Open access peer-reviewed chapter

Interactive Effects of Salinity, Drought, and Heat Stresses on Physiological Process and Selection Criteria for Breeding Stress-Resistant Cotton

Written By

Volkan Mehmet Cinar, Serife Balci and Aydın Unay

Submitted: 24 January 2022 Reviewed: 26 May 2022 Published: 18 June 2022

DOI: 10.5772/intechopen.105576

From the Edited Volume

Advances in Plant Defense Mechanisms

Edited by Josphert Ngui Kimatu

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Abstract

The cotton crop is adversely affected by the combination of salinity, drought, and heat stress during all growth stages in cultivated areas. The negative impacts of salinity together with water scarcity on osmotic stress dramatically increased the sensitivity of reproductive development. After membrane integrity and signaling networks are depressed under stress at the cell level, the metabolic and physiological processes are disrupted in the next stage. The restricted root growth, ion and water uptake, phloem, photosynthetic and respiratory capacity, incompatible hormonal balance, and reduction in yield due to lower boll retention are the most important symptoms. The seed treatments and foliar applications of osmoprotectant and fertilization appear to reduce multiple stress factors in possible climate change conditions. The osmotic adjustment, antioxidative ability, electrolyte leakage in the membrane, and chlorophyll fluorescence are evaluated as selection criteria for improving genotypes. Direct selection of plants with high yield under stress conditions may increase the success of cotton breeding. It is important to know the molecular approaches and gene functions responsible for abiotic stress. In this chapter, the effects of high temperature, salinity, and drought on cotton plants and characteristics associated with tolerance were focused on cotton improvement. The classical breeding methods and molecular approaches should be combined for breeding new cotton varieties.

Keywords

  • breeding
  • high temperature
  • saline condition
  • stress physiology
  • water deficiency

1. Introduction

Agricultural areas with saline soils are estimated at 1100 Mha in the World. These areas are classified as saline (60%), sodic (26%), and saline-sodic soils (14%). Areas suffering from salinity are mostly in the Middle East, Australia, North Africa, and Eurasia. However, 20 to 50 percent of irrigated soils in arid or semi-arid climates are salt-affected [1]. A report produced by Cotton 2040 emphasized that cotton affected by heat stress and drought will reach 40% and 50%, respectively by 2040 [2]. Although heat stress is a problem along with drought [3], cotton-growing face many stress factors such as drought (Figure 1B), high temperature (Figure 2C), salinity (Figure 1A), alkalinity (Figure 1C), and heavy metal contamination, which are seen together in the same ecology. A single stressor can play a predominant or protective role depending on stress resources [4]. The long-term effect of climatic change resulted in higher pH values with drought, unfavorable soil organic C, total N and P, and available N [5], and soil salinity due to increasing the sea level [6].

Figure 1.

Cotton plants under saline soil (A), stunted cotton plants under drought conditions (B), and cotton plants under alkali soil (C).

Figure 2.

A picture of a plastic tunnel under heat stress at 11.00 am–17.00 pm (A), non-affected plants by heat stress (B), and aborted/shed bolls affected by heat stress (C).

Cotton’s response to stress is different at plant growth and development stages. Germination and seedling growth are adversely affected by high salinity and low temperature at the early stages in cotton (Gossypium hirsutum L.) [7, 8]. Although progenitors of cotton have spread in adverse conditions, modern cotton varieties are non-resistant to stresses from the squaring stage to boll retention stages, in which yield formation occurs [9]. Water deficiency [10, 11, 12, 13] and heat stress [13, 14, 15], especially during flowering and boll formation can reduce yield and fiber quality. In many areas of the world where cotton is cultivated, as in the Aegean, Mediterranean, and Southeast Anatolia regions of Turkey, maximum temperatures during the reproductive stage in July and August are above 40°C with low humidity and precipitation. The boll components such as seed and boll weight cannot be affected by short-term temperature changes, whereas the effects of sudden temperature rises during anthesis on seed number per boll are very high [16].

This review presents the physiological mechanism of stress tolerance and principles of genotype improvement, both classically and by genetic engineering in cotton. Also, it discussed the effects of agronomic management such as seed and foliar treatments to alleviate drought, salt and heat stress.

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2. Roles of osmolytes

The stress-induced protein breakdown and amino acid synthesis caused the accumulation of amino acids in cells under abiotic stress conditions. The overall accumulation of amino acids has two different symptoms: cell damage and the beneficial effect of specific amino acids such as proline during stress acclimation [17, 18]. Drought, salinity, and heavy metals caused an increase in the accumulation of proline in resistant plants. Although proline was not synthesized in tobacco exposed to heat stress [19], the combination of heat and drought stress [20], and drought stress [21] induced an increase in the proline content of some cotton cultivars. In contrast with this finding, genotypic differences were non-significant for proline content under drought conditions [22].

The first approach to stress resilience is to restrict ethylene synthesis, which triggers the abscission of leaves and all reproductive organs under stress conditions [23]. Boll retention and yield increased by application of ethylene inhibitors such as aminoethoxyvinylglycine (AVG) or by downregulation of genes responsible for ethylene synthesis [24]. Abscisic acid (ABA), another growth-inhibitory hormone, is an important regulator of abiotic stress tolerance. The main function of ABA is to stimulate stomatal closure and gene expression to respond to the drought, salinity, and excessive temperature in the adaptive mechanism of plants [25, 26]. Under stress conditions, plants can alter their metabolism such as the synthesis of compatible solutes. In cell metabolism, the different unfavorable conditions increased the concentration of γ-aminobutyric acid (GABA) through enhanced activity of enzymes involved in GABA biosynthesis [27]. Under alkali stress conditions, the accumulation of GABA and putrescine in young and old leaves of cotton increased [28]. Polyamines such as putrescine, spermidine, and spermine have a protective role against salinity, drought and heat stress, and putrescine and spermine could be evaluated as selection criteria for stress-tolerant genotype breeding in cotton [29]. Exogenously applied polyamines increased the stress tolerance and yield under drought and salinity stress conditions in cotton [30, 31].

The osmoprotectant solutes such as glycine betaine (GB) are evaluated in three different ways. Firstly, the increase in GB under stress conditions is analyzed as biochemical. Secondly, the selection of the plants with high GB levels is used for improving the stress-tolerant cotton genotypes in conventional breeding and genetic engineering. Thirdly, the success of seed treatment with GB and foliar application of GB were examined in stress conditions. The accumulation of GB increased in transgenic crops, whereas foliar application of GB or seed dressed are common applications in plants where a certain amount of synthesis is insufficient [32, 33]. Generally localized in the chloroplast, GB has an important role in protecting photosystem II, stabilizing membranes, and alleviating ROS (reactive oxygen species) damage [34] and chilling damage [35], enhancing tolerance to lead (Pb) [36]. Cottonseed coating with GB enhanced seed cotton yield by approximately 20% [37]. Also, combined foliar application with GB and salicylic acid (SA) increased the tolerance to salt stress due to an increase in the leaf gas exchange with positively correlated stomatal properties and stimulate antioxidant enzyme activity in cotton seedlings [38, 39].

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3. Heat stress

On the cell basis, heat stress firstly induced structural changes in chloroplast protein [40] and plasma membrane [41, 42], and these changes stimulated cell elongation, expansion, and differentiation. ROS, as a signaling molecule, induced excessive MDA (malondialdehyde) synthesis by increasing lipid peroxidation, and membrane structure disintegrated under heat stress conditions [43, 44, 45]. Also, molecular chaperones and heat shock proteins (HSP) accumulate to protect the membrane integrity by the expression of the sHSP coding gene in leaves under drought and heat stress conditions [46]. Heat stress caused the accumulation of specific isoforms of activase in cotton leaves [47, 48]. Although the efficiency of Rubisco activase decreased under heat stress, an enzyme can develop an alternative function by relocating on a thylakoid membrane [49], and this contributes to the acclimation of photosynthesis during high temperatures in cotton [50]. In addition, the synthesis of phytohormones such as SA and jasmonic acid (JA) to respond to heat stress is enhanced by ROS and Ca2+ [51]. The association between heat stress and plant nutrition demonstrated that the fertilization containing some macro and microelements such as K, Zn, and B could be used to alleviate the harmful effects of higher temperatures. These elements play an important role in chlorophyll synthesis and delay senescence in cotton [52, 53]. Zn spray eliminates the adverse effects of heat stress in cotton [54].

The reproductive period is synchronized with many processes in cotton. Although square retention is less affected by heat stress [55], sexual reproduction is defined as both sensitive to high temperature and thermotolerant during flowering – boll retention depending on the time, length, and severity of stress in cotton [56, 57, 58]. Heat stress during this period adversely affected the development of both sexual organs and caused flower abnormalities such as small flower, elongated stigmas (Figure 3A and B) [55], gametophyte, pollen germination, and pollen tube growth [59, 60]. Abiotic stress limited fertilization by preventing pollen tube development [61]. The tapetum, the innermost layer of somatic cells in anther lobes, is responsible for microsporogenesis and secretion of enzymes for the release of microspores from tetrads [62]. Programmed cell death (PCD) induced tapetal degeneration by heat stress, resulting in male sterility [63]. The regulation of tapetal PCD and anther dehiscence were controlled by the GhCKI gene (G. hirsutum casein kinase I) in the heat stress tolerance of cotton [64].

Figure 3.

Elongated stigma affected by heat stress (A), and normal flower structure (B).

