Leaf morphology,rather than plant water status,underlies genetic variation of rice leaf rolling under drought

Soil drying causes leaf rolling in rice, but the relationship between leaf rolling and drought tolerance has historically confounded selection of drought‐tolerant genotypes. In this study on tropical japonica and aus diversity panels (170–220 genotypes), the degree of leaf rolling under drought was more affected by leaf morphology than by stomatal conductance, leaf water status, or maintenance of shoot biomass and grain yield. A range of canopy temperature and leaf rolling (measured as change in normalized difference vegetation index [ΔNDVI]) combinations were observed among aus genotypes, indicating that some genotypes continued transpiration while rolled. Association mapping indicated colocation of genomic regions for leaf rolling score and ΔNDVI under drought with previously reported leaf rolling genes and gene networks related to leaf anatomy. The relatively subtle variation across these large diversity panels may explain the lack of agreement of this study with earlier reports that used small numbers of genotypes that were highly divergent in hydraulic traits driving leaf rolling differences. This study highlights the large range of physiological responses to drought among rice genotypes and emphasizes that drought response processes should be understood in detail before incorporating them into a varietal selection programme.     

辣椒开花结果期对干旱胁迫的形态与生理响应

在遮雨网室选用抗旱性较强的农城椒二号和抗旱性较弱的陕蔬2001,研究辣椒在轻度、中度和重度干旱胁迫下不同时间的生长、产量、渗透调节物质、保护酶活性的变化规律及其生理调节机制。结果表明:随干旱胁迫时间的延长,辣椒的株高、分枝数、叶面积、单位面积产量、叶绿素含量和叶片相对含水量的抗旱系数呈下降趋势,下降速率与干旱胁迫程度呈正相关,与品种的抗旱性呈负相关;脯氨酸、丙二醛含量和细胞膜透性相对值随干旱胁迫时间的延长呈上升趋势;POD、SOD、CAT活性和可溶性蛋白相对值随着干旱胁迫时间的延长先升高后下降,抗旱性强的材料增加幅度低于抗旱性弱的材料;可溶性糖含量的相对值在轻度和中度干旱胁迫下呈上升趋势,在重度干旱胁迫下呈上升—下降趋势,且抗旱性强的材料上升速度大于抗旱性弱的材料。相关分析表明,干旱胁迫下,产量与株高、分枝数、叶片叶绿素含量、叶面积、叶片相对含水量抗旱系数呈显著正相关;与细胞膜透性、CAT活性和可溶性蛋白含量抗旱系数呈显著负相关。主成分分析表明,用作辣椒抗旱性鉴定的主要指标是单株产量、株高、叶面积、分枝数、可溶性蛋白、可溶性糖、MDA、叶绿素含量和细胞膜透性及叶片相对含水量,叶片POD、SOD、CAT活性、脯氨酸含量可做为辣椒抗旱性鉴定的次要鉴选指标。     

Genotypic variation in drought stress response and subsequent recovery of wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

Three wheat (Triticum aestivum L.) genotypes, Sadovo, Katya and Prelom, with different tolerance to drought were comparatively evaluated in terms of leaf respiratory responses to progressing dehydration and consecutive rewatering. Under drought stress, the respiration of all varieties gradually decreased, as the drought-tolerant Katya showed the most pronounced decline at earlier stages of dehydration. When water stress intensified, this genotype gave relatively stable respiration rates compared with the drought-sensitive varieties. Additionally, dehydrated Katya leaves displayed lower stomatal conductance and higher photosynthesis values, which resulted in greater water use efficiency during the dehydration period. Combination of drought stress and short-term changes in leaf temperature also induced genotype-specific response that differed from the response to drought only. Over the whole temperature range, the leaves of Katya exposed to dehydration for 14 days, showed higher respiration rates compared to the drought-sensitive varieties. The sensitive varieties maintained higher respiration rates under control conditions and mild dehydration, and very low rates under severe drought. In Katya, respiration and photosynthesis were fully restored from the stress within the first day of rewatering. The drought-sensitive genotypes displayed a considerably slower recovering capacity. The results are discussed in terms of possible physiological mechanisms underlying plant tolerance to drought.     

