated with production of new crown roots from each developing tiller. Such profuse rooting is at the expense of the growth of existing roots, deeper into soil. Hence, limited tillering in cereals and grasses has been repeatedly observed to be associated with relatively deeper root extension.

In certain soils a hardpan can limit deep root growth and the capacity for hardpan penetration by roots becomes a critical factor in drought resistance. The factors, which may support axial root force and hardpan penetration, are not known and most research in this area has been performed with seedlings only. In mature plants the penetration of hardpan by roots seems to be better in plants that constitutively develop fibrous and thick roots.

Many drought environments present a situation where rainfall is low and soil depth that contains moisture is permanently shallow. For example, in many of the drier Mediterranean wheat-growing regions the wetted soil depth of around 60-80 cm is shallower than the maximum root depth of wheat (>100 cm). Under such conditions a deep root is not an issue. Other plant factors may then become far more important in the control and use of the limited soil moisture, such as shoot developmental characteristics (e.g. leaf area development or growth duration), osmotic adjustment, leaf surface properties, etc.'

Another scenario of seasonal soil moisture status is where the crop is grown on stored soil moisture and there is little effective rainfall during the growing season. Under such conditions the main consideration is to manage seasonal soil moisture use such that sufficient moisture will remain for carrying the crop to maturity. It is to be expected that with the available moisture the crop might grow luxuriously leading to a large leaf area and an even greater water requirement towards the latter part of the season. Hence, short growth duration, small leaf area and perhaps a higher root hydraulic resistance can achieve the control of seasonal water use. The last option has been researched at the CSIRO Australia and an increase wheat root hydraulic resistance was effectively attained by selection for smaller xylem element diameter. It is not known whether this approach has found its way into actual application in wheat breeding

Whatever may be the constitutive form and function of roots, the environment can modify the root in a pronounced way. Offcourse, soil conditions in terms of topsoil moisture and deep soil hardness alter root growth and depth. Drought stress generall