The decrease in the photosynthetic capacity of cotton at a temperature above 32°C reduces the accumulation of sucrose [65]. The decrease in sucrose accumulation adversely affected the fiber quality by inhibiting cellulose synthesis in higher temperature conditions [66]. Hereby, sucrose transport from subtending leaf of boll to developing boll negatively affected, and boll number per plant and boll weight reduced [67]. Plant growth is mostly regulated by sucrose, which is a source of substrates’ energy production and biosynthesis by decomposing into hexoses. Furthermore, the plant responses are determined by ROS scavenging capacity and the signal pathway of sucrose [68]. In many crops, drought increased carbohydrate deprivation and ABA levels, whereas the ability of reproductive sinks to use sucrose and starch was reduced [69].

Auxin, an essential hormone, is very important for stress tolerance [70] and plays a role as a coordinator of plant growth and development [71]. The inhibition of auxin synthesis by overexpressing miR157 caused the sensitivity to heat stress and anther sterility in the reproductive period of cotton [72]. The auxin signaling pathway is controlled by the suppression of the sucrose synthase gene, and the antagonistic relationship between auxin and sucrose regulated plant growth and development [73, 74]. The favorable balance between auxin and sucrose is indispensable for the response of anther to heat stress [75].

Shedding is the formation of the abscission layer by natural or stress between sympodial or monopodial branches and reproductive organs such as square, flower, and boll in cotton (Figure 2B and C) [76]. The amount of hormones and regulation among hormone-controlled shedding, as auxin inhibits, and abscisic acid (ABA) promotes. Shedding is induced by increasing ABA and decreasing auxin under drought conditions [77]. In the reproductive period, failed fertilization caused by non-available pollen in higher temperatures resulted in flower shedding.

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4. Salinity stress

The physiological drought via osmotic stress and ion toxicity caused by Na+, Cl, and SO42− are two major forms of damage. Although cotton is classified as moderately tolerant to salt stress (7.7 dS m−1), salinity caused a decrease in seed cotton yield [78], boll number per plant [79, 80, 81], and an increase in early maturity [82]. Seed germination and early seedling growth in cotton are the most sensitive stages [83]. Both forms of salt damage caused abnormal plant growth such as stunted root and shoot growth by reducing photosynthetic capacity (Figure 1A) [84]. The excessive salt accumulation under salinity caused cellular injury in transpiring leaves [85]. However, cotton was successfully cultivated as a monoculture crop in saline-alkali soil. Plant cells are affected by low amounts of available water, loss of membrane functions, and ionic toxicity under excess sodium (Na+). As with most abiotic stress, salinity contains an osmotic component, and cellular dehydration causes and disrupts the internal balance (homeostasis). The plant’s first reaction is to reduce the Na+ level in the cytosol by restricting influx, increasing efflux, and accumulating Na+ in the vacuole for maintaining the cell metabolism. Transporters such as antiporter, uniporter, and symporter localized in the membrane are responsible for reducing Na+ [86, 87, 88]. In a study conducted by [89], the Na+/H+ antiporter gene (GhSOS1) was detected in the plasma membrane of cotton (G. hirsutum L.). This gene is a Salinity Overly Sensitive, which has an important role to synthesize protein in upregulating under stress conditions such as salinity and drought. The suberization and lignification may occur around endodermal cells to inhibit apoplastic absorption of toxic ions in cotton [90, 91].

Many researchers focused on a root-associated microorganism to alleviate salinity stress in cotton. The arbuscular mycorrhizal fungi (AMF) are capable of increasing P and Zn uptake and promoting leaf proline accumulation [92, 93] but it should be noted that Glomus mosseae, AMF species, isolated from saline soil found to be less successful in alleviating salt stress compared to that of non-saline soil [94]. The microorganisms with the PGPR effect such as Pseudomonas fluorescens have IAA producing ability involved in the synthesis of important compounds under salinity stress [95]. In addition, the higher rhizosphere colonization of PGPR induced moderate N application due to the signaling molecule role of nitric oxide (NO) in the denitrification process [96]. Also, melatonin, as an indole hormone, alleviated the adverse effects of salt stress to reduce ROS production and ion toxicity and increases proline content in cotton seedlings [97].

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5. Drought stress

The increase in the tensile force of the xylem, due to drought, caused a greater probability of rupture in the water column and formation of gas emboli in the xylem [98], and embolism decreased carbon assimilation depending on the linkage between water and CO2 exchange. Furthermore, stomatal closure and decrease in the stomatal area take place [99], and desiccation of all plants is induced in the further process (Figure 1B) [100]. Thus, the most negative impacts occurred in photosynthetic capacity and transpiration rate [101]. The environmentally induced PCD (programmed cell death) occurred in response to drought, and PCD increased ROS accumulation, DNA fragmentation, organelle degeneration, and cytoplasm shrinkage [102]. The tylose formation and xylem inhabitation by wilt pathogens in water-limiting conditions may cause drought sensitivity to stress the susceptibility [103]. Primarily, MAPK (Mitogen-activated protein kinase), and secondly, ROS play an important role in intracellular signaling [104, 105]. GhMKK3 and GhMPK2 from G. hirsutum increased root hair development and ROS production by regulating ethylene synthesis, respectively under drought conditions [106].

Arbuscular mycorrhizal fungi (AMF) have an important role to alleviate drought stress by spreading on the soil and water transport by hyphae [107]. Similarly, PGPR coated phosphorus exhibited high performance due to the increasing stomatal conductance, net photosynthetic capacity, and yield of cotton under osmotic stress [108]. As a result of studies investigating the relationship between nutrients and drought, it was suggested that high N concentrations may decrease the effects of drought through nitrogen metabolism, proline synthesis antioxidant capacity, and osmotic regulation in cotton [109, 110]. However, K application has osmotic regulation due to increasing the osmoprotectant and regulates N metabolism [53, 111, 112] and photo-assimilation and translocation process [113] in drought-stressed cotton plants. Also, supplemental Zn alleviated the negative effects of drought stress by increasing antioxidant capacity and decreasing MDA content in cotton [112, 114]. The nanoparticles such as nano-TiO2 and nano-SiO2 could alleviate drought stress because their foliar applications increase photosynthetic pigments, antioxidant capacity, and proline content in cotton [115].

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6. Heritability and breeding for stress tolerance in cotton

The combination of multiple stress factors and the identification of the model plant is complex at the physiological and genetic levels [116]. Growth chamber, greenhouse, and field studies were conducted by many researchers to screen the cotton genotypes in different stress studies. Stress tolerant genotypes can be bred through a selection of promising single plants under stress conditions followed by testing in either stress and non-stress conditions or selection under more optimum conditions for effective selection criteria [117]. Previous studies recommended the selection of cotton plants with higher seedling vigor, enhanced early root development, and lower root/shoot ratio for drought tolerance, pollen carrying a dominant heat-tolerance allele for heat stress, and salinity barrier for salt stress [118].

In stress-tolerant cotton breeding, genetic stock in strains with the D genome rather than those with the A genome should be considered. Transcriptomic analyses indicated that thick cuticles and a double layer palisade layer of D-genome species such as G. harknessii, G. armourianum, and G. turneri are important germplasm resources for water deficiency. Similarly, G. gossypioides and G. thurberi with aggressive and deep root structures for drought tolerance and G. aridum, G. davidsonii, and G. klotzschianum for both drought and salt tolerance can be used in cotton breeding [84, 119, 120, 121, 122]. In support of this knowledge, [123] revealed that QTLs (quantitative trait locus) responsible for salt tolerance are usually localized on the D-subgenome. At the same time, novel genes and alleles in wild relatives are important to overcome the abiotic stress tolerance caused by narrow gene pools in cultivated cotton species [124].

The selection of plants with thick cuticle and waxy surfaces is important to reduce solar radiation in the breeding of heat tolerance [125]. The intense absorbency of cotton cultivars increased sensitivity to heat stress [126]. The most important cultural management is to arrange the sowing time to avoid the higher temperature in the reproductive period. However, planting cotton before the recommended time faced the problem of low temperature during the early growing stage. Therefore, breeding of high-temperature tolerant cotton varieties has been suggested as the best method [127].

A pre-screening of cotton germplasm and evaluation of hybrids by constructing a polythene tunnel at the reproductive stage is a very common method in cotton (Figure 2A) [128]. However, some researchers preferred delayed planting set of cotton plants to test genotypes in heat tolerance breeding [129, 130, 131, 132]. In addition, some researchers have used tetrazolium chloride for identifying heat tolerance in both vegetative tissues and pollen viability in cotton [133, 134, 135]. Various studies have emphasized the success of wild cotton species, their stacking progenitor alleles, and Gossypium tomentosum (heat-resistant species) for stress tolerance [13, 136, 137]. The pollen characteristics, germination ability, and tube length have been screened to determine the tolerant and susceptible genotypes under higher temperatures in cotton [13, 138]. On the other hand, the increase in fiber wax content of susceptible cotton genotypes under heat and drought stress indicated that acceptable fiber wax levels could be used to improve tolerant genotypes in conventional breeding [139].

In quantitative genetic studies about heritability and gene action for different selection criteria of heat stress, multigenic inheritance and both additive and non-additive gene action in controlling cellular membrane thermostability was found to be higher under heat stress conditions [140]. The results of the scaling test, which is an important biometrical analysis method, indicated the significant dominance, additive x dominance, and dominance x dominance referred to as non-additive gene action for relative cell injury under heat stress [141]. Similarly, non-additive gene actions were estimated for fiber quality characters and ginning out-turn in heat stress (~38–39°C) at peak flowering time under field conditions by arranged sowing time [142]. In contrast, high heritability associated with high genetic advance for hydrogen peroxide content, catalase activity, total soluble proteins, carotenoids, and chlorophyll contents were found to have significant additive gene action under heat stress conditions [143]. In conventionally breeding of drought and heat stress tolerance, it was emphasized that instead of single plant selection in the F2 generation, superior plants should be selected in further generations such as F4 [144, 145].