Evaluation of physiological screening tests for breeding drought resistant triticale (x <Emphasis Type="Italic">Triticosecale</Emphasis> wittmack)

Effects of soil drought on crop yield of 4 strains and 7 cultivars of spring triticale was investigated under field condition. The Drought Susceptibility Index (DSI) was evaluated in a two year experiment by the determination of grain loss in conditions of two soil moisture levels (drought-D and irrigated-IR). In the experiment response to drought was evaluated by different screening tests (leaf gaseous exchange, leaf water potential, chlorophyll content and fluorescence, leaf injury by drought and by simulated drought and heat temperature and water loss by excited leaf. The DSI values and the results of screening tests showed the genetic variation in the degree of drought tolerance. The values of DSI enabled the ranking of the tested triticale genotypes with respect to their drought tolerance and allow to divide them into three groups of drought susceptibility. Large differences among studied forms were observed also in changes of leaf water potential, fluorescence and leaf injury. For plants in vegetative stage of growth the tested breeding forms were easily separated into groups of different drought tolerance. Changes of ψ, Fv/Fm and LI as a screening tests were the most suitable techniques for estimation of degree of drought tolerance for triticale. Laboratory screening tests (leaf injury by simulated drought (LI_DS) and high temperature (LI_HT) and water loss (WL) of excited leaf conducted for nonstressed plants in most cases were significantly correlated with DSI. The statistically significant correlation between leaf water potential (ψ) was observed only with leaf fluorescence (Fv/Fm). Changes of Fv/Fm were significantly correlated with ψ, LI and LI_HT for 50 °C. Index of leaf injury (LI) by soil drought were significantly correlated with Fv/Fm, LI_DS (−1.0, −1.5 MPa), LI_HT (45 and 50°C) and water loss (WL). The correlation coefficient between the tests LI_DS and LI_HT were most of the considered cases statistically significant which indicate that the mechanism of membranes injury resulted from simulated drought or high temperature were similar in triticale. Water loss (WL) of excited leaves was the most suitable test for screening drought tolerance in triticale population. Changes of gaseous exchange parameters were not useful as screening test in this research.     

The physiological characteristics of the low canopy temperature wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) genotypes under simulated drought condition

Crop canopy temperature reflects the interactions among plants, soil and atmosphere. Through a long-term investigation into physiological characteristics of different canopy temperature wheat, it is found that some wheat exist a high canopy temperature while others a low one, which was closely correlated with their corresponding performance. Under simulated drought conditions, the physiological aspects of different canopy temperature wheat, such as leaf functional duration, chlorophyll content, activities of superoxide dismutase, protein content, transpiration rate, net photosynthesis rate, etc., were investigated, the result showed that wheat with low canopy temperature could maintain superiority to wheat with high canopy temperature in those physiological traits. Therefore, the low canopy temperature in wheat could be used as an index to evaluate physiological capacities of wheat under drought conditions and also as a useful marker for wheat breeding for drought tolerance.     

Overexpression of Arabidopsis Molybdenum Cofactor Sulfurase Gene Confers Drought Tolerance in Maize (Zea mays L.)

Abscisic acid (ABA) is a key component of the signaling system that integrates plant adaptive responses to abiotic stress. Overexpression of Arabidopsis molybdenum cofactor sulfurase gene (LOS5) in maize markedly enhanced the expression of ZmAO and aldehyde oxidase (AO) activity, leading to ABA accumulation and increased drought tolerance. Transgenic maize (Zea mays L.) exhibited the expected reductions in stomatal aperture, which led to decreased water loss and maintenance of higher relative water content (RWC) and leaf water potential. Also, transgenic maize subjected to drought treatment exhibited lower leaf wilting, electrolyte leakage, malondialdehyde (MDA) and H₂O₂ content, and higher activities of antioxidative enzymes and proline content compared to wild-type (WT) maize. Moreover, overexpression of LOS5 enhanced the expression of stress-regulated genes such as Rad 17, NCED1, CAT1, and ZmP5CS1 under drought stress conditions, and increased root system development and biomass yield after re-watering. The increased drought tolerance in transgenic plants was associated with ABA accumulation via activated AO and expression of stress-related gene via ABA induction, which sequentially induced a set of favorable stress-related physiological and biochemical responses.     

