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Drought Stress in Plants a Review on Morphological Characteristics and Pigments Composition

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Drought Stress: Manifestation and Mechanisms of Alleviation in Plants

Submitted: December 13th, 2021 Reviewed: January 19th, 2022 Published: March 14th, 2022

DOI: 10.5772/intechopen.102780

From the Edited Book

Drought [Working Championship]

Associate Prof. Murat Eyvaz, Dr. Ahmed Albahnasawi, MSc. Mesut Tekbaş and Thousand.Sc. Ercan Gürbulak

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Abstruse

Drought can be referred to equally a meteorological period without pregnant rainfall and it is one of such major abiotic stresses that contributes to a huge reduction in crop yield throughout the world. Plant shows a wide range of physiological, morphological, and biochemical changes such every bit reduced photosynthetic accumulation, altered gene expression, etc. Under the drought stress which ultimately causes reduced growth every bit well equally poor grain yield. Drought stressconditions trigger production of ROS, which disrupts the dynamic balance between ROS production and ROS scavenging systems and its accumulation depends on the intensity besides as duration of water stress, and it varies among species. A constitute species that has a better inherited genetic response allowing information technology to rapidly deploy its antioxidant enzymatic and non-enzymatic defense system, can tolerate drought amend than a plant species with a poor antioxidant defense system. Furthermore, enzyme and protein encoding drought specific genes have the ability to enhance drought tolerance. These ii enzymatic and genetic engineering strategies are unique and vital tools, which tin be used to help alleviate the world's hereafter problems related to free energy, food, and environmental stresses, particularly drought. This chapter attempts to discuss developments in understanding effects of drought stress and underlying mechanisms in plants for its alleviation.

Keywords

  • ABA signaling
  • antioxidant
  • drought
  • ROS
  • stress

1. Introduction

Whatsoever inimical status or substance that affects found's metabolism, growth and development is referred equally stress. Basically, stress is an altered physiological status caused by different living and non-living factors which disturb the equilibrium. Plants are oft posed with a plethora of stress conditions such as drought, salinity, heat stress, depression temperature, heavy metal toxicity, flooding and extremes of soil pH. Plants also confront challenges from biotic factors similar pathogens, insects etc. These types of abiotic and biotic factors limit plants growth and productivity. The non-living variable must impact the environment beyond its normal range of variation to unfavorably impact the population performance or individual physiology of the organism in a meaning way.

Drought is a meteorological term and defined equally a period without significant rainfall. Generally, drought stress occurs when the available soil-h2o becomes scanty and atmospheric conditions cause continuous loss of water past transpiration or evaporation. Water deficit is i of the major abiotic stresses, which adversely affects ingather growth and yield. These changes are mainly associated with altered metabolic functions, one of those is either loss of or diminished synthesis of photosynthetic pigments, uptake and translocation of ion, saccharide biosynthesis, nutrient metabolism and synthesis of growth promoters. These changes in the metabolic functions and synthesis of photosynthetic pigments are closely related to biomass production in constitute [1]. A common agin effect of water stress on crop plants is the reduction in fresh and dry biomass [2]. Establish productivity under moisture stress is strongly associated with the processes of dry out affair partitioning and temporal biomass distribution [iii]. Previous study about different crop species faces huge yield reduction due to drought stress (Table 1). Nosotros have aimed to discuss the crops' response and adaptive mechanisms to gainsay drought stress and also genetic interventions which may help developing cultivars suitable for h2o-deficient weather.

Crop Yield reduction (%) References
Rice 53–92 [4]
Maize 79–81 [five]
Barley 49–57 [half-dozen]
Chickpea 45–69 [7]
Pigeonpea twoscore–55 [8]
Soybean 46–71 [9]
Sunflower lx [10]
Tater 13 [xi]
Canola xxx [12]
Cowpea 55–65 [thirteen]
Wheat 64.46 [xiv]

Tabular array 1.

Yield reduction attributable to drought stress in different crops.

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ii. Physiological changes during drought stress

During drought, Water scarcity occurs generally because of absence of water in the soil. Simply Physiological drought caused both lack of h2o in the soil, and as well occurs when excess water is present in the soil. Thus, physiological drought is a situation where the constitute cannot receive water [15, 16]. The responses of plants to h2o stress are diverse and may involve the contribution of various defense mechanisms or modification of physiology, morphology, anatomy, biochemistry, as well as short and long-term developmental and growth related accommodation processes [17].

