Evolutionary Origins of Drought Tolerance in Spermatophytes

It is commonly known that drought stress is a major constraint limiting crop production. Drought stress and associated drought tolerance mechanisms are therefore under intense investigation with the view to future production of drought tolerant crops. With an ever-growing population and variable climate, novel approaches need to be considered to sustainably feed future generations. In this context, definitions of drought tolerance are highly variable, which poses a major challenge for the systematic assessment of this trait across the plant kingdom. Furthermore, drought tolerance is a polygenic trait and understanding the evolution of this complex trait may inform us about patterns of gene gain and loss in relation to diverse drought adaptations. We look at the transition of plants from water to land, and the role of drought tolerance in enabling this transition, before discussing the first drought tolerant plant and common drought responses amongst vascular plants. We reviewed the distribution of a combined “drought tolerance” trait in very broad terms to encompass different experimental systems and definitions used in the current literature and assigned a binary trait “tolerance vs. sensitivity” in 178 extant plant species. By simplifying drought responses of plants into this “binary” trait we were able to explore the evolution of drought tolerance across the wider plant kingdom, compared to previous studies. We show how this binary “drought tolerance/sensitivity” trait has evolved and discuss how incorporating this information into an evolutionary genomics framework could provide insights into the molecular mechanisms underlying extreme drought adaptations.

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Improving Drought Tolerance in Mungbean (Vigna radiata L. Wilczek): Morpho-Physiological, Biochemical and Molecular Perspectives

Agronomy ◽

10.3390/agronomy11081534 ◽

2021 ◽

Vol 11 (8) ◽

pp. 1534

Author(s):

Chandra Mohan Singh ◽

Poornima Singh ◽

Chandrakant Tiwari ◽

Shalini Purwar ◽

Mukul Kumar ◽

...

Keyword(s):

Drought Stress ◽

Drought Tolerance ◽

Crop Production ◽

Mitigation Strategies ◽

Theoretical Research ◽

Whole Genome Sequence ◽

Complex Nature ◽

Drought Tolerant ◽

Breeding Programmes ◽

The Impact

Drought stress is considered a severe threat to crop production. It adversely affects the morpho-physiological, biochemical and molecular functions of the plants, especially in short duration crops like mungbean. In the past few decades, significant progress has been made towards enhancing climate resilience in legumes through classical and next-generation breeding coupled with omics approaches. Various defence mechanisms have been reported as key players in crop adaptation to drought stress. Many researchers have identified potential donors, QTLs/genes and candidate genes associated to drought tolerance-related traits. However, cloning and exploitation of these loci/gene(s) in breeding programmes are still limited. To bridge the gap between theoretical research and practical breeding, we need to reveal the omics-assisted genetic variations associated with drought tolerance in mungbean to tackle this stress. Furthermore, the use of wild relatives in breeding programmes for drought tolerance is also limited and needs to be focused. Even after six years of decoding the whole genome sequence of mungbean, the genome-wide characterization and expression of various gene families and transcriptional factors are still lacking. Due to the complex nature of drought tolerance, it also requires integrating high throughput multi-omics approaches to increase breeding efficiency and genomic selection for rapid genetic gains to develop drought-tolerant mungbean cultivars. This review highlights the impact of drought stress on mungbean and mitigation strategies for breeding high-yielding drought-tolerant mungbean varieties through classical and modern omics technologies.

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Genome-wide transcriptional changes triggered by water deficit on a drought-tolerant common bean cultivar

BMC Plant Biology ◽

10.1186/s12870-020-02664-1 ◽

2020 ◽

Vol 20 (1) ◽

Cited By ~ 1

Author(s):

Josefat Gregorio Jorge ◽

Miguel Angel Villalobos-López ◽

Karen Lizeth Chavarría-Alvarado ◽

Selma Ríos-Meléndez ◽

Melina López-Meyer ◽

...