空气湿度与土壤水分胁迫对紫花苜蓿叶表皮蜡质特性的影响

选用2个抗旱性不同的紫花苜蓿品种,敖汉(强抗旱)和三得利(弱抗旱),设置空气湿度(45%-55%和75%-85%)和土壤水分胁迫(75%和35%田间持水量)处理,分析紫花苜蓿叶表皮蜡质含量、组分及晶体结构、气体交换参数、水势及脯氨酸含量的变化规律。结果表明,单独土壤水分胁迫时,紫花苜蓿叶表皮蜡质晶体结构及蜡质总量无显著变化;敖汉蜡质组分中烷类、酯类含量增加,醇类含量下降;三得利醇类含量下降,烷类、酯类含量变化不显著。低空气湿度胁迫时,两品种蜡质总量无显著变化,烷类和酯类含量显著增加,醇类含量显著下降,叶表皮片状蜡质晶体结构熔融呈弥漫性,扩大了对叶表面积的覆盖,其蒸腾速率显著低于正常湿度。复合胁迫处理时,叶表皮片状蜡质晶体结构继续呈弥漫性,烷类、酯类、未知蜡质组分含量均高于单独胁迫处理,醇类含量最低,而蜡质总量除三得利显著高于对照外,其余均无显著差异。紫花苜蓿叶表皮蜡质各组分含量(除醇类)及蜡质总量与光合速率呈显著负相关,与蒸腾速率无显著相关关系。蜡质总量与叶水势呈显著正相关。总体上,敖汉蜡质总量显著高于三得利,蜡质组分中烷类物质的增加有助于提高植株的抗旱性。在复合胁迫下,强抗旱品种主要通过气孔因素控制水分散失,而弱抗旱品种通过气孔和非气孔因素共同控制植物水分散失。     

Genotypic Differences in Leaf Water Potential, Abscisic Acid and Proline Concentrations in Spring Wheat during Drought Stress

Recent work with spring wheat has revealed significant genotypicvariation in changes of water potential and abscisic acid (ABA)concentration in response to drought Two experiments with eightspring wheat genotypes have been carried out to check the earlierwork on relationships between water potential and ABA concentrationand to examine causes of genotypic variation in the rate ofdecline of water potential during drought Changes in prolineconcentration were also studied Plants were grown in controlled environment cabinets with nutrientsolution culture and were stressed by withholding water as thefifth or sixth leaf on the main stem emerged. Plants were harvested4, 5 and 6 days after the treatment commenced and measurementsof leaf water potential, stomatal conductance, ABA and prolineconcentrations, and tissue d wts were taken. Significant genotypic variation was found in the decrease ofwater potential with time and in the slopes of linear regressionsof ABA concentration on water potential, confirming earlierresults When differences between leaf areas at the start of the treatmentwere minimised by varying the genotype sowing date significantgenotypic variation in water potentials at harvest was stillobtained. The change in water potential was significantly positivelycorrelated with shoot root d wt ratios at harvest and pre-treatmentstomatal conductances. Proline concentrations were significantly correlated with waterpotential for every genotype, although there was no clear evidenceof genotypic variation in proline concentrations at a givenwater potential The possible role of ABA concentration in drought resistanceof cereals is discussed Triticum aestivum L, spring wheat, water potential, abscisic acid, proline, drought stress     

Exploration of Distinct Physiological and Biochemical Alterations in Chickpea (<i>Cicer arietinum</i> L.) under Varying Organic Materials,Drought and Proline Application

Chickpea yield is decreasing day by day due to drought stress, which could be an immense risk for future food security in developing countries. Management practices could be the most excellent approach to diminish loss due to this abiotic factor. The current research work was designed to explore the tolerance reaction of chickpea genotypes against management practices, through morphological and biochemical parameters and evaluate yield performance across drought prone location of Bangladesh. Four genotypes BD-6048, BD-6045, BD-6090, BD-6092 and eight management practices, e.g., severe water stress (SWS), i.e., without irrigation, 10 cm thick mulching with rice straw (MRS), 10 cm thick mulching with water hyacinth (MWH), organic amendment through compost (OAC) @ 3 t ha⁻¹, organic amendment through cow dung @ 5 t ha⁻¹ (OACD), organic amendment through poultry manure @ 2 t ha⁻¹ (OAPM), inorganic amendment through proline application (IAPA) as foliar spray and 16 h hydro-priming (HP). The study revealed that the genotypes BD-6048 showed excellent performance because of the highest chlorophyll, carotenoids, phosphorus, potassium, proline and protein content. The highest pod number plant⁻¹ also increased seed yield in BD-6048. Considering management practices, IAPA increased relative water content, carotenoids, leaf phosphorus and potassium compared to other management practices and severe water stress. Finally, BD-6084 was selected as best genotype because of a significant increase in chlorophyll a and b, carotenoids, and relative water content with IAPA. Identified top performing genotypes can be used for releasing variety and cultivated for sustainable production in drought prone area of Bangladesh.     