PEG 6000 (polyethylene glycol) is defined as a rapid and effective method to observe the response of cotton genotypes for selection in drought tolerance breeding [146, 147]. Drought-tolerant species are defined with low maximum transpiration and photosynthetic rate, stomatal conductance, specific leaf area, small leaf size high leaf longevity, root mass ratio, and small leaf size [148]. The higher chlorophyll stability and relative water content exhibited drought tolerance due to photosynthate, which results in higher biomass [149] and fiber quality [150]. Furthermore, the presence of late embryogenesis abundant proteins is important for drought tolerance of cotton [151]. The physiological characteristics used for drought tolerance should be combined with yield, yield attributes, and fiber quality parameters in the breeding of cotton varieties with high adaptability [152]. Drought responsive genes were classified as induced (stress-related, metabolism, transcription factor, proline, and cellular transport) and repressed (mainly comprising metabolism, cellular transport, and stress-related) [153]. Drought tolerance genes such as RD2 (rice drought-responsive), HAT22 (homeobox from Arabidopsis thaliana), PIP2, PIP2C (plasma membrane intrinsic proteins), and GaTOP6B (encoding DNA topoisomerase from G. arboreum) were associated with drought in cotton [104]. Similarly, QTL analysis revealed that genes responsive to drought tolerance are spread over nine chromosomes while one QTL hotspot is concentrated on the eighth chromosome [154]. Anwar et al. [155] evaluated the selected varieties by molecular under drought stress, and MNH-886 cotton cultivar with high boll retention percentage, photosynthesis rate, and stomatal conductance was recommended against drought stress. The ratio of general combining ability and specific combining ability variance in line x tester analysis showed significant non-additive gene effects for proline content, total chlorophyll, canopy temperature, and cell membrane stability in drought stress conditions [156].

NaCl treatment at different doses is the most used method for screening cotton genotypes in order to determine suitable genotypes. Many researchers revealed significant variations in morphological, physiological, and biochemical characters under salinity stress [157, 158, 159]. The performance of genotypes, depending on genetic factors, compared with environmental factors and genotype x environment interactions, and high genetic gain by suitable selection increased the breeding success because of high additive effects under saline [160] and drought conditions [161]. On the other hand, non-additive gene actions were found significantly higher for chlorophyll content, K+/Na+ ratio and within boll yield components under saline conditions [144, 162]. At the same time, selection for K+, Na+, and K+/Na+ were recommended for salinity tolerance according to the results of factor analysis in cotton [163]. In addition, a reciprocal effect found in some studies indicated cytoplasmic and cytoplasmic x nuclear genes interaction for salinity tolerance breeding [160, 164]. Besides, the fact that genes with pleiotropic effects are effective for both salinity and drought indicated physiological characters are used in indirect selection can increase the success of multi-stress breeding [165, 166].

In many studies about engineering abiotic stress-tolerant crops, genes responsible for multiple stress factors found to confer tolerance in different plants are defined [167]. Co-overexpressing of SUMO E3 ligase (OsSIZ) and Vacuolar H+− pyrophosphatase (AVP1) from Arabidopsis and vacuolar Na+/H+ antiporter genes performed significantly in increasing tolerance to multiple stressors such as drought, salinity, and higher temperature [168]. Overexpression of mitogen-activated protein kinases (MAPK) from cotton GhMPK2 performed stress tolerance when induced by salt, ABA, and water scarcity [169]. Eight genes belonging to the GhHSP20 family are responsible for heat, drought, and salinity [170], and the expression of GHSP26 caused a significant increase in proteins under drought stress [171]. Similarly, it was verified that GhCIPK6a overexpressed cotton lines can reduce the negative effects of salinity, and the seeds of these lines exhibited higher water absorption capacity at the germination stage [172]. In another study, JAZ proteins were identified as inhibitors of JA, and overexpression of JAZ genes resulted in a higher performance to salt stress [173]. GaJAZ1 transgenic plants carrying genes transferred from G. arboreum to G. hirsutum were found to be different from wilt type in terms of salinity tolerance [174]. GhPHD genes (plant homeodomain genes from G. hirsutum) increased plant tolerance in adverse environmental conditions to alleviate abiotic and phytohormonal stresses [175]. Also, the overexpression of the 14–3-3 gene GF14λ from Arabidopsis resulted in a stay-green phenotype due to decreased wilting, delayed senescence, and higher photosynthetic capacity. It was suggested that this gene can be used successfully in drought resistance [176, 177].

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7. Inferences (future considerations)

The nutrient relations of cotton plants were greatly affected by abiotic stress. The nitrogen, silicon, magnesium, and calcium uptake are limited under high temperatures, drought, and salinity. In addition, the architecture of cotton changed to capture the less mobile nutrients. Therefore, fertilization should be reconsidered under stress conditions. To reduce the negative effects of abiotic stress, the use of some osmoprotectant and hormones may be a solution. The sowing time and plant density should be rearranged especially for high temperatures and drought. Cluster-type cotton varieties suitable for High-Density Planting System (HDPS) should be developed. These varieties will also reduce the density of weeds and dry matter content that compete for available water under drought conditions.

The most effective way to overcome the adverse effects of multiple abiotic stress factors is to improve tolerant cultivars. The possible effects of drought, salinity, and high-temperature stress on physiological, morphological, and yield should be well resolved to mitigate stress and develop varieties. The crossing between standard cultivars and donor suitable parents is a basic stage for genetic variation in conventional breeding. Multiple stress factors are very complex and polygenic characters and are controlled by non-additive gene effects with low heritability. Therefore, bulk selection should be useful for traits with low heritability such as stress tolerance, and the selection of a single plant should be postponed to F4 or F5 generation. Drought and high temperature are effective stress factors from flowering and during the first boll formation period, whereas salinity is an important problem in the early development period. In areas where these conditions are created, indirect selection can be made in terms of physiological characteristics, whereas direct selection can be made for yield. The evaluation of genotypes to stress tolerance under the plastic tunnel at the reproductive stage and in saline and drought conditions is a more accurate approach compared to artificial conditions and delayed sowing date in cotton.

Genetic engineering studies are promising, but manipulation of a single gene does not seem to be sufficient as resistance to multiple stress conditions is controlled by multiple genes. However, future transgenic cultivar breeding may be a good approach for multiple stress tolerance. In both classical breeding and genetic engineering studies, tolerance to multiple stress factors should be pyramided with yield and fiber characteristics. The non-genetically CRISPR-Cas system could be evaluated for engineering multiple stress tolerance in future cotton cultivars for all cotton-growing regions.

It could be highlighted that temperature extremes and fluctuations in the cotton-growing season along with other stress appear to be major factors of future yield reductions.

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Acknowledgments

Volkan Mehmet CINAR thanks the Higher Education Council of Turkey (YOK) for 100/2000 PhD scholarship.

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Conflict of interest

The authors declare no conflict of interest.

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Author statement

Volkan Mehmet CINAR: Conceptualization, Literature survey, Writing – Review & Editing. Serife BALCI: Writing – Review & Editing. Aydın UNAY: Conceptualization, Literature survey, Writing – Review, Supervision.

References

  1. 1. FAO. Global soil partnership. Soil salinization as a global major challenge. [Internet]. 2021. Available from: https://www.fao.org/global-soil-partnership/resources/highlights/detail/en/c/1412475/ [Accessed: April 11, 2021]
  2. 2. Anonymous. Physical climate risk for global cotton production: Global analysis. [Internet]. 2021. Available from: https://www.acclimatise.uk.com/wp-content/uploads/2021/06/Cotton2040-GAReport-FullReport -highres.pdf [Accessed: October 11, 2021]
  3. 3. Rahman HU. Number and weight of cotton lint fibres: Variation due to high temperatures in the field. Australian Journal of Agricultural Research. 2006;57(5):583-590. DOI: 10.1071/AR05135
  4. 4. Zhou R, Yu X, Ottosen CO, Rosenqvist E, Zhao L, Wang Y, et al. Drought stress had a predominant effect over heat stress on three tomato cultivars subjected to combined stress. BMC Plant Biology. 2017;17(1):1-3. DOI: 10.1186/s12870-017-0974-x
  5. 5. Jiao F, Shi XR, Han FP, Yuan ZY. Increasing aridity, temperature and soil pH induce soil CNP imbalance in grasslands. Scientific Reports. 2016;6(1):1-9. DOI: 10.1038/srep19601
  6. 6. Rahman AK, Ahmed KM, Butler AP, Hoque MA. Influence of surface geology and micro-scale land use on the shallow subsurface salinity in deltaic coastal areas: A case from southwest Bangladesh. Environmental Earth Sciences. 2018;77(12):1-8. DOI: 10.1007/s12665-018-7594-0
  7. 7. Gorham J, Läuchli A, Leidi EO. Plant responses to salinity. In: Stewart JM, Oosterhuis DM, Heitholt JJ, Mauney JR, editors. Physiology of Cotton. Dordrecht: Springer; 2010. pp. 129-141. DOI: 10.1007/978-90-481-3195-2_13
  8. 8. Rehman A, Kamran M, Afzal I. Production and processing of quality cotton seed. In: Ahmad S, Hasanuzzaman M, editors. Cotton Production and Uses. Singapore: Springer; 2020. pp. 547-570. DOI: 10.1007/978-981-15-1472-2_27
  9. 9. Oosterhuis DM, Loka D. Polyamines and cotton flowering. In: Oosterhuis DM, Cothren JT, editors. Flowering and Fruiting of Cotton. Memphis: The Cotton Foundation; 2012. pp. 109-132
  10. 10. Basal H, Dagdelen N, Unay A, Yilmaz E. Effects of deficit drip irrigation ratios on cotton (Gossypium hirsutum L.) yield and fibre quality. Journal of Agronomy and Crop Science. 2009;195(1):19-29. DOI: 10.1111/j.1439-037X.2008.00340.x
  11. 11. Loka DA, Oosterhuis DM. Water stress and reproductive development in cotton. In: Oosterhuis DM, Cothren JT, editors. Flowering and Fruiting in Cotton. Candova: The Cotton Foundaiton; 2012. pp. 51-58
  12. 12. Lokhande S, Reddy KR. Reproductive and fiber quality responses of upland cotton to moisture deficiency. Agronomy Journal. 2014;106(3):1060-1069. DOI: 10.2134/agronj13.0537
  13. 13. Reddy KR, Bheemanahalli R, Saha S, Singh K, Lokhande SB, Gajanayake B, et al. High-temperature and drought-resilience traits among interspecific chromosome substitution lines for genetic improvement of upland cotton. Plants. 2020;9(12):1747. DOI: 10.3390/plants9121747
  14. 14. Reddy KR, Davidonis GH, Johnson AS, Vinyard BT. Temperature regime and carbon dioxide enrichment alter cotton boll development and fiber properties. Agronomy Journal. 1999;91(5):851-858. DOI: 10.2134/agronj1999.915851x
  15. 15. Zhao D, Reddy KR, Kakani VG, Koti S, Gao W. Physiological causes of cotton fruit abscission under conditions of high temperature and enhanced ultraviolet-B radiation. Physiologia Plantarum. 2005;124(2):189-199. DOI: 10.1111/j.1399-3054.2005.00491.x
  16. 16. Reddy KR, Brand D, Wijewardana C, Gao W. Temperature effects on cotton seedling emergence, growth, and development. Agronomy Journal. 2017;109(4):1379-1387. DOI: 10.2134/agronj2016.07.0439
  17. 17. Patterson JH, Newbigin ED, Tester M, Bacic A, Roessner U. Metabolic responses to salt stress of barley (Hordeum vulgare L.) cultivars, Sahara and Clipper, which differ in salinity tolerance. Journal of Experimental Botany. 2009;60(14):4089-4103. DOI: 10.1093/jxb/erp243
  18. 18. Krasensky J, Jonak C. Drought, salt, and temperature stress-induced metabolic rearrangements and regulatory networks. Journal of Experimental Botany. 2012;63(4):1593-1608. DOI: 10.1093/jxb/err460
  19. 19. Dobra J, Motyka V, Dobrev P, Malbeck J, Prasil IT, Haisel D, et al. Comparison of hormonal responses to heat, drought and combined stress in tobacco plants with elevated proline content. Journal of Plant Physiology. 2010;167(16):1360-1370. DOI: 10.1016/j.jplph.2010.05.013
  20. 20. De Ronde JA, Van Der Mescht A, Steyn HS. Proline accumulation in response to drought and heat stress in cotton. African Crop Science Journal. 2000;8(1):85-92
  21. 21. Ahmed AH, Darwish E, Alobaidy MG. Impact of putrescine and 24-epibrassinolide on growth, yield and chemical constituents of cotton (Gossypium barbadense L.) plant grown under drought stress conditions. Asian Journal of Plant Sciences. 2017;16(1):9-23. DOI: 10.3923/ajps.2017.9.23
  22. 22. Zahid Z, Khan MK, Hameed A, Akhtar M, Ditta A, Hassan HM, et al. Dissection of drought tolerance in upland cotton through morpho-physiological and biochemical traits at seedling stage. Frontiers in Plant Science. 2021;12:627107. DOI: 10.3389/fpls.2021.627107
  23. 23. Rahman MU, Majeed A, Zulfiqar S, Ishfaq S, Mohsan M, Ahmad N. In: Rahman M, Zafar Y, Zhang T, editors. Cotton Precision Breeding. Cham: Springer; 2021. pp. 137-156. DOI: 10.1007/978-3-030-64504-5_6
  24. 24. Najeeb U, Bange MP, Tan DK, Atwell BJ. Consequences of waterlogging in cotton and opportunities for mitigation of yield losses. AoB Plants. 2015;1:1-17. DOI: 10.1093/aobpla/plv080
  25. 25. Wani SH, Kumar V. Plant stress tolerance: Engineering ABA: A potent phytohormone. Transcriptomics. 2015;3(2):1000113. DOI: 10.4172/2329-8936.1000113
  26. 26. Fujita Y, Fujita M, Shinozaki K, Yamaguchi-Shinozaki K. ABA-mediated transcriptional regulation in response to osmotic stress in plants. Journal of Plant Research. 2011;124(4):509-525. DOI: 10.1007/s10265-011-0412-3
  27. 27. Renault H, Roussel V, El Amrani A, Arzel M, Renault D, Bouchereau A, et al. The Arabidopsis pop2-1 mutant reveals the involvement of GABA transaminase in salt stress tolerance. BMC Plant Biology. 2010;10(1):1-6. DOI: 10.1186/1471-2229-10-20
  28. 28. Guo R, Shi L, Yang C, Yan C, Zhong X, Liu Q , et al. Comparison of ionomic and metabolites response under alkali stress in old and young leaves of cotton (Gossypium hirsutum L.) seedlings. Frontiers in Plant Science. 2016;25(7):1785. DOI: 10.3389/fpls.2016.01785
  29. 29. Loka DA, Oosterhuis DM, Pilon C. Effect of water-deficit stress on polyamine metabolism of cotton flower and their subtending leaf. In: Oosterhuis D, editor. Summaries of Arkansas Cotton Research. Arkansas: Arkansas Agricultural Experiment Station; 2011. pp. 70-75
  30. 30. Shallan MA, Hassan HM, Namich AA, Ibrahim AA. Effect of sodium nitroprusside, putrescine and glycine betaine on alleviation of drought stress in cotton plant. American-Eurasian Journal of Agricultural and Environmental Science. 2012;12(9):1252-1265. DOI: 10.5829/idosi.aejaes.2012.12.09.1902
  31. 31. Ahmed AH, Darwish ES, Hamoda SA, Alobaidy MG. Effect of putrescine and humic acid on growth, yield and chemical composition of cotton plants grown under saline soil conditions. American-Eurasian Journal of Agricultural and Environmental Science. 2013;13(4):479-497. DOI: 10.5829/idosi.aejaes.2013.13.04.1965
  32. 32. Sakamoto A, Murata N. The role of glycine betaine in the protection of plants from stress: Clues from transgenic plants. Plant, Cell & Environment. 2002;25(2):163-171. DOI: 10.1046/j.0016-8025.2001.00790.x
  33. 33. Naidu BP, Williams RL. Seed Treatment and Foliar Application of Osmoprotectants to Increase Crop Establishment and Cold Tolerance at Flowering in Rice: A Report for the Rural Industries Research and Development Corporation. Brisbane, Australia: CSIRO Tropical Agriculture; 2004
  34. 34. Chen TH, Murata N. Glycine betaine protects plants against abiotic stress: Mechanisms and biotechnological applications. Plant, Cell & Environment. 2011;34(1):1-20. DOI: 10.1111/j.1365-3040.2010.02232.x
  35. 35. Cheng C, Pei LM, Yin TT, Zhang KW. Seed treatment with glycine betaine enhances tolerance of cotton to chilling stress. The Journal of Agricultural Science. 2018;156(3):323-332. DOI: 10.1017/S0021859618000278
  36. 36. Bharwana SA, Ali S, Farooq MA, Iqbal N, Hameed A, Abbas F, et al. Glycine betaine-induced lead toxicity tolerance related to elevated photosynthesis, antioxidant enzymes suppressed lead uptake and oxidative stress in cotton. Turkish Journal of Botany. 2014;38(2):281-292. DOI: 10.3906/bot-1304-65