Keyword(s):

Cell Wall ◽

Drought Tolerance ◽

Common Bean ◽

Crop Production ◽

Molecular Mechanisms ◽

Molecular Data ◽

Transcriptional Level ◽

Cell Periphery ◽

Enriched Categories ◽

Drought Tolerant

Abstract Background Common bean (Phaseolus vulgaris L.) is a relevant crop cultivated over the world, largely in water insufficiency vulnerable areas. Since drought is the main environmental factor restraining worldwide crop production, efforts have been invested to amend drought tolerance in commercial common bean varieties. However, scarce molecular data are available for those cultivars of P. vulgaris with drought tolerance attributes. Results As a first approach, Pinto Saltillo (PS), Azufrado Higuera (AH), and Negro Jamapa Plus (NP) were assessed phenotypically and physiologically to determine the outcome in response to drought on these common bean cultivars. Based on this, a Next-generation sequencing approach was applied to PS, which was the most drought-tolerant cultivar to determine the molecular changes at the transcriptional level. The RNA-Seq analysis revealed that numerous PS genes are dynamically modulated by drought. In brief, 1005 differentially expressed genes (DEGs) were identified, from which 645 genes were up-regulated by drought stress, whereas 360 genes were down-regulated. Further analysis showed that the enriched categories of the up-regulated genes in response to drought fit to processes related to carbohydrate metabolism (polysaccharide metabolic processes), particularly genes encoding proteins located within the cell periphery (cell wall dynamics). In the case of down-regulated genes, heat shock-responsive genes, mainly associated with protein folding, chloroplast, and oxidation-reduction processes were identified. Conclusions Our findings suggest that secondary cell wall (SCW) properties contribute to P. vulgaris L. drought tolerance through alleviation or mitigation of drought-induced osmotic disturbances, making cultivars more adaptable to such stress. Altogether, the knowledge derived from this study is significant for a forthcoming understanding of the molecular mechanisms involved in drought tolerance on common bean, especially for drought-tolerant cultivars such as PS.

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Fine-tuning of SUMOylation modulates drought tolerance

10.1101/2021.03.29.437542 ◽

2021 ◽

Author(s):

Qingmei Guan ◽

Xuewei Li ◽

Shuang-Xi Zhou ◽

Liyuan Lu ◽

Huan Dang ◽

...

Keyword(s):

Drought Stress ◽

Drought Tolerance ◽

Molecular Mechanisms ◽

Plant Biology ◽

Fine Tuning ◽

Wild Type ◽

Tight Control ◽

Drought Tolerant ◽

Relationship Of ◽

The Relationship

SUMOylation is involved in various aspects of plant biology, including drought stress. However, the relationship between SUMOylation and drought stress tolerance is complex; whether SUMOylation has a crosstalk with ubiquitination in response to drought stress remains largely unclear. In this study, we found that both increased and decreased SUMOylation led to increased survival of apple (Malus × domestica) under drought stress: both transgenic MdSUMO2A overexpressing (OE) plants and MdSUMO2 RNAi plants exhibited enhanced drought tolerance. We further confirmed that MdDREB2A is one of the MdSUMO2 targets. Both transgenic MdDREB2A OE and MdDREB2AK192R OE plants (which lacked the key site of SUMOylation by MdSUMO2A) were more drought tolerant than wild-type plants. However, MdDREB2AK192R OE plants had a much higher survival rate than MdDREB2A OE plants. We further showed SUMOylated MdDREB2A was conjugated with ubiquitin by MdRNF4 under drought stress, thereby triggering its protein degradation. In addition, MdRNF4 RNAi plants were more tolerant to drought stress. These results revealed the molecular mechanisms that underlie the relationship of SUMOylation with drought tolerance and provided evidence for the tight control of MdDREB2A accumulation under drought stress mediated by SUMOylation and ubiquitination.

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Comparative Proteomics Analysis of the Seedling Root Response of Drought-sensitive and Drought-tolerant Maize Varieties to Drought Stress

International Journal of Molecular Sciences ◽

10.3390/ijms20112793 ◽

2019 ◽

Vol 20 (11) ◽

pp. 2793 ◽

Cited By ~ 3

Author(s):

Wenjing Zeng ◽

Yunling Peng ◽

Xiaoqiang Zhao ◽

Boyang Wu ◽

Fenqi Chen ◽

...

Keyword(s):

Drought Stress ◽

Drought Tolerance ◽

Molecular Mechanisms ◽

Mrna Level ◽

Differentially Expressed ◽

Water Retention Capacity ◽

Coding Region ◽

Retention Capacity ◽

Drought Tolerant ◽

Maize Varieties

The growth and development of maize roots are closely related to drought tolerance. In order to clarify the molecular mechanisms of drought tolerance between different maize (Zea mays L.) varieties at the protein level, the isobaric tags for relative and absolute quantitation (iTRAQ) quantitative proteomics were used for the comparative analysis of protein expression in the seedling roots of the drought-tolerant Chang 7-2 and drought-sensitive TS141 maize varieties under 20% polyethylene glycol 6000 (PEG 6000)-simulated drought stress. We identified a total of 7723 differentially expressed proteins (DEPs), 1243 were significantly differentially expressed in Chang 7-2 following drought stress, 572 of which were up-regulated and 671 were down-regulated; 419 DEPs were identified in TS141, 172 of which were up-regulated and 247 were down-regulated. In Chang 7-2, the DEPs were associated with ribosome pathway, glycolysis/gluconeogenesis pathway, and amino sugar and nucleotide sugar metabolism. In TS141, the DEPs were associated with metabolic pathway, phenylpropanoid biosynthesis pathway, and starch and sucrose metabolism. Compared with TS141, the higher drought tolerance of Chang 7-2 root system was attributed to a stronger water retention capacity; the synergistic effect of antioxidant enzymes; the strengthen cell wall; the osmotic stabilization of plasma membrane proteins; the effectiveness of recycling amino acid; and an improvement in the degree of lignification. The common mechanisms of the drought stress response between the two varieties included: The promotion of enzymes in the glycolysis/gluconeogenesis pathway; cross-protection against the toxicity of aldehydes and ammonia; maintenance of the cell membrane stability. Based on the proteome sequencing information, the coding region sequences of eight DEP-related genes were analyzed at the mRNA level by quantitative real-time PCR (qRT-PCR). The findings of this study can inform the future breeding of drought-tolerant maize varieties.

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The Plastid-Localized AtFtsHi3 Pseudo-Protease of Arabidopsis thaliana Has an Impact on Plant Growth and Drought Tolerance

Frontiers in Plant Science ◽

10.3389/fpls.2021.694727 ◽

2021 ◽

Vol 12 ◽

Author(s):

Laxmi S. Mishra ◽

Sanatkumar Mishra ◽

Daniel F. Caddell ◽

Devin Coleman-Derr ◽

Christiane Funk

Keyword(s):

Arabidopsis Thaliana ◽

Drought Tolerance ◽

Plant Growth ◽

Crop Production ◽

Molecular Mechanisms ◽

Wild Type ◽

Drought Tolerant ◽

Darwinian Fitness ◽

Intrinsic Water Use Efficiency ◽

Stomata Size

While drought severely affects plant growth and crop production, the molecular mechanisms of the drought response of plants remain unclear. In this study, we demonstrated for the first time the effect of the pseudo-protease AtFtsHi3 of Arabidopsis thaliana on overall plant growth and in drought tolerance. An AtFTSHi3 knock-down mutant [ftshi3-1(kd)] displayed a pale-green phenotype with lower photosynthetic efficiency and Darwinian fitness compared to wild type (Wt). An observed delay in seed germination of ftshi3-1(kd) was attributed to overaccumulation of abscisic acid (ABA); ftshi3-1(kd) seedlings showed partial sensitivity to exogenous ABA. Being exposed to similar severity of soil drying, ftshi3-1(kd) was drought-tolerant up to 20 days after the last irrigation, while wild type plants wilted after 12 days. Leaves of ftshi3-1(kd) contained reduced stomata size, density, and a smaller stomatic aperture. During drought stress, ftshi3-1(kd) showed lowered stomatal conductance, increased intrinsic water-use efficiency (WUEi), and slower stress acclimation. Expression levels of ABA-responsive genes were higher in leaves of ftshi3-1(kd) than Wt; DREB1A, but not DREB2A, was significantly upregulated during drought. However, although ftshi3-1(kd) displayed a drought-tolerant phenotype in aboveground tissue, the root-associated bacterial community responded to drought.

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Drought Tolerant near Isogenic Lines (NILs) of Pusa 44 Developed through Marker Assisted Introgression of qDTY2.1 and qDTY3.1 Enhances Yield under Reproductive Stage Drought Stress

Agriculture ◽

10.3390/agriculture11010064 ◽

2021 ◽

Vol 11 (1) ◽

pp. 64

Author(s):

Priyanka Dwivedi ◽

Naleeni Ramawat ◽

Gaurav Dhawan ◽

Subbaiyan Gopala Krishnan ◽

Kunnummal Kurungara Vinod ◽

...

Keyword(s):

Drought Stress ◽

Drought Tolerance ◽

Grain Quality ◽

Reproductive Stage ◽

Recurrent Parent ◽

Near Isogenic Lines ◽

Improvement Project ◽

Isogenic Lines ◽

Drought Tolerant ◽

Rice Improvement

Reproductive stage drought stress (RSDS) is detrimental for rice, which affects its productivity as well as grain quality. In the present study, we introgressed two major quantitative trait loci (QTLs), namely, qDTY2.1 and qDTY3.1, governing RSDS tolerance in a popular high yielding non-aromatic rice cultivar, Pusa 44, through marker-assisted backcross breeding (MABB). Pusa 44 is highly sensitive to RSDS, which restricts its cultivation across drought-prone environments. Foreground selection was carried out using markers, RM520 for qDTY3.1 and RM 521 for qDTY2.1. Background selection was achieved with 97 polymorphic SSR markers in tandem with phenotypic selection to achieve faster recurrent parent genome (RPG) recovery. Three successive backcrosses followed by three selfings aided RPG recoveries of 98.6% to 99.4% among 31 near isogenic lines (NILs). Fourteen NILs were found to be significantly superior in yield and grain quality under RSDS with higher drought tolerance efficiency (DTE) than Pusa 44. Among these, the evaluation of two promising NILs in the multilocational trial during Kharif 2019 showed that they were significantly superior to Pusa 44 under reproductive stage drought stress, while performing on par with Pusa 44 under normal irrigated conditions. These di-QTL pyramided drought-tolerant NILs are in the final stages of testing the All India Coordinated Rice Improvement Project varietal trials for cultivar release. Alternately, the elite drought-tolerant Pusa 44 NILs will serve as an invaluable source of drought tolerance in rice improvement.

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Genetic Diversity of Selected Rice Genotypes under Water Stress Conditions

Plants ◽

10.3390/plants10010027 ◽

2020 ◽

Vol 10 (1) ◽

pp. 27

Author(s):

Mahmoud M. Gaballah ◽

Azza M. Metwally ◽

Milan Skalicky ◽

Mohamed M. Hassan ◽

Marian Brestic ◽

...

Keyword(s):

Genetic Diversity ◽

Water Stress ◽

Drought Stress ◽

Drought Tolerance ◽

Ssr Markers ◽

Similarity Index ◽

Stress Conditions ◽

Yield Potential ◽

Drought Tolerant ◽

Rice Genotypes

Drought is the most challenging abiotic stress for rice production in the world. Thus, developing new rice genotype tolerance to water scarcity is one of the best strategies to achieve and maximize high yield potential with water savings. The study aims to characterize 16 rice genotypes for grain and agronomic parameters under normal and drought stress conditions, and genetic differentiation, by determining specific DNA markers related to drought tolerance using Simple Sequence Repeats (SSR) markers and grouping cultivars, establishing their genetic relationship for different traits. The experiment was conducted under irrigated (normal) and water stress conditions. Mean squares due to genotype × environment interactions were highly significant for major traits. For the number of panicles/plants, the genotypes Giza179, IET1444, Hybrid1, and Hybrid2 showed the maximum mean values. The required sterility percentage values were produced by genotypes IET1444, Giza178, Hybrid2, and Giza179, while, Sakha101, Giza179, Hybrid1, and Hybrid2 achieved the highest values of grain yield/plant. The genotypes Giza178, Giza179, Hybrid1, and Hybrid2, produced maximum values for water use efficiency. The effective number of alleles per locus ranged from 1.20 alleles to 3.0 alleles with an average of 1.28 alleles, and the He values for all SSR markers used varied from 0.94 to 1.00 with an average of 0.98. The polymorphic information content (PIC) values for the SSR were varied from 0.83 to 0.99, with an average of 0.95 along with a highly significant correlation between PIC values and the number of amplified alleles detected per locus. The highest similarity coefficient between Giza181 and Giza182 (Indica type) was observed and are susceptible to drought stress. High similarity percentage between the genotypes (japonica type; Sakha104 with Sakha102 and Sakha106 (0.45), Sakha101 with Sakha102 and Sakha106 (0.40), Sakha105 with Hybrid1 (0.40), Hybrid1 with Giza178 (0.40) and GZ1368-S-5-4 with Giza181 (0.40)) was also observed, which are also susceptible to drought stress. All genotypes are grouped into two major clusters in the dendrogram at 66% similarity based on Jaccard’s similarity index. The first cluster (A) was divided into two minor groups A1 and A2, in which A1 had two groups A1-1 and A1-2, containing drought-tolerant genotypes like IET1444, GZ1386-S-5-4 and Hybrid1. On the other hand, the A1-2 cluster divided into A1-2-1 containing Hybrid2 genotype and A1-2-2 containing Giza179 and Giza178 at coefficient 0.91, showing moderate tolerance to drought stress. The genotypes GZ1368-S-5-4, IET1444, Giza 178, and Giza179, could be included as appropriate materials for developing a drought-tolerant variety breeding program. Genetic diversity to grow new rice cultivars that combine drought tolerance with high grain yields is essential to maintaining food security.

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Drought Stress in Grain Legumes: Effects, Tolerance Mechanisms and Management

Agronomy ◽

10.3390/agronomy11122374 ◽

2021 ◽

Vol 11 (12) ◽

pp. 2374

Author(s):

Marium Khatun ◽

Sumi Sarkar ◽

Farzana Mustafa Era ◽

A. K. M. Mominul Islam ◽

Md. Parvez Anwar ◽

...

Keyword(s):

Drought Stress ◽

Drought Tolerance ◽

Association Studies ◽

Stomatal Closure ◽

Grain Legumes ◽

Grain Legume ◽

Developmental Cycle ◽

Genome Wide Association Studies ◽

Tolerance Mechanisms ◽

Drought Tolerant

Grain legumes are important sources of proteins, essential micronutrients and vitamins and for human nutrition. Climate change, including drought, is a severe threat to grain legume production throughout the world. In this review, the morpho-physiological, physio-biochemical and molecular levels of drought stress in legumes are described. Moreover, different tolerance mechanisms, such as the morphological, physio-biochemical and molecular mechanisms of legumes, are also reviewed. Moreover, various management approaches for mitigating the drought stress effects in grain legumes are assessed. Reduced leaf area, shoot and root growth, chlorophyll content, stomatal conductance, CO2 influx, nutrient uptake and translocation, and water-use efficiency (WUE) ultimately affect legume yields. The yield loss of grain legumes varies from species to species, even variety to variety within a species, depending upon the severity of drought stress and several other factors, such as phenology, soil textures and agro-climatic conditions. Closure of stomata leads to an increase in leaf temperature by reducing the transpiration rate, and, so, the legume plant faces another stress under drought stress. The biosynthesis of reactive oxygen species (ROS) is the most detrimental effect of drought stress. Legumes can adapt to the drought stress by changing their morphology, physiology and molecular mechanism. Improved root system architecture (RSA), reduced number and size of leaves, stress-induced phytohormone, stomatal closure, antioxidant defense system, solute accumulation (e.g., proline) and altered gene expression play a crucial role in drought tolerance. Several agronomic, breeding both conventional and molecular, biotechnological approaches are used as management practices for developing a drought-tolerant legume without affecting crop yield. Exogenous application of plant-growth regulators (PGRs), osmoprotectants and inoculation by Rhizobacteria and arbuscular mycorrhizal fungi promotes drought tolerance in legumes. Genome-wide association studies (GWASs), genomic selection (GS), marker-assisted selection (MAS), OMICS-based technology and CRISPR/Cas9 make the breeding work easy and save time in the developmental cycle to get resistant legumes. Several drought-resistant grain legumes, such as the chickpea, faba bean, common bean and pigeon pea, were developed by different institutions. Drought-tolerant transgenic legumes, for example, chickpeas, are developed by introgressing desired genes through breeding and biotechnological approaches. Several quantitative trait loci (QTLs), candidate genes occupying drought-tolerant traits, are identified from a variety of grain legumes, but not all are under proper implementation. Hence, more research should be conducted to improve the drought-tolerant traits of grain legumes for avoiding losses during drought.