Physiological reactions to moisture stress provides some escape mechanisms to the water stress comprise physiological and morphological adaptations [eighteen]. Decreased foliage area (Figure 1), reduced stomatal number and conductance, enlargement of root organization, increased leaf thickness, and leaf folding to lessen evapotranspiration are strictly associated with an adaptive response [17, 19, twenty, 21]. Institute growth and productivity decreased under moisture stress, which are caused by alterations in plantwater relations, CO2 assimilation reduction, membrane damage of affected tissues, cellular oxidative stress, and inhibition of enzymes activeness.

Effigy 1.

Effects of unlike levels of drought stress on ricebean seedlings.

Plants can change water relations to keep cellular mechanisms under drought stress conditions. Plants show osmotic adjustment past accumulating and integrating compatible solutes likely, proline, sugars and costless amino acids [22]. Maintenance of turgor pressure besides every bit cell volume at low water potential is facilitated by osmotic adjustment and is vital for metabolic functions. Osmotic aligning also plays role in recovery of metabolic activities post drought stress [23]. Previously, there are lot of studies investigated which showed the recovery of photosynthesis from moisture stress in various crop species and also recovered from drought stress in terms of oxidative stress, membrane stability alphabetize and antioxidative mechanisms [16, 24]. Osmolytes too accept a pregnant role in drought stress recovery.

Drought stress at college intensity decreases the activities of photosynthetic enzymes as well as foliage chlorophyll content which ultimately hampers the procedure of photosynthesis [20, 25]. Chlorophyll a / b proportion and synthesis of leaf chlorophyll altered during drought stress. A lower content of chlorophyll (Figure two), inactivation of central proteins linked to the photosynthesis procedure, and alteration of thylakoid membranes happen equally a upshot of drought stress. The decline in chlorophyll content is due to over production of O2 − and H2Oii production, which ultimately results meaning chlorophyll degradation and lipid peroxidation. During drought stress, in stomata and mesophyll cell the CO2 conductance declined equally the decrease in the photosynthetic process. The decrease in photosynthetic action also may be because of the reduction of stomatal movement [27]. Rubisco activity greatly afflicted by the loss of COtwo uptake, similarly information technology decreases the activity of sucrose phosphate synthase, nitrate reductase and RuBP production [20, 28]. Refuse in photosynthetic activity, loss of photosystem Ii photochemical efficiency, reduction in chlorophyll content and amending in stomatal movement results the reduction in found productivity. As a effect of reduction in photosynthetic activity in drought stress, it dismantles the product of sugar in diverse way likely prevents the transport of sucrose into sink organs and reduces the level of sucrose in leaves, which in turn limits reproductive development. The costless sugars and different metabolites thereof support plant growth nether drought, and have up osmolytic role and compatible solutes to mitigate the drastic effect of the stress [29, thirty].

Figure 2.

Visual effects of drought stress in rice. Source: [26].

The relative leaf h2o content (RLWC) is an guess of leafage's hydration status relative to its maximal water holding chapters at full turgid state. The relative leafage water content (RLWC) is one of the reliable parameters to know the water status in plants and it decreases gradually with increases in the severity of drought stress weather condition. The decline of RLWC every bit a response to osmotic stress was before reported by several investigators under different stress conditions [31, 32, 33, 34]. The physiological traits considered for evaluating drought stress tolerance include root trait characteristics (root length, root density, root biomass, root length density, delayed canopy wilting (DCW) and leaf pubescence density (LPD) [35], delayed leaf senescence (DLS) [36], and recovery ability after wilting (RAW) [37]. Drought stress drastically affects seed germination and decreases the speed of germination (Figure 3). Autonomously from these, stomatal conductance, chlorophyll content and use of carbon isotope discrimination are too constructive screening methods for drought stress tolerance and has been used for some food legumes.

Figure 3.

Ricebean response to varying levels of PEG every bit drought induction agent.

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3. Plants adaptive responses to drought stress

Plants accept adult various adaptive mechanisms conferring tolerance to drought stress induced adversities through evolution [38]. Their survival strategies for drought stress tin broadly exist classified as escape, abstention and tolerance. Hence, their drought stress response varies from molecular to plant level [39]. The mechanisms of found escape, avoidance and tolerance (Figure 4) confronting drought stress are discussed as follows.

Figure four.

Overall drought stress response in crop species.

3.1 Escape, avoidance and tolerance mechanisms

To escape the pernicious effects of drought stress on plant wellness and productivity, some plants utilize mechanisms involving shortening of the life wheel by rapid establish evolution, self-reproduction, and seasonal growth before the beginning of the drought season (Effigy 4) [40]. Amongst all, early flowering is perhaps the best possible escape adaptive mechanism in plants [41]. However, this machinery can connote a considerable reduction in the plant'due south growing period compromising plant productivity in some cases [42].

In abstention strategy, loftier plant water potential is maintained through transpiration loss reduction and the increased water uptake from well-established root systems [43]. Xeromorphic features such every bit the presence of hairy structure on leaves and cuticles in some cases practise help to maintain high water potentials in plant tissues [44]. It is notable that overdevelopment of these structures may lead to reduced productivity and reduced decreased size of vegetative and reproductive parts [45]. On the contrary, an adaptive tolerance mechanism at the photosynthetic level involves reductions in the institute's total leaf area and express expansion of new leaves. Likewise, trichrome production on leaves is an aspect that enables the plant to tolerate water deficits in dry environments [46]. There is an increase in charge per unit of lite reflection in the foliage reducing the leaf temperature too equally trichomes provide additional layer of resistance to the water loss thereby reducing the rate of h2o loss through transpiration [47]. Changes in root system-size, density, length, proliferation, expansion and growth rate, constitute the preliminary strategy for drought-tolerant plants to cope against drought [48]. Osmotic adjustment, antioxidant defense force mechanism, metabolic and biochemical dynamics of stomatal closure, solute accumulation and increase in root shoot ratio are other mutual strategies that aid to drought stress resilience [49].

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4. Biochemical responses to drought

4.1 Oxidative damage

Drought stress triggers an array of biochemical mechanisms including fluidity of the plasma membranes, osmolytes product, lipid peroxidation, reactive oxygen species (ROS) generation, rigidity of the cellular membranes and activation of unlike enzymes which are involved in oxidative defence force organisation [50, 51]. Previously, in various crop species ROS generation instigates pregnant damage to cellular components and also causing damages to lipid peroxidation, proteins [52]. The drought stress induced ROS generation had calamitous furnishings on lipid membrane and protein. Among all the ROS superoxide radical (O2•−), hydrogen peroxide (HtwoOii), singlet oxygen (iO2) and hydroxyl radical (OH•) are mainly produced by enzymatic or non-enzymatic processes during photosynthesis (Figure 5). Their production occurs also in components of electron transport system in the mitochondria by partial reduction or oxidation of atmospheric oxygen [53]. In some electric current studies, it has been shown that ROS have dual part in plant biology; involvement in vital signaling processes and as toxic by-products of aerobic metabolism [53].

Figure 5.

Production of various ROS by energy transfer (or) sequential univalent reduction of ground state triplet oxygen.

4.2 Enzymatic and non-enzymatic antioxidants

There are several componentsutilized by plantsto cope up with oxidative stress, which are involved in ROS homeostasis modulation [54]. Plants produces diverse reactive oxygen species (ROS) continuously equally bi-products of diverse metabolic pathways in unlike cellular compartments similar chloroplast, mitochondria, and peroxisome. ROS have partially reduced forms of atmospheric oxygen and nether normal conditions, their production in constitute cells is balanced by their constructive scavenging through enzymatic and non-enzymatic cascade (Figure 6). ROS can cause damage to different biomolecules namely Deoxyribonucleic acid, proteins and lipids, and therefore by creating oxidative injury; it leads to a reduction in found growth and development [56]. The equilibrium between the production and the scavenging of ROS may be perturbed past diverse stress factors. Thus, the disturbances of cellular homeostasis resulted in a sudden rise in intracellular levels of ROS leading to oxidative stress which in turn tin can cause substantial impairment to prison cell structure and membrane integrity. To mask themselves from these toxic oxygen intermediates, plant cells contain both enzymatic and not-enzymatic components. Amid them enzymatic antioxidantsare superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX), dehydroascorbate reductase (DHAR), monodehydroascorbate reductase (MDHAR) and ascorbate (AsA), glutathione (GSH), carotenoids, glycine betaine, proline, α-tocopherol and flavonoids are the non-enzymatic antioxidants [51, 57]. Hence, stress induced oxidative harm of ROSs can only be counteracted by increased level of enzymatic and nonenzymatic antioxidants [54].

Effigy six.

ROS scavenging machinery by antioxidant defense organisation in different stress weather condition. Source: [55].

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5. Molecular and genomic prospects for improvement of drought tolerance

Traditionally, at that place have been several efforts to develop drought-tolerant ingather genotypes through usual convenance methods [58, 59]. In this method, two groups of plants with desirable traits are selected and crossed to obtain offsprings having new genetic arrangements [threescore]. Drought resistance is directly or indirectly incorporated in the crop species via genetic variability of traits and thus selection in convenance is ought to be useful. Important traits to target in plant breeding might include water-extraction efficiency, water-employ efficiency, conductance of water, osmo-rubberband adjustments and leaf surface area modulation [15]. Genetic information improves the efficiency of the convenance method. Polymorphisms based on molecular markers that occur naturally in the DNA like restriction fragment length polymorphisms (RFLPs), sequence characteristic amplified regions (SCARs), random amplified polymorphic Dna (RAPDs), simple sequence repeats (SSRs), amplified fragment length polymorphism (AFLPs), and others have been effectively utilized. The use of plant convenance methods has an enormous potential to accelerate drought-tolerant found production and help drought management aid these plants [15].

Marker assisted option (MAS) and genomic selection (GS) are the 2 well versed approaches of genomic assisted breeding. For the first approach, foremost step is to place the molecular markers linked to the trait of interest so that selection tin can exist performed in breeding programs. However, GS depends on progress of selection models based on genetic markers present on the whole genome and selection of genome estimated breeding values (GEBVs) in breeding populations through phenotyping of "training population".

MAS utilizes molecular markers in identification of quantitative trait loci (QTL) or specific genes that are linked with the target trait and are used to place the individual with desirable alleles (Effigy vii) [61]. Through these methods, QTLs for the traits linked with drought resistance are identified in various crops i.eastward., rice, wheat, maize, sorghum, pearl millet, soybean and many other crops [62, 63, 64, 65, 66, 67].

Figure seven.

Model for the role of signaling factors in stomatal closure and retrograde signaling during water stress. Source: [sixteen].

Genomic selection utilizes all the markers bachelor for a population of GEBVs and GS models are used for selection of aristocracy lines without phenotyping [61]. Reverse to MAS, the information nigh QTLs is not the prerequisite for GS [68]. However, GS requires denser mark data than MAS. GS is being applied for breeding in maize tolerant to drought by the international maize and wheat improvement center (CIMMYT) [69]. Research efforts through this arroyo are progressing in other crops i.e., sugarcane, legumes and wheat [seventy, 71, 72].

Many studies take elucidated molecular responses in plants related to drought-induced transcription signaling pathways. In recent times, various stress-responsive genes and transcription factors having potential to mitigate drought-induced oxidative stress have been identified [73]. The TFs operate specific interaction with the cis-elements present in the genes' promoter region and, stimulate the expression of stress-inducible genes of various signaling pathways upon bounden [74, 75]. These TFs are categorized into unlike families based on their conserved motifs that lawmaking their DNA bounden domain (DBD), viz., APETALA two (AP2)/ethylene-responsive element binding cistron (ERF); dehydration-responsive chemical element bounden protein (DREB); no apical meristem/ Arabidopsis transcription activation factor, cup-shaped cotyledon (NAC); related to abscisic acid insensitive (ABI3)/VIVIPAROUS 1 (VP1) (RAV); WRKY; auxin response gene (ARF); and SQUAMOSA-promoter binding protein (SBP). The DBDs of the AP2/ERF, DREB, NAC, SBP, and WRKY are named as per the names of their respective TFs family, whereas DBDs of ABI3/VP1 and ARF family of TFs are named every bit B3 family [76].

Biochemical and molecular factors involved in the induction of processes to alleviate the detrimental impacts of h2o stress include transcription, stress responsive genes similar TaNAC69 (wheat), AP37 & OSNAC10 (rice), NF-YB2 (maize) and abscisic acid [16]. Transgenic expression of different stress responsive genes has been also utilized to confer increased tolerance to draught defecits. [77, 78]. The increased expression of these genes is oft associated with a decreased institute growth rate and this could narrow downwards its practical use (Table 2) [79]. In this sense, genomic and related molecular tools could accentuate the genes that mitigate the stress effect so that efforts may help maintaining those genes in breeding programs [104]. Marking assisted breeding combined with traditional breeding as an integrated approach is the best approach for the improvement of the drought stress tolerance in plants. [105, 106].

Establish species Genes Pathway involved/activated Role References
Oryza sativa OsNAC5/half-dozen/9/10 ABA responsive genes drought avoidance [80, 81, 82, 83]
NAC1/five/022 drought avoidance and activation of transcriptional regulation of various other genes [84]
bZIP23, ZFP252 Non identified drought avoidance and activation of transcriptional regulation of various other genes [85, 86]
OsMYB2 drought avoidance and activation of transcriptional regulation of various other genes [87]
DREB1F/DREB2A/EREBP1 mediates dehydration-inducible transcription Enhanced ROS scavenging induced drought tolerance [88]
Triticumaestivum TaNAC69, NAC2 ABA responsive gene drought avoidance and activation of transcriptional regulation of various other genes [89, 90]
TaPIMP1 Not identified drought avoidance and activation of transcriptional regulation of various other genes [91]
DREB2/DREB3 Overexpression causes strengthening of the antioxidant defense system in response to drought stress [92]
Arabidopsis thaliana NCED3 ABA responsive gene key enzyme of ABA biosynthesis [93]
AtABCG25 drought tolerance [94]
AHK1 positive regulator of osmosensing and drought tolerance [95]
OSCA1 membrane protein mediating osmotic stress responses [96]
Zat10 Not identified drought avoidance and activation of transcriptional regulation of diverse other genes [97]
AREB1/ AREB2/ ABF3/ ABF4 Induce drought tolerance by trifurcating feed forward pathway [98]
bZIP28 stress sensor and transducer in ER stress signaling pathway Activates brassinosteroid signaling and promotes acclimation to drought stress [99]
Zea mays bZIP17 noninducible expression of multiple genes involved in cell growth Induced drought tolerance by promoting jail cell differentiation [100, 101]
Solanumtuberosum StMYB1R-ane ABA responsive cistron drought avoidance and activation of transcriptional regulation of various other genes [102]
Gossypiumhirsutum SnRK2 ABA responsive gene imparts cellular adaptation in response to dehydration stress. [103]

Table ii.

Transcription factors involved in drought stress response in diverse crop species.

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six. Conclusion

Sustainable crop production to feed exponentially growing population is the major challenge to the scientific communities in the electric current global climate change scenario. Out of many productivity-limiting factors, drought stress is one of the most disquisitional factor and of prime importance in the context of decreasing water availability for crop production. Water deficit leads to cellular damage and triggers an array of signaling pathways which in turn activate synthesis of factor transcripts associated with protective functions. In general, wilting occurs owing to physiological responses such every bit reduced turgor pressure, gaseous exchange, mineral assimilation and overall growth. The prominent result of these is reduced photosynthetic rate Many plant species are inherently equipped with drought tolerance mechanisms such equally reduction in leaf expanse and canopy resistance. Both these mechanisms induce tolerance by cutting off excessive absorption of indecent lite as a event of reduced expanse exposed to the incident radiations. In order to select for a tolerant genotype and/or traits conferring tolerance, robust phenotyping is a must. Marker assisted breeding to comprise drought tolerance conferring quantitative trait loci (QTL) has proven to be constructive and efficient. In addition, the knowledge generated by "OMICS" techniques (genomics, proteomics, transcriptomics, epigenomics and metabolomics) and transgenomics are stiff and pregnant tools that would enable a researcher to develop an effective strategy for crop comeback programs in a less time-consuming cost-effective manner. So, an integrated approach will provide better agreement of mechanisms underlying drought stress and plants' response to information technology, and help in developing genotypes for dry out environments in order to reduce the threat to global nutrient security.

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Acknowledgments

The author would like to give thanks to the co-authors for their valuable inputs.

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Disharmonize of interest

The authors declare no conflict of interest.

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Kousik Atta, Aditya Pratap Singh, Saju Adhikary, Subhasis Mondal and Sujaya Dewanjee

Submitted: December 13th, 2021 Reviewed: January 19th, 2022 Published: March 14th, 2022

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