  37. 37. Naidu BP, Cameron DF, Konduri SV. Improving drought tolerance of cotton by glycinebetaine application and selection. In: Michalk DL, Pratley JE, editors. Agronomy, Growing a Greener Future? Proceedings of the 9th Australian Agronomy Conference; Charles Sturt University, Wagga Wagga, Australia. Serpentine, Australia: The Australian Society of Agronomy Inc., Gosford, NSW, Australia: Published online by The Regional Institute Ltd.; 20-23 July 1998;1-5. Available from: http://www.regional.org.au/au/asa/1998/4/221naidu.htm
  38. 38. Hamani AK, Wang G, Soothar MK, Shen X, Gao Y, Qiu R, et al. Responses of leaf gas exchange attributes, photosynthetic pigments and antioxidant enzymes in NaCl-stressed cotton (Gossypium hirsutum L.) seedlings to exogenous glycine betaine and salicylic acid. BMC Plant Biology. 2020;20(1):1-4. DOI: 10.1186/s12870-020-02624-9
  39. 39. Hamani AK, Li S, Chen J, Amin AS, Wang G, Xiaojun S, et al. Linking exogenous foliar application of glycine betaine and stomatal characteristics with salinity stress tolerance in cotton (Gossypium hirsutum L.) seedlings. BMC Plant Biology. 2021;21(1):1-2. DOI: 10.1186/s12870-021-02892-z
  40. 40. Ahmad N, Michoux F, Nixon PJ. Investigating the production of foreign membrane proteins in tobacco chloroplasts: Expression of an algal plastid terminal oxidase. PLoS One. 2012;7(7):e41722. DOI: 10.1371/journal.pone.0041722
  41. 41. Potters G, Pasternak TP, Guisez Y, Jansen MA. Different stresses, similar morphogenic responses: Integrating a plethora of pathways. Plant, Cell & Environment. 2009;32(2):158-169. DOI: 10.1111/j.1365-3040.2008.01908.x
  42. 42. Rasheed R. Salinity and Extreme Temperature Effects on Sprouting Buds of Sugarcane (Saccharum officinarum L.): Some Histological and Biochemical Studies. Faisalabad: University of Agriculture; 2006
  43. 43. Majeed S, Rana IA, Mubarik MS, Atif RM, Yang SH, Chung G, et al. Heat stress in cotton: A review on predicted and unpredicted growth-yield anomalies and mitigating breeding strategies. Agronomy. 2021;11(9):1825. DOI: 10.3390/agronomy11091825
  44. 44. Burke JJ, Hatfield JL, Klein RR, Mullet JE. Accumulation of heat shock proteins in field-grown cotton. Plant Physiology. 1985;78(2):394-398. DOI: 10.1104/pp.78.2.394
  45. 45. Sable A, Rai KM, Choudhary A, Yadav VK, Agarwal SK, Sawant SV. Inhibition of heat shock proteins HSP90 and HSP70 induce oxidative stress, suppressing cotton fiber development. Scientific Reports. 2018;8(1):1-7. DOI: 10.1038/s41598-018-21866-0
  46. 46. Sarwar M, Saleem MF, Ullah N, Rizwan M, Ali S, Shahid MR, et al. Exogenously applied growth regulators protect the cotton crop from heat-induced injury by modulating plant defense mechanism. Scientific Reports. 2018;8(1):1-5. DOI: 10.1038/s41598-018-35420-5
  47. 47. Law RD, Crafts-Brandner SJ. High temperature stress increases the expression of wheat leaf ribulose-1, 5-bisphosphate carboxylase/oxygenase activase protein. Archives of Biochemistry and Biophysics. 2001;386(2):261-267. DOI: 10.1006/abbi.2000.2225
  48. 48. Law DR, Crafts-Brandner SJ, Salvucci ME. Heat stress induces the synthesis of a new form of ribulose-1, 5-bisphosphate carboxylase/oxygenase activase in cotton leaves. Planta. 2001;214(1):117-125. DOI: 10.1007/s004250100592
  49. 49. Rokka A, Zhang L, Aro EM. Rubisco activase: An enzyme with a temperature-dependent dual function? The Plant Journal. 2001;25(4):463-471. DOI: 10.1046/j.1365-313x.2001.00981.x
  50. 50. DeRidder BP, Salvucci ME. Modulation of Rubisco activase gene expression during heat stress in cotton (Gossypium hirsutum L.) involves post-transcriptional mechanisms. Plant Science. 2007;172(2):246-254. DOI: 10.1016/j.plantsci.2006.08.014
  51. 51. Rai KK, Pandey N, Rai SP. Salicylic acid and nitric oxide signaling in plant heat stress. Physiologia Plantarum. 2020;168(2):241-255. DOI: 10.1111/ppl.12958
  52. 52. Corrales I, Poschenrieder C, Barceló J. Boron-induced amelioration of aluminium toxicity in a monocot and a dicot species. Journal of Plant Physiology. 2008;165(5):504-513. DOI: 10.1016/j.jplph.2007.03.014
  53. 53. Zahoor R, Zhao W, Dong H, Snider JL, Abid M, Iqbal B, et al. Potassium improves photosynthetic tolerance to and recovery from episodic drought stress in functional leaves of cotton (Gossypium hirsutum L.). Plant Physiology and Biochemistry. 2017a;119:21-32. DOI: 10.1016/j.plaphy.2017.08.011
  54. 54. Sarwar M, Saleem MF, Ullah N, Ali S, Rizwan M, Shahid MR, et al. Role of mineral nutrition in alleviation of heat stress in cotton plants grown in glasshouse and field conditions. Scientific Reports. 2019;9(1):1-7. DOI: 10.1038/s41598-019-49404-6
  55. 55. Brown PW. Cotton Heat Stress [Internet]. 1999. Available from: https://cals.arizona.edu/AZMET/az1448.pdf [Accessed: November 13, 2021]
  56. 56. Hedhly A, Hormaza JI, Herrero M. Global warming and sexual plant reproduction. Trends in Plant Science. 2009;14(1):30-36. DOI: 10.1016/j.tplants.2008.11.001
  57. 57. Zinn KE, Tunc-Ozdemir M, Harper JF. Temperature stress and plant sexual reproduction: Uncovering the weakest links. Journal of Experimental Botany. 2010;61(7):1959-1968. DOI: 10.1093/jxb/erq053
  58. 58. Reddy KR, Hodges HF, Reddy VR. Temperature effects on cotton fruit retention. Agronomy journal. 1992;84(1):26-30. DOI: 10.2134/agronj1992.00021962008400010006x
  59. 59. Snider JL, Oosterhuis DM, Kawakami EM. Diurnal pollen tube growth rate is slowed by high temperature in field-grown Gossypium hirsutum pistils. Journal of Plant Physiology. 2011;168(5):441-448. DOI: 10.1016/j.jplph.2010.08.003
  60. 60. Echer FR, Oosterhuis DM, Loka DA, Rosolem CA. High night temperatures during the floral bud stage increase the abscission of reproductive structures in cotton. Journal of Agronomy and Crop Science. 2014;200(3):191-198. DOI: 10.1111/jac.12056
  61. 61. Oosterhuis DM, Snider JL. High temperature stress on floral development and yield of cotton. In: Oosterhuis DM, editor. Stress Physiology in Cotton. Cordova: The Cotton Foundation; 2011. pp. 1-24
  62. 62. Sanders PM, Bui AQ , Weterings K, McIntire KN, Hsu YC, Lee PY, et al. Anther developmental defects in Arabidopsis thaliana male-sterile mutants. Sexual Plant Reproduction. 1999;11(6):297-322. DOI: 10.1007/s004970050158
  63. 63. Li X, Gao X, Wei Y, Deng L, Ouyang Y, Chen G, et al. Rice APOPTOSIS INHIBITOR5 coupled with two DEAD-box adenosine 5′-triphosphate-dependent RNA helicases regulates tapetum degeneration. The Plant Cell. 2011;23(4):1416-1434. DOI: 10.1105/tpc.110.082636
  64. 64. Min L, Zhu L, Tu L, Deng F, Yuan D, Zhang X. Cotton Gh CKI disrupts normal male reproduction by delaying tapetum programmed cell death via inactivating starch synthase. The Plant Journal. 2013;75(5):823-835. DOI: 10.1111/tpj.12245
  65. 65. Crafts-Brandner SJ, Salvucci ME. Rubisco activase constrains the photosynthetic potential of leaves at high temperature and CO2. Proceedings of the National Academy of Sciences. 2000;97(24):13430-13435. DOI: 10.1073/pnas.230451497
  66. 66. Tian JS, Hu YY, Gan XX, Zhang YL, Hu XB, Ling GO, et al. Effects of increased night temperature on cellulose synthesis and the activity of sucrose metabolism enzymes in cotton fiber. Journal of Integrative Agriculture. 2013;12(6):979-988. DOI: 10.1016/S2095-3119(13)60475-X
  67. 67. Peng J, Liu J, Zhang L, Luo J, Dong H, Ma Y, et al. Effects of soil salinity on sucrose metabolism in cotton leaves. PLoS One. 2016;11(5):e0156241. DOI: 10.1371/journal.pone.0156241
  68. 68. Ruan YL, Jin Y, Yang YJ, Li GJ, Boyer JS. Sugar input, metabolism, and signaling mediated by invertase: Roles in development, yield potential, and response to drought and heat. Molecular Plant. 2010;3(6):942-955. DOI: 10.1093/mp/ssq044
  69. 69. Andersen MN, Asch F, Wu Y, Jensen CR, Næsted H, Mogensen VO, et al. Soluble invertase expression is an early target of drought stress during the critical, abortion-sensitive phase of young ovary development in maize. Plant Physiology. 2002;130(2):591-604. DOI: 10.1104/pp.005637
  70. 70. Xiao YH, Li DM, Yin MH, Li XB, Zhang M, Wang YJ, et al. Gibberellin 20-oxidase promotes initiation and elongation of cotton fibers by regulating gibberellin synthesis. Journal of Plant Physiology. 2010;167(10):829-837. DOI: 10.1016/j.jplph.2010.01.003
  71. 71. Leyser O. Auxin signaling. Plant Physiology. 2018;176(1):465-479. DOI: 10.1104/pp.17.00765