Restriction of transpiration rate under high vapour pressure deficit and non‐limiting water conditions is important for terminal drought tolerance in cowpea

Drought stress is a major constraint on cowpea productivity, since the crop is grown under warm conditions on sandy soils having low water‐holding capacity. For enhanced performance of crops facing terminal drought stress, like cowpea, water‐saving strategies are crucial. In this work, the growth and transpiration rate (TR) of 40 cowpea genotypes with contrasting response to terminal drought were measured under well‐watered conditions across different vapour pressure deficits (VPD) to investigate whether tolerant and sensitive genotypes differ in their control of leaf water loss. A method is presented to indirectly assess TR through canopy temperature (CT) and the index of canopy conductance (Ig). Overall, plants developed larger leaf area under low than under high VPD, and there was a consistent trend of lower plant biomass in tolerant genotypes. Substantial differences were recorded among genotypes in TR response to VPD, with tolerant genotypes having significantly lower TR than sensitive ones, especially at times with the highest VPD. Genotypes differed in TR response to increasing VPD, with some tolerant genotypes exhibiting a clear VPD breakpoint at about 2.25 kPa, above which there was very little increase in TR. In contrast, sensitive genotypes presented a linear increase in TR as VPD increased, and the same pattern was found in some tolerant lines, but with a smaller slope. CT, estimated with thermal imagery, correlated well with TR and Ig and could therefore be used as proxy for TR. These results indicate that control of water loss discriminated between tolerant and sensitive genotypes and may, therefore, be a reliable indicator of terminal drought stress tolerance. The water‐saving characteristics of some genotypes are hypothesised to leave more soil water for pod filling, which is crucial for terminal drought adaptation.     

Brassinolide alleviated the adverse effect of water deficits on photosynthesis and the antioxidant of soybean (<Emphasis Type="Italic">Glycine max</Emphasis> L.)

Brassinolide (BR) is a relatively new plant growth regulator. To test whether BR could be used to increase tolerance to water deficits in soybean, the effects of BR application on photosynthesis, assimilate distribution, antioxidant enzymes and seed yield were studied. BR at 0.1 mg l⁻¹ was foliar applied at the beginning of bloom. Two levels of soil moisture (80% field capacity for well-watered control and 35% for drought-stressed treatment) were applied at pod initiation. BR treatment increased biomass accumulation and seed yield for both treatments. Drought stress inhibited translocation of assimilated ¹⁴C from the labeled leaf, but BR increased the translocation for both treatments. Drought stress depressed chlorophyll content and assimilation rate (A), while chlorophyll content and A of BR-treated plants were greater than that of drought-stressed plants. BR treatment increased maximum quantum yield of PS II, the activity of ribulose-1,5-bisphosphate carboxylase, and the leaf water potential of drought-stressed plants. Treatment with BR also increased the concentration of soluble sugars and proline, and the activities of peroxidase and superoxide dismutase of soybean leaves when drought-stressed. However, it decreased the malondialdehyde concentration and electrical conductivity of leaves under drought stress. This study show that BR can be used as a plant growth regulator to enhance drought tolerance and minimize the yield loss of soybean caused by water deficits.     

Photosynthetic apparatus activity in relation to high and low contents of cell wall-bound phenolics in triticale under drought stress

Cell wall-bound phenolics (CWP) play an important role in the mechanisms of plant acclimation to soil drought. The study involved CWP analyses in 50 strains and 50 doubled haploid (DH) lines of winter triticale exposed to drought at their vegetative and generative stages. CWP in the plants experiencing drought at the generative stage positively correlated with their leaf water contents. The strains and DH lines characterized by high content of CWP showed higher leaf water content and higher activity of photosynthetic apparatus when exposed to drought at the generative stage compared to the strains and DH lines with the low CWP content. Furthermore, when drought subsided at the generative stage, the strains and DH lines richer in CWP demonstrated higher regeneration potential and their grain yield loss was smaller.     

UPLC‐HRMS‐based untargeted metabolic profiling reveals changes in chickpea (Cicer arietinum) metabolome following long‐term drought stress

Genetic improvement for drought tolerance in chickpea requires a solid understanding of biochemical processes involved with different physiