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Asati, R.;  Tripathi, M.K.;  Tiwari, S.;  Yadav, R.K.;  Tripathi, N. Drought in Chickpea. Encyclopedia. Available online: https://encyclopedia.pub/entry/36702 (accessed on 20 April 2024).
Asati R,  Tripathi MK,  Tiwari S,  Yadav RK,  Tripathi N. Drought in Chickpea. Encyclopedia. Available at: https://encyclopedia.pub/entry/36702. Accessed April 20, 2024.
Asati, Ruchi, Manoj Kumar Tripathi, Sushma Tiwari, Rakesh Kumar Yadav, Niraj Tripathi. "Drought in Chickpea" Encyclopedia, https://encyclopedia.pub/entry/36702 (accessed April 20, 2024).
Asati, R.,  Tripathi, M.K.,  Tiwari, S.,  Yadav, R.K., & Tripathi, N. (2022, November 28). Drought in Chickpea. In Encyclopedia. https://encyclopedia.pub/entry/36702
Asati, Ruchi, et al. "Drought in Chickpea." Encyclopedia. Web. 28 November, 2022.
Drought in Chickpea
Edit

Chickpea productivity has been found to be around 995 kg ha−1 on a global scale, which is quite low. Drought, terminal heat, excessive salt, and cold are abiotic variables. Several factors are responsible for complexity of drought stress, including severity of drought, stage of crop, and duration of drought stress.

abiotic stress candidate genes drought tolerance

1. Introduction

Chickpea is a diploid annual crop that is extremely self-pollinated [1]. After the faba bean and field pea, it is the world’s third most significant pules crop [2]. It is a popular cool-season legume crop with a 738-megabyte genome size [3]. With an annual production of 10.13 million tonnes from a land area of 9.44 million hectares and a productivity of 1073 kg ha−1, India is the greatest producer of chickpeas in the world [4]. Chickpeas are grown in 52 countries, together with Africa, Asia, Australia, and South Europe [5]. Mexico, Turkey, Canada, Iran, Australia, Tanzania, Ethiopia, Spain, and Burma are also notable producers of chickpea. Its seeds come in two varieties. The ‘desi’ chickpea is hardy in character, while the Kabuli chickpea has a delicate seed coat and appears to have evolved from the desi varieties [6][7]. In semi-arid zones, chickpea is cultivated in the form of a dry weather crop [8]; however, in cold climatic zones, it is grown as a rainfed crop [9][10]. In actuality, about 90% of the chickpea crop is cultivated in a rainfed environment [11][12][13]. Without irrigation, the crop is affected [14] at vegetative as well as reproductive phases. After illnesses, drought is the second most significant constraint to the yield of chickpea crop [15]. Drought has been reported as a factor of 40–50 percent yield reduction in chickpea [11][12][16][17][18].
The chickpea is also termed as the “poor man’s meat” [19], since it is important for supplying protein sources [20]. Nutritionists have also highlighted its importance due to high nutritional contents in it [21]. Chickpea is high in lysine and arginine [22], but low in methionine and cystine [23]. In general, the Kabuli type contains more protein than the desi kinds. It contains more calcium and phosphorus than most other pulse crops [24][25]. Chickpea seeds comprise 23% protein, 64% total carbohydrates (47% starch, 6% soluble sugar), 5% fat, 6% crude fibre, and 2% ash on average, as well as micronutrients, for example phosphorus, calcium, magnesium, iron, and zinc [26]. Recently, Singh et al. [21] also reported chickpea as good source of Fe and Zn. Consequently, Samineni et al. [27] examined the effects of drought stress on nutritional parameters of chickpea and observed significant differences in the nutritional contents due to stress.
Chickpea is mostly cultivated in the post-rainy season [28], using soil moisture that has been retained from the previous rainy season [29]. As a result, the crop is frequently subjected to severe heat and drought pressure [12][13][22]. Drought, among other abiotic factors, has a significant impact on chickpea output [30]. Drought and heat stress have been reported to have reduced chickpea yields by about 50% due to the damaging effects of the membrane and reduced photosynthesis [31].
The four climatic elements that are changing will have an impact on how much water plants consume [32]. These elements include rising CO2 concentrations and temperatures, more erratic precipitation, and changes in humidity. Due to the increased variability in precipitation during the growing season and more so in soils with low water holding capacity, these climate changes may result in an increase in the atmospheric water demand by crops and an increase in the potential for limitations in the availability of water in the soil. In the long run, breeding cultivars with high water use efficiency (WUE) is a more realistic and cost-effective strategy for raising yields in drought-prone locations. WUE promotes modest water absorption while maintaining elevated WUE, which is a crucial component of breeding programmes because of its yields in drought-prone areas. Any WUE is impacted by changes above the soil surface because they have an impact on the soil water balance by the evaporation and penetration of soil water. The majority of the chickpea crop is grown on residual moisture; however, additional irrigation can increase yields. At some sites in India, irrigation during the pre-flowering stage and at the beginning of the pod fill led to an increase in yield. Chickpeas’ reproductive cycle was prolonged by irrigation, which also increased plant biomass and increased the number of pods per plant.
The greatest sustained surface winds of tropical storms range from 39 to 73 mph, and they are fast rotating storm systems with an organized centre over warm tropical oceans. These storms have a wide range in size and can cause a variety of dangers for the impacted areas, including tornadoes, catastrophic winds, coastal floods, and inland flooding. The effects of tropical cyclones on drought have been extensively studied, but less research has been conducted on how smaller tropical storms affect the severity of drought. According to research, rainfall is not necessarily inversely correlated with the strength of a tropical cyclone; therefore, tropical storms can sometimes provide more rain than expected. The question of whether tropical storms can help to lessen and mitigate drought conditions is now being researched. Water deficit and surplus are related to drought and tropical storms (TS), respectively. When it comes to monitoring dryness, soil moisture is a crucial element of the hydrological cycle, since it reflects the water that TS rainfall has penetrated or stored. Soil moisture data can be used to determine whether TS can alleviate extremely severe drought situations [33]. The authors calculated the frequency of TS afflicted places in the US, including the ratio of droughts that TS exacerbated and alleviated, and the regions where TS have a significant impact on the offset of drought. Based on a high-resolution data set, the findings demonstrate extensive spatial information about the offset of drought conditions and offer potential guidance for future drought and TS mitigation.
Drought has a substantial influence on crop growth and photosynthesis, both of which are directly related to production [34][35]. Drought researchers must assess growth as well as physiological responses such as chlorophyll index [36], relative water content [37], membrane stability index, and biomass when determining the influence of drought on various crop metrics. The quantitative character of attributes and the prevalence of linkage between desired and undesired genes make developing drought-tolerant agricultural variants difficult [38]. Many experiments on the effects of drought on numerous chickpea features, such as root attributes, shoot biomass, and early maturity, have been conducted [39]. In this crop, various experiments have been performed successfully and published with specific conclusions on different aspects, such as morphological, physiological, biochemical, and molecular characteristics [40][41][42].
Advances genomics has made it possible to tag genes [43] associated to agronomic qualities, as well as the tolerance/resistance to abiotic and biotic challenges [44]. It is playing a significant role in the transfer of labelled genes through molecular breeding [45][46], quickly and accurately. In chickpeas, microsatellite and sequence-tagged microsatellite site markers have been found to be more beneficial [47][48]. As stress resistance/tolerance is governed by numerous genes [49], quantitative trait loci (QTL) mapping has proven to be effective in identifying and tagging the genes [50] involved for disease resistance/tolerance in plants. Foreground, recombinant and background selection are all examples of marker-assisted backcrossing. Linkage disequilibrium (LD) and association mapping are also determined using markers [51].
Next-generation sequencing (NGS) is a segment of revolutionary biology being a frontier area in crop science and produces correct data, with the results of significant throughput [52] and reduction in the need for fragment-cloning processes, which were the initial requirement for Sanger sequencing. NGS is used for the identification and mapping of mutations in targeted genotype [53][54]. Aside from whole genome sequencing (WGS), NGS also provides a platform for whole transcriptome shotgun sequencing, which is also termed as RNA sequencing (RNA-seq) [55][56] and whole-exome sequencing [57], which exhibits for functional variations [58], targeted or candidate gene sequencing [59].In the examination of large numbers of samples, RNA-seq enables a more precise and sensitive measurement of gene expression levels than microarrays [60].
Transcriptomics is the technology used to study the transcriptome of an organism [61]. Transcriptome is the complete set of genes [62] expressed under specific conditions by the genome of the targeted organism. MicroRNA (miRNA), transfer RNA (tRNA), messenger RNA (mRNA), ribosomal RNA (rRNA) and other non-coding RNA are all found in the transcriptome (ncRNA). Transcriptomics of chickpea [63] has provided insight into mechanisms of drought tolerance/avoidance, as well as pathogenesis-related and developmental processes [64]. Transcriptomics may undoubtedly have a greater impact on chickpea breeding in the future, including the use of microarrays.
Proteomics is the study of whole protein complement in a cell, tissue or organism in detail [65]. Mass spectrometry and protein microarrays can be used to analyse the proteome [66].
Role of various genes of plants under drought stress conditions have been recognized clearly [67]. Drought responsive mechanisms are activated in response to drought stress, which is a regular occurrence in plants. Morphological and structural changes [68], drought-resistant gene expression, hormonal and other biochemical changes are among these pathways [69]. Environmental stresses have the ability to change the developmental behaviour of plants. These alterations in plant growth and development [70][71] mostly resulted in lower yields [72].

2. Drought in Chickpea

Chickpea productivity has been found to be around 995 kg ha−1 on a global scale, which is quite low [73]. Drought, terminal heat [74], excessive salt, and cold are abiotic variables [75], whereas Ascochyta blight, Fusarium wilt, and Helicoverpa are biotic factors that have been recognised as key drivers of yield reduction in chickpea [76]. Drought stress was identified as a major cause in around 50% of chickpea output losses worldwide.
Several factors are responsible for complexity of drought stress (Table 1), including severity of drought, stage of crop, and duration of drought stress [77]. Two types of drought stresses, i.e., terminal and intermittent, have been reported with their impacts on crop plants [78]. During terminal drought, soil water availability diminishes over time, potentially leading to severe drought stress later in crop development. Intermittent drought is defined as a series of short episodes of insufficient rain or irrigation that occur at different times during the growing season [79]. Due to its limited cultivation on marginal terrain, chickpea is suffering from terminal drought stress. Intermittent and terminal drought stress is caused by breaks in rainfall combined with less moisture in terminal growth stages [80]. Apart from morpho-physiological factors various genes and proteins are also responsible for drought tolerance in chickpea crop (Table 2).
Table 1. Relevance of various physiological traits contributing to drought adaptation in chickpea.
Table 2. List of some genes conferring adaptation to drought and other abiotic stresses in chickpea.
Due to the overabundance of wheat in irrigated areas in India, chickpea growth is primarily limited to rainfed areas. Crops in rainfed areas are experiencing water shortages, particularly during the sowing and terminal growth periods.
Soil and plant management are important for minimizing water stress. For this purpose, various experiments have been conducted with the applications of different agents. Gypsum can enhance overall plant growth, since it is a moderately soluble source of the crucial plant nutrients, calcium and sulphur. Gypsum supplements can also enhance the physical and chemical characteristics of soils, hence lowering nutrient concentrations in surface water runoff and reducing soil erosion losses. The most often used addition for reclaiming sodic soil is gypsum, which can also be found in synthetic soils used in nursery, greenhouse, and landscaping applications. Gypsum can be used for a variety of purposes in agriculture and horticulture, which could be advantageous to users. There are currently no recognized standards that outline the broad best management practices for using gypsum in agricultural applications.
Drought tolerance is a complex phenomenon that involves defence mechanisms as well as stress-induced signal responses [114][115]. Drought stress triggers a number of physiological, biochemical, and molecular responses (Figure 1) that can be classified into six categories: drought escape [116], avoidance [117], tolerance [118], resistance [119], abandonment [120], and drought adaptation [12][121]. Some chickpea genotypes have been identified as drought sensitive [122] and others as drought tolerant [123][124]. Plant breeders apply different ways of selection and development of drought tolerant crop genotypes. Different strategies are important to protect plants from harmful effects of drought.
Figure 1. Diagrammatic representation of effects of drought stress on chickpea.
Drought escape is the capacity of the plant to complete its life cycle before experiencing a major water deficit. Drought escape causes early flowering and maturity, as well as better yield potential, allowing plants to finish reproduction before drought strikes [125]. Crop longevity is governed in part by genotype and in part by the environment, and it impacts the crop’s ability to withstand climatic conditions such as drought. To achieve large seed yields, it is necessary to match the plant growth time to soil moisture availability. The genotypes with early maturity have the capacity to escape the terminal drought stress, whereas the genotypes with late maturity generally needs well-watered environments. The length of the growing phase and yield potential are positively associated with each other. In this consequence, the development of shorter duration crop is important for the reliable management of drought stress in the chickpea. The timing of flowering is a key feature of a plant’s response to extreme drought and high temperatures [126]. Early maturity lets the crop circumvent the passé of stress, hence short duration cultivars may be produced to minimise production loss from terminal dryness. However, under ideal growth conditions, the yield is often associated with crop duration, and any decrease in crop interval underneath the optimal will tax yield [29][52].
Drought escape is a critical strategy for preventing chickpea crop from drought [12]. Water supply is harmonised with phenological development in drought escape. Early maturity aids in escaping terminal dryness and is a key feature in germplasm screening. However, growers are frequently incapable to reorganize for early planting owing to climatic factors [52].
Drought avoidance is explained as a plant’s aptitude to retain a high tissue water potential contempt in a lack of soil moisture [125]. Processes involved in the enhancement of water intake, its storage in plant cells, and limiting water loss are associated with drought avoidance. Other mechanisms, including deep rooting, increased level of hydraulic conductance, reduced level of epidermal conductance, radiation absorption and reduced leaf area have also been reported to be linked with drought avoidance in plants. Deep rooting promotes water intake, which is helpful in reducing water losses. In the chickpea, the stomata remains closed during the day to minimise water loss during drought, and as a result, the carbon assimilation is impeded, lowering production [126][127].
Root biomass plays major role in absorbing water [128], as it is advantageous even in the condition of less moisture in the soil. It means there is a linkage between the root system and water stress tolerance [129][130], thus, in the current scenario, breeders are focused in the development of cultivars with larger root systems [131]. From integrating large root features, cultivars have been developed by chickpea breeders with increased drought tolerance [43][132]. Because root size is governed by intrinsic genetic variables [133][134] and modified by multiple environmental signals, such as nutrition and moisture accessibility in the soil, it is a complicated feature [135]. During the vegetative growth stage, susceptible genotypes absorb more water than tolerant genotypes, whereas tolerant genotypes absorb more water during the reproductive stage [136]. The intake of water during the vegetative as well as reproductive stages of plants has a direct relation with seed yield [137]. The importance of roots, rather than just root growth, is determined by their temporal water intake [138]. The best method for screening the germplasm for water usage competence (WUE) is carbon isotope discernment (13C), and this method has also been adopted in the chickpea [139][140].
One of the important impacts of drought stress is stomatal closure. Drought stress reduces the stomatal conductance and transpiration rate. This declines the CO2 fixation and photosynthesis due to the reduction in the internal CO2 concentration of the leaf (Ci). All of these factors have their role on the reduction in yield due to the reduced rate of photosynthesis [141].
The reduced rate of photosynthesis is directly related to extreme drought stress, and it is a result of the decreased chlorophyll content. Because of the lowered chlorophyll content, continuous poor moisture availability reduces light collecting capacity, triggering the generation of reactive oxygen species due to excessive energy absorption [142]. This is also a cause of damaged photosynthetic machinery. The principal cause of chlorophyll depletion is reactive oxygen species [143]. Reduction in photosynthetic activities under drought stress, have been experimented in chickpea genotypes [55] and this reduction was found to be linked to reduced ATP synthesis [144][145]. The yield reduction in chickpea genotypes due to the flower and pod drop under heat and drought stress circumstances was also noticed [146][147].
The leaf surface is also an imperative characteristic of plants in relation to drought stress. As tiny leaf surfaces lose less water [148], waxy leaves have high water preservation potential. Waxy leaves have the ability of a reflectance of irradiation and the reduction of water loss. This helps in the reduction of leaf temperature and provides tolerance against drought condition. The preservation of water in leaves with a reduced leaf temperature are directly related to the drought tolerant behaviour of plants. Drought stress raises leaf temperature in a variable manner, as tolerance genotypes have lower leaf temperatures than sensitive genotypes [149]. One drought tolerant chickpea variety ‘Gokce’ has been developed by ICARDA through the gene pyramiding method, which can be survived under severe drought conditions. This variety possess some other important features, such as early maturity, resistance to Ascochyta blight, increased seed size, and suitability for mechanised harvesting [150].
The drought tolerance refers to a plant’s ability to maintain its metabolism in a water shortage [35] condition with low tissue water potential [58]. Two types of traits are responsible for the drought tolerance in plants, i.e., constitutive characters and acquired behaviours. The constitutive traits affect the yield at mild to moderate levels of drought stress, whereas the acquired traits affect the yield at severe levels of drought stress. Drought tolerance features are largely concerned with cellular structural protection against the effects of cellular dehydration. Due to a reduction in the plant tissue water content, dehydrins and late embryogenesis of abundant (LEA) proteins accumulate [151]. These proteins act as chaperones [152].
In recent years, the role of reactive oxygen species (ROS) in stress signalling has been widely researched and evaluated [153][154]. The extreme creation of ROS causes oxidative damage and, lastly, cell death [155]. The role of ROS as a signalling molecule or in the oxidative damage depends upon the equilibrium between production and the scavenging of them [156]. The scavenging of ROS under drought stress depends upon the action of antioxidants in the cell [157][158].
Pushpavalli et al. [159] emphasised the need of selecting chickpea genotypes that can withstand various shocks rather than simply one. High temperature stress, in addition to drought stress, is a new threat to chickpea production [33][160]. According to Kalra et al. [161], a temperature increase of 18 °C above a particular threshold causes a significant loss in chickpea output. Furthermore, it is predicted that a global temperature increase of 2–38 °C, along with erratic rainfall patterns, would pose a threat to chickpea yield. In agriculture, the yield is the most important parameter for crops, and a reduction in yield cannot be compromised at any level. There is a strong association between drought tolerance with yield in a crop [122][162]. This is because yield-related traits of crops have been found to be sensitive under drought stress [163].

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