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Physiological Responses under Drought Stress of Improved Drought-Tolerant Rice Lines and their Parents

Notulae Botanicae Horti Agrobotanici Cluj-Napoca ◽

10.15835/nbha46211188 ◽

2018 ◽

Vol 46 (2) ◽

pp. 679-687 ◽

Cited By ~ 6

Author(s):

Preeyanuch LARKUNTHOD ◽

Noppawan NOUNJAN ◽

Jonaliza L SIANGLIW ◽

Theerayut TOOJINDA ◽

Jirawat SANITCHON ◽

...

Keyword(s):

Drought Stress ◽

Drought Tolerance ◽

Water Status ◽

Physiological Responses ◽

High Water ◽

Recurrent Parent ◽

Potential Candidate ◽

Substitution Lines ◽

Chromosome Segment Substitution Lines ◽

Drought Tolerant

Many of the economically important rice cultivars including ‘Khao Dawk Mali 105’ (KDML105) or jasmine rice, one of the world’s famous rice exported from Thailand suffers from drought due to erratic rainfalls and limited irrigation. To improve drought tolerance and reserve genetic background of KDML105, chromosome segment substitution lines (CSSL) containing drought tolerant quantitative trait loci (DT-QTL) has been previously developed by backcrossing between KDML105 and drought tolerant donor, IR58586-F2-CA-143 (DH212). To understand the physiological responses related to drought tolerance in CSSL lines compared to parents, two CSSLs namely CSSL1-16 and CSSL1-18, respectively were used in this study. Twenty-one-d-old hydroponically grown plants were subjected to 20% PEG for 0, 7, 14 d and then recovered from stress for 3 d. The results indicated that CSSL lines especially, CSSL1-16 showed better performance under drought stress compared to their recurrent parent. Drought tolerance superior CSSL1-16 line was indicated by high water status (high relative water content and leaf water potential), good osmotic adjustment, high proline and greater membrane stability. Moreover, this line was able to resume growth after stress recovery whereas other lines/cultivar could not recover. Similarly, drought tolerant donor showed high water status suggesting that well-maintained plant water status was associated with drought tolerant trait. It could be concluded that the highest drought tolerant line was CSSL1-16 followed by DH212, CSSL1-18 and KDML105. It would be interesting to go further into introgressed section in CSSL1-16 to identify potential candidate genes in DT-QTL for breeding drought tolerant rice in the future.

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Phytohormone cross-talk and antioxidant gadgets tussles osmotic stress in indigenous rice landraces during seed germination

10.21203/rs.3.rs-80159/v1 ◽

2020 ◽

Author(s):

R. Anupriya ◽

Sugitha Thankappan ◽

A. Senthil ◽

D. Rajakumar ◽

Geetha S ◽

...

Keyword(s):

Drought Stress ◽

Seed Germination ◽

Drought Tolerance ◽

Cross Talk ◽

Tamil Nadu ◽

Critical Role ◽

Hydrolytic Enzyme ◽

Rice Landraces ◽

Drought Tolerant ◽

Root And Shoot Length

Abstract Seed germination plays a critical role in determining rice productivity under drought stress. We evaluated 100 traditional rice landraces originated from different agro-ecological zones of Tamil Nadu along with drought- susceptible (IR 64) and drought- tolerant (IR 64 DRT) checks. Moisture stress was induced using PEG 6000 and screening done over a range of osmotic potentials (-) 10 bars, (-) 12.5 bars and (-)15 bars for a period of 5 d. Physio-morphological traits such as germination rate, survival per cent, root and shoot length, vigor index, RS ratio and relative water content (RWC) were assessed during early drought stress. We observed significant changes in the seed macromolecules, phytohormone levels (GA and IAA), osmolytes and antioxidant responses (catalase and superoxide dismutase) between drought stress and control treatments. Kuliyadichan registered significantly higher IAA and GA (44% and 35% respectively over drought tolerant check IR 64 DRT) at drought stress, whereas all the landraces showed an elevated catalase activity. In PC analysis, first three PCs captured 88.93% of the total variation; significant differences were detected among genotypes with respect to the studied parameters. Six traditional landraces such as Kuliyadichan, Rajalakshmi, Sabhagidhan, Nootripathu, Chandaikar and Mallikar were selected and their inherent drought tolerance was associated with metabolic responses viz., triggered hydrolytic enzyme activities, hormonal cross-talk, ROS signaling and catalase under drought stress compared to drought sensitive IR64. Hence, these genotypes can be used as potential donor candidates towards genetic improvement of drought tolerance in rice.

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