  72. 72. Ding Y, Ma Y, Liu N, Xu J, Hu Q , Li Y, et al. micro RNA s involved in auxin signalling modulate male sterility under high-temperature stress in cotton (Gossypium hirsutum). The Plant Journal. 2017;91(6):977-994. DOI: 10.1111/tpj.13620
  73. 73. Stokes ME, Chattopadhyay A, Wilkins O, Nambara E, Campbell MM. Interplay between sucrose and folate modulates auxin signaling in Arabidopsis. Plant Physiology. 2013;162(3):1552-1565. DOI: 10.1104/pp.113.215095
  74. 74. Yuan Y, Mei L, Wu M, Wei W, Shan W, Gong Z, et al. SlARF10, an auxin response factor, is involved in chlorophyll and sugar accumulation during tomato fruit development. Journal of Experimental Botany. 2018;69(22):5507-5518. DOI: 10.1093/jxb/ery328
  75. 75. Min L, Li Y, Hu Q , Zhu L, Gao W, Wu Y, et al. Sugar and auxin signaling pathways respond to high-temperature stress during anther development as revealed by transcript profiling analysis in cotton. Plant Physiology. 2014;164(3):1293-1308. DOI: 10.1104/pp.113.232314
  76. 76. Tariq M, Yasmeen A, Ahmad S, Hussain N, Afzal MN, Hasanuzzaman M. Shedding of fruiting structures in cotton: Factors, compensation and prevention. Tropical and Subtropical Agroecosystems. 2017;20(2):251-262
  77. 77. Guinn G, Brummett DL. Changes in abscisic acid and indoleacetic acid before and after anthesis relative to changes in abscission rates of cotton fruiting forms. Plant Physiology. 1988;87(3):629-631. DOI: 10.1104/pp.87.3.629
  78. 78. Zhang L, Zhang G, Wang Y, Zhou Z, Meng Y, Chen B. Effect of soil salinity on physiological characteristics of functional leaves of cotton plants. Journal of Plant Research. 2013;126(2):293-304. DOI: 10.1007/s10265-012-0533-3
  79. 79. Naderiarefi A, Ahmadi A, Tavakoli AR, Vafaietabar MAR, Sabikdast M. The effect of issue stress on physiological characteristics of leaf and drought resistance of different cotton genotypes. To Agricultural Crop Journal. 2016;18(4):987-999
  80. 80. Anagholi A, Rousta MJ, Azari A. Selection of salt tolerance varitieis of cotton by using of tolerant indices. Arid Bio Scientific and Research Journal. 2016;2:1-9
  81. 81. Afrasiab P, Delbari M, Asadi R, Mohammadi E. Effect of soil section and water salinity on yield and components yield of cotton. Journal of Plant Production Research. 2015;22(3):295-311
  82. 82. Razaji A, Paknejad F, Moarefi M, Mahdavi Damghani A, Ilkaee M. Meta-analysis of the effects of salinity stress on cotton (Gossypium spp.) growth and yield in Iran. Journal of. Agricultural Sciences. 2020;26(1):94-103
  83. 83. Wang Z, Wang J, Bao Y, Wu Y, Zhang H. Quantitative trait loci controlling rice seed germination under salt stress. Euphytica. 2011;178(3):297-307. DOI: 10.1007/s10681-010-0287-8
  84. 84. Zhang F, Zhu G, Du L, Shang X, Cheng C, Yang B, et al. Genetic regulation of salt stress tolerance revealed by RNA-Seq in cotton diploid wild species, Gossypium davidsonii. Scientific Reports. 2016;6(1):1-5. DOI: 10.1038/srep20582
  85. 85. Khan AN, Qureshi RH, Ahmad N, Rashid A. Response of cotton cultivars to salinity at various growth development stages. Sarhad Journal of Agriculture. 1995;11(6):729-731
  86. 86. Niu X, Bressan RA, Hasegawa PM, Pardo JM. Ion homeostasis in NaCl stress environments. Plant Physiology. 1995;109(3):735. DOI: 10.1104/pp.109.3.735
  87. 87. Roy SJ, Negrão S, Tester M. Salt resistant crop plants. Current Opinion in Biotechnology. 2014;26:115-124. DOI: 10.1016/j.copbio.2013.12.004
  88. 88. Deinlein U, Stephan AB, Horie T, Luo W, Xu G, Schroeder JI. Plant salt-tolerance mechanisms. Trends in Plant Science. 2014;19(6):371-379. DOI: 10.1016/j.tplants.2014.02.001
  89. 89. Chen X, Lu X, Shu N, Wang D, Wang S, Wang J, et al. GhSOS1, a plasma membrane Na+/H+ antiporter gene from upland cotton, enhances salt tolerance in transgenic Arabidopsis thaliana. PLoS One. 2017;12(7):e0181450. DOI: 10.1371/journal.pone.0181450
  90. 90. Faiyue B, Al-Azzawi MJ, Flowers TJ. A new screening technique for salinity resistance in rice (Oryza sativa L.) seedlings using bypass flow. Plant, Cell & Environment. 2012;35(6):1099-1108. DOI: 10.1111/j.1365-3040.2011.02475.x
  91. 91. Thorne SJ, Hartley SE, Maathuis FJ. Is silicon a panacea for alleviating drought and salt stress in crops? Frontiers in plant science. 2020;11:1221. DOI: 10.3389/fpls.2020.01221
  92. 92. Liu S, Guo X, Feng G, Maimaitiaili B, Fan J, He X. Indigenous arbuscular mycorrhizal fungi can alleviate salt stress and promote growth of cotton and maize in saline fields. Plant and Soil. 2016;398(1-2):195-206. DOI: 10.1007/s11104-015-2656-5
  93. 93. Khaitov B, Teshaev S. The effect of arbuscular mycorrhiza fungi on cotton growth and yield under salinated soil condition. Cotton Genomics and Genetics. 2015;17:6. DOI: 10.5376/cgg.2015.06.0003
  94. 94. Evelin H, Kapoor R, Giri B. Arbuscular mycorrhizal fungi in alleviation of salt stress: A review. Annals of Botany. 2009;104(7):1263-1280. DOI: 10.1093/aob/mcp251
  95. 95. Egamberdieva D, Jabborova D, Hashem A. Pseudomonas induces salinity tolerance in cotton (Gossypium hirsutum) and resistance to Fusarium root rot through the modulation of indole-3-acetic acid. Saudi Journal of Biological Sciences. 2015;22(6):773-779. DOI: 10.1016/j.sjbs.2015.04.019
  96. 96. Kang A, Zhang N, Xun W, Dong X, Xiao M, Liu Z, et al. Nitrogen fertilization modulates beneficial rhizosphere interactions through signaling effect of nitric oxide. Plant Physiology. 2021;1:kiab555
  97. 97. Jiang D, Lu B, Liu L, Duan W, Meng Y, Li J, et al. Exogenous melatonin improves the salt tolerance of cotton by removing active oxygen and protecting photosynthetic organs. BMC Plant Biology. 2021;21(1):1-9. DOI: 10.1186/s12870-021-03082-7
  98. 98. Venturas MD, Sperry JS, Hacke UG. Plant xylem hydraulics: What we understand, current research, and future challenges. Journal of Integrative Plant Biology. 2017;59(6):356-389. DOI: 10.1111/jipb.12534
  99. 99. Li X, Smith R, Choat B, Tissue DT. Drought resistance of cotton (Gossypium hirsutum) is promoted by early stomatal closure and leaf shedding. Functional Plant Biology. 2019;47(2):91-98. DOI: 10.1071/FP19093
  100. 100. Choat B, Brodribb TJ, Brodersen CR, Duursma RA, López R, Medlyn BE. Triggers of tree mortality under drought. Nature. 2018;558(7711):531-544. DOI: 10.1038/s41586-018-0240-x
  101. 101. Azhar MT, Rehman A. Overview on Effects of Water Stress on Cotton Plants and Productivity. In: Biochemical, Physiological and Molecular Avenues for Combating Abiotic Stress Tolerance in Plants. Cambridge: Academic Press; 2018. pp. 297-316. DOI: 10.1016/B978-0-12-813066-7.00016-4
  102. 102. Hameed A, Goher M, Iqbal N. Drought induced programmed cell death and associated changes in antioxidants, proteases, and lipid peroxidation in wheat leaves. Biologia Plantarum. 2013;57(2):370-374. DOI: 10.1007/s10535-012-0286-9
  103. 103. Fradin EF, Thomma BP. Physiology and molecular aspects of Verticillium wilt diseases caused by V. dahliae and V. albo-atrum. Molecular Plant Pathology. 2006;7(2):71-86. DOI: 10.1111/j.1364-3703.2006.00323.x
  104. 104. Hou S, Zhu G, Li Y, Li W, Fu J, Niu E, et al. Genome-wide association studies reveal genetic variation and candidate genes of drought stress related traits in cotton (Gossypium hirsutum L.). Frontiers. Plant Science. 2018;9:1276. DOI: 10.3389/fpls.2018.01276
  105. 105. Zhang JB, Wang XP, Wang YC, Chen YH, Luo JW, Li DD, et al. Genome-wide identification and functional characterization of cotton (Gossypium hirsutum) MAPKKK gene family in response to drought stress. BMC Plant Biology. 2020;20(1):1-4. DOI: 10.1186/s12870-020-02431-2
  106. 106. Wang C, Lu W, He X, Wang F, Zhou Y, Guo X, et al. The cotton mitogen-activated protein kinase 3 functions in drought tolerance by regulating stomatal responses and root growth. Plant and Cell Physiology. 2016;57(8):1629-1642. DOI: 10.1093/pcp/pcw090
  107. 107. Li J, Meng B, Chai H, Yang X, Song W, Li S, et al. Arbuscular mycorrhizal fungi alleviate drought stress in C3 (Leymus chinensis) and C4 (Hemarthria altissima) grasses via altering antioxidant enzyme activities and photosynthesis. Frontiers in Plant Science. 2019;10:499. DOI: 10.3389/fpls.2019.00499
  108. 108. Majid M, Ali M, Shahzad K, Ahmad F, Ikram RM, Ishtiaq M, et al. Mitigation of osmotic stress in cotton for the ımprovement in growth and yield through ınoculation of rhizobacteria and phosphate solubilizing bacteria coated diammonium phosphate. Sustainability. 2020;12(24):10456. DOI: 10.3390/su122410456
  109. 109. Iqbal A, Dong Q , Wang X, Gui H, Zhang H, Zhang X, et al. High nitrogen enhance drought tolerance in cotton through antioxidant enzymatic activities, nitrogen metabolism and osmotic adjustment. Plants. 2020;9(2):178. DOI: 10.3390/plants9020178
  110. 110. Singh M, Singh VP, Prasad SM. Responses of photosynthesis, nitrogen and proline metabolism to salinity stress in Solanum lycopersicum under different levels of nitrogen supplementation. Plant Physiology and Biochemistry. 2016;109:72-83. DOI: 10.1016/j.plaphy.2016.08.021
  111. 111. Zahoor R, Zhao W, Abid M, Dong H, Zhou Z. Potassium application regulates nitrogen metabolism and osmotic adjustment in cotton (Gossypium hirsutum L.) functional leaf under drought stress. Journal of Plant Physiology. 2017b;215:30-38. DOI: 10.1016/j.jplph.2017.05.001
  112. 112. Wu S, Hu C, Tan Q , Li L, Shi K, Zheng Y, et al. Drought stress tolerance mediated by zinc-induced antioxidative defense and osmotic adjustment in cotton (Gossypium hirsutum). Acta Physiologiae Plantarum. 2015;37(8):1-9. DOI: 10.1007/s1173 8-015-1919-3
  113. 113. Zahoor R, Dong H, Abid M, Zhao W, Wang Y, Zhou Z. Potassium fertilizer improves drought stress alleviation potential in cotton by enhancing photosynthesis and carbohydrate metabolism. Environmental and Experimental Botany. 2017c;137:73-83. DOI: 10.1016/j.envexpbot.2017.02.002
  114. 114. Ratnayaka HH, Molin WT, Sterling TM. Physiological and antioxidant responses of cotton and spurred anoda under interference and mild drought. Journal of Experimental Botany. 2003;54(391):2293-2305. DOI: 10.1093/jxb/erg251
  115. 115. Shallan MA, Hassan HM, Namich AA, Ibrahim AA. Biochemical and physiological effects of TiO2 and SiO2 nanoparticles on cotton plant under drought stress. Research Journal of Pharmaceutical Biological and Chemical Sciences. 2016;7(4):1540-1551
  116. 116. Dabbert TA, Gore MA. Challenges and perspectives on improving heat and drought stress resilience in cotton. Journal of Cotton Science. 2014;18(3):393-409
  117. 117. Dever JK. Cotton breeding and agro-technology. In: Handbook of Natural Fibres. Sawston: Woodhead Publishing; 2012. pp. 469-507. DOI: 10.1533/9780857095503.2.469
  118. 118. Ashraf M, Harris P. Breeding for abiotic stress tolerance in cotton. In: Ashraf M, Harris PJC, editors. Abiotic stresses: Plant Resistance through Breeding and Molecular Approaches. Binghamton: Food Products Press; 2005. pp. 595-613
  119. 119. Wei Y, Xu Y, Lu P, Wang X, Li Z, Cai X, et al. Salt stress responsiveness of a wild cotton species (Gossypium klotzschianum) based on transcriptomic analysis. PLoS One. 2017;12(5):e0178313. DOI: 10.1371/journal.pone.0178313
  120. 120. Chen Y, Feng S, Zhao T, Zhou B. Overcoming obstacles to interspecific hybridization between Gossypium hirsutum and G. turneri. Euphytica. 2018;214(2):1-5. DOI: 10.1007/s10681-018-2118-2
  121. 121. Hu G, Grover CE, Yuan D, Dong Y, Miller E, Conover JL, et al. Evolution and diversity of the cotton genome. In: Rahman M, Zafar Y, Zhang T, editors. Cotton Precision Breeding. Cham: Springer; 2021. pp. 25-78. DOI: 10.1007/978-3-030-64504-5_2
  122. 122. Fan X, Guo Q , Xu P, Gong Y, Shu H, Yang Y, et al. Transcriptome-wide identification of salt-responsive members of the WRKY gene family in Gossypium aridum. PLoS One. 2015;10(5):e0126148. DOI: 10.1371/journal.pone.0126148
  123. 123. Oluoch G, Zheng J, Wang X, Khan MK, Zhou Z, Cai X, et al. QTL mapping for salt tolerance at seedling stage in the interspecific cross of Gossypium tomentosum with Gossypium hirsutum. Euphytica. 2016;209(1):223-235. DOI: 10.1007/s10681-016-1674-6
  124. 124. Mammadov J, Buyyarapu R, Guttikonda SK, Parliament K, Abdurakhmonov IY, Kumpatla SP. Wild relatives of maize, rice, cotton, and soybean: Treasure troves for tolerance to biotic and abiotic stresses. Frontiers in Plant Science. 2018;9:886. DOI: 10.3389/fpls.2018.00886
  125. 125. CICR. Abiotic Stresses in Cotton: A Physiological Approach [Internet]. 2000. CICR Tech Bull. 2:1-13. Available from: http://www.cicr.org.in/pdf/abiotic_stress.pdf [Accessed: November 16, 2021]
  126. 126. Ahmad F, Perveen A, Mohammad N, Ali MA, Akhtar MN, Shahzad K, et al. Heat stress in cotton: Responses and adaptive mechanisms. In: Ahmad S, Hasanuzzaman M, editors. Cotton Production and Uses. Singapore: Springer; 2020. pp. 393-428. DOI: 10.1007/978-981-15-1472-2_20
  127. 127. Moreno AA, Orellana A. The physiological role of the unfolded protein response in plants. Biological Research. 2011;44(1):75-80. DOI: 10.4067/S0716-97602011000100010
  128. 128. Zafar MM, Manan A, Razzaq A, Zulfqar M, Saeed A, Kashif M, et al. Exploiting agronomic and biochemical traits to develop heat resilient cotton cultivars under climate change scenarios. Agronomy. 2021 Sep;11(9):1885. DOI: 10.3390/agronomy11091885
  129. 129. Hassan IS, El-Shaarawy SA, Abou-Tour HB. Response of some Egyptian genotypes to varied climatic measurements over varied environments. In: Proceedings Beltwide Cotton Conferences; 2000; San Antonio, USA, 553-560.
  130. 130. Abro S, Rajput MT, Khan MA, Sial MA, Tahir SS. Screening of cotton (Gossypium hirsutum L.) genotypes for heat tolerance. Pakistan Journal of Botany. 2015;47(6):2085-2091
  131. 131. Ekinci R, Başbağ S, Karademir E, Karademir C. The effects of high temperature stress on some agronomic characters in cotton. Pakistan Journal of Botany. 2017;49(2):503-508
  132. 132. Goren H. Comparison of Heat Stress Response in Cotton (Gossypium spp.) Genotypes in Diyarbakir Conditions. Aydin-Turkey: Aydin Adnan Menderes University; 2017
  133. 133. McDowell AJ, Bange MP, Tan DK. Cold temperature exposure at 10 C for 10 and 20 nights does not reduce tissue viability in vegetative and early flowering cotton plants. Australian Journal of Experimental Agriculture. 2007;47(2):198-207. DOI: 10.1071/ea05371
  134. 134. Cottee NS, Tan DK, Bange MP, Cothren JT, Campbell LC. Multi-level determination of heat tolerance in cotton (Gossypium hirsutum L.) under field conditions. Crop Science. 2010;50(6):2553-2564. DOI: 10.2135/cropsci2010.03.0182
  135. 135. Jaconis SY, Thompson AJ, Smith SL, Trimarchi C, Cottee NS, Bange MP, et al. A standardised approach for determining heat tolerance in cotton using triphenyl tetrazolium chloride. Scientific Reports. 2021;11(1):1-4. DOI: 10.1038/s41598-021-84798-2
  136. 136. Saha S, Jenkins JN, Wu J, McCarty JC, Gutiérrez OA, Percy RG, et al. Effects of chromosome-specific introgression in upland cotton on fiber and agronomic traits. Genetics. 2006;172(3):1927-1938. DOI: 10.1534/genetics.105.053371
  137. 137. Zhai H, Gong W, Tan Y, Liu A, Song W, Li J, et al. Identification of chromosome segment substitution lines of Gossypium barbadense introgressed in G. hirsutum and quantitative trait locus mapping for fiber quality and yield traits. PLoS One. 2016;11(9):e0159101
  138. 138. Kakani VG, Reddy KR, Koti S, Wallace TP, Prasad PV, Reddy VR, et al. Differences in in vitro pollen germination and pollen tube growth of cotton cultivars in response to high temperature. Annals of Botany. 2005;96(1):59-67. DOI: 10.1093/aob/mci149
  139. 139. Birrer KF, Conaty WC, Cottee NS, Sargent D, Francis ME, Cahill DM, et al. Can heat stress and water deficit affect cotton fiber wax content in field-grown plants? Industrial Crops and Products. 2021;168:113559. DOI: 10.1016/j.indcrop.2021.113559
  140. 140. Ur-Rahman H. Environmental interaction, additive and non-additive genetic variability is involved in the expression of tissue and whole-plant heat tolerance in upland cotton (Gossypium hirsutum L). Genetics and Molecular Biology. 2006;29(3):525-532. DOI: 10.1590/S1415-47572006000300022
  141. 141. Salman M, Zia ZU, Rana IA, Maqsood RH, Ahmad S, Bakhsh A, et al. Genetic effects conferring heat tolerance in upland cotton (Gossypium hirsutum L.). Journal of Cotton Research. 2019;2(1):1-8. DOI: 10.1186/s42397-019-0025-2
  142. 142. Amjad F, Amir S, Saghir A, Khan TM, Afzal AI. Evaluation of breeding potential of cotton germplasm of Pakistan origin for fiber quality traits under heat stress. International Journal of Agriculture and Biology. 2020;23(2):311-318. DOI: 10.17957/IJAB/15.1290
  143. 143. Manan A, Zafar MM, Ren M, Khurshid M, Sahar A, Rehman A, et al. Genetic analysis of biochemical, fiber yield and quality traits of upland cotton under high-temperature. Plant Production Science. 2021;9:1-5. DOI: 10.1080/1343943X.2021.1972013
  144. 144. Shakeel A, Naeem M, Imtiaz AL, Allah SU, Afzal I, Saeed A, et al. Genetic mechanism controlling selected within boll yield components and physiological traits of Gossypium hirsutum L. under salinity stress. Turkish Journal of Field Crops. 2017;22(1):89-95
  145. 145. Blum A. Breeding methods for drought resistance. In: Jones HG, Flowers TJ, Jones MB, editors. Plant under Stress. Cambridge: Cambridge University Press; 1989. pp. 197-216
  146. 146. Xue-yan ZH, Chuan-liang LI, Jun-juan WA, Fu-Guang L, Wu-wei YE. Drought-tolerance evaluation of cotton with PEG water-stress method. 棉花学报. 2008;20(1):56-61
  147. 147. Jaafar KS, Mohammed MA, Mohammed SM. Screening for drought tolerance in cotton (Gossypium hirsutum L.) using in vitro technique. Journal of Dryland Agriculture. 2021;7(4):52-59. DOI: 10.5897/JODA2021.0067
  148. 148. Lambers H, Chapin FS, Pons TL. Photosynthesis. In: Plant Physiological Ecology. New York: Springer; 2008. pp. 11-99. DOI: 10.1007/978-0-387-78341-3_2
  149. 149. Mohan MM, Narayanan SL, Ibrahim SM. Chlorophyll stability index (CSI): Its impact on salt tolerance in rice. International Rice Research Notes. 2000;25(2):38-39
  150. 150. Baytar AA, Peynircioğlu C, Sezener V, Basal H, Frary A, Frary A, et al. Identification of stable QTLs for fiber quality and plant structure in Upland cotton (G. hirsutum L.) under drought stress. Industrial Crops and Products. 2018;124:776-786. DOI: 10.1016/j.indcrop.2018.08.054
  151. 151. Magwanga RO, Lu P, Kirungu JN, Lu H, Wang X, Cai X, et al. Characterization of the late embryogenesis abundant (LEA) proteins family and their role in drought stress tolerance in upland cotton. BMC Genetics. 2018;19(1):1-31. DOI: 10.1186/s12863-017-0596-1
  152. 152. Basal H, Unay A. Water stress in cotton (Gossypium hirsutum L.). Journal of Agricultural Faculty of Ege University. 2006;43(3):101-111
  153. 153. Rodriguez-Uribe L, Abdelraheem A, Tiwari R, Sengupta-Gopalan C, Hughs SE, Zhang J. Identification of drought-responsive genes in a drought-tolerant cotton (Gossypium hirsutum L.) cultivar under reduced irrigation field conditions and development of candidate gene markers for drought tolerance. Molecular Breeding. 2014;34(4):1777-1796. DOI: 10.1007/s11032-014-0138-8
  154. 154. Shukla RP, Tiwari GJ, Joshi B, Song-Beng K, Tamta S, Boopathi NM, et al. GBS-SNP and SSR based genetic mapping and QTL analysis for drought tolerance in upland cotton. Physiology and Molecular Biology of Plants. 2021;27(8):1731-1745. DOI: 10.1007/s12298-021-01041-y
  155. 155. Anwar M, Saleem MA, Dan M, Malik W, Ul-Allah S, Ahmad MQ , et al. Morphological, physiological and molecular assessment of cotton for drought tolerance under field conditions. Saudi Journal of Biological Sciences. 2022;29(1):444-452. DOI: 10.1016/j.sjbs.2021.09.009
  156. 156. Mahmood T, Ahmar S, Abdullah M, Iqbal MS, Yasir M, Khalid S, et al. Genetic potential and inheritance pattern of phenological growth and drought tolerance in cotton (Gossypium hirsutum L.). Frontiers in Plant Science. 2021;12:705392. DOI: 10.3389/fpls.2021.705392
  157. 157. Gouia H, Ghorbal MH, Touraine B. Effects of NaCl on flows of N and mineral ions and on NO3-reduction rate within whole plants of salt-sensitive bean and salt-tolerant cotton. Plant Physiology. 1994;105(4):1409-1418. DOI: 10.1104/pp.105.4.1409
  158. 158. Higbie SM, Wang F, Stewart JM, Sterling TM, Lindemann WC, Hughs E, et al. Physiological response to salt (NaCl) stress in selected cultivated tetraploid cottons. International Journal of Agronomy. 2010;4:643475. DOI: 10.1155/2010/643475
  159. 159. Munawar W, Hameed A, Khan MK. Differential morphophysiological and biochemical responses of cotton genotypes under various salinity stress levels during early growth stage. Frontiers in Plant Science. 2021;12:622309. DOI: 10.3389/fpls.2021.622309
  160. 160. Ashraf M, Ahmad S. Genetic effects for yield components and fibre characteristics in upland cotton (Gossypium hirsutum L.) cultivated under salinized (NaCl) conditions. Agronomie. 2000;20(8):917-926
  161. 161. Abdelraheem A, Kuraparthy V, Hinze L, Stelly D, Wedegaertner T, Zhang J. Genome-wide association study for tolerance to drought and salt tolerance and resistance to thrips at the seedling growth stage in US Upland cotton. Industrial Crops and Products. 2021;169:113645. DOI: 10.1016/j.indcrop.2021.113645
  162. 162. Zafar MM, Razzaq A, Farooq MA, Rehman A, Firdous H, Shakeel A, et al. Genetic variation studies of ıonic and within boll yield components in cotton (Gossypium hirsutum L.) under salt stress. Journal of Natural Fibers. 2020;25:1-20. DOI: 10.1080/15440478.2020.1838996
  163. 163. Hosseini G, Thengane RJ. Salinity tolerance in cotton (Gossypium hirsutum L.) genotypes. International Journal of Botany. 2007;3(1):48-55
  164. 164. Dehdari A, Rezai A, Maibody MM. Nuclear and cytoplasmic inheritance of salt tolerance in bread wheat plants based on ion contents and biological yield. Iran Agricultural Research. 2007;24(1):15-26
  165. 165. Fan Y, Shabala S, Ma Y, Xu R, Zhou M. Using QTL mapping to investigate the relationships between abiotic stress tolerance (drought and salinity) and agronomic and physiological traits. BMC Genomics. 2015;16(1):43. DOI: 10.1186/s12864-015-1243-8
  166. 166. Abdelraheem A, Fang DD, Zhang J. Quantitative trait locus mapping of drought and salt tolerance in an introgressed recombinant inbred line population of upland cotton under the greenhouse and field conditions. Euphytica. 2018;214(1):1-20. DOI: 10.1007/s10681-017-2095-x
  167. 167. Turan S, Cornish K, Kumar S. Salinity tolerance in plants: Breeding and genetic engineering. Australian Journal of Crop Science. 2012;6(9):1337-1348
  168. 168. Esmaeili N, Cai Y, Tang F, Zhu X, Smith J, Mishra N, et al. Towards doubling fibre yield for cotton in the semiarid agricultural area by increasing tolerance to drought, heat and salinity simultaneously. Plant Biotechnology Journal. 2021;19(3):462-476. DOI: 10.1111/pbi.13476
  169. 169. Zhang L, Xi D, Li S, Gao Z, Zhao S, Shi J, et al. A cotton group C MAP kinase gene, GhMPK2, positively regulates salt and drought tolerance in tobacco. Plant Molecular Biology. 2011;77(1):17-31. DOI: 10.1007/s11103-011-9788-7
  170. 170. Ma W, Zhao T, Li J, Liu B, Fang L, Hu Y, et al. Identification and characterization of the GhHsp20 gene family in Gossypium hirsutum. Scientific Reports. 2016;6(1):1-3. DOI: 10.1038/srep32517
  171. 171. Maqbool A, Zahur M, Irfan M, Qaiser U, Rashid B, Husnain T, et al. Identification, characterization and expression of drought related alpha-crystalline heat shock protein gene (GHSP26) from Desi Cotton. Crop Science. 2007;47(6):2437-2444. DOI: 10.2135/cropsci2007.03.0120
  172. 172. Su Y, Guo A, Huang Y, Wang Y, Hua J. GhCIPK6a increases salt tolerance in transgenic upland cotton by involving in ROS scavenging and MAPK signaling pathways. BMC Plant Biology. 2020;20(1):1-9. DOI: 10.1186/s12870-020-02548-4
  173. 173. Yao D, Zhang X, Zhao X, Liu C, Wang C, Zhang Z, et al. Transcriptome analysis reveals salt-stress-regulated biological processes and key pathways in roots of cotton (Gossypium hirsutum L.). Genomics. 2011;98(1):47-55. DOI: 10.1016/j.ygeno.2011.04.007
  174. 174. Delgado C, Mora-Poblete F, Ahmar S, Chen JT, Figueroa CR. Jasmonates and plant salt stress: Molecular players, physiological effects, and improving tolerance by using genome-associated tools. International Journal of Molecular Sciences. 2021;22(6):3082. DOI: 10.3390/ijms22063082
  175. 175. Wu H, Zheng L, Qanmber G, Guo M, Wang Z, Yang Z. Response of phytohormone mediated plant homeodomain (PHD) family to abiotic stress in upland cotton (Gossypium hirsutum spp.). BMC Plant Biology. 2021;21(1):1-20. DOI: 10.1186/s12870-020-02787-5
  176. 176. Yan J, He C, Wang J, Mao Z, Holaday SA, Allen RD, et al. Overexpression of the Arabidopsis 14-3-3 protein GF14λ in cotton leads to a “stay-green” phenotype and improves stress tolerance under moderate drought conditions. Plant and Cell Physiology. 2004;45(8):1007-1014. DOI: 10.1093/pcp/pch115
  177. 177. Abdelrahman M, El-Sayed M, Jogaiah S, Burritt DJ, Tran LS. The “STAY-GREEN” trait and phytohormone signaling networks in plants under heat stress. Plant Cell Reports. 2017;36(7):1009-1025. DOI: 10.1007/s00299-017-2119-y

Written By

Volkan Mehmet Cinar, Serife Balci and Aydın Unay

Submitted: 24 January 2022 Reviewed: 26 May 2022 Published: 18 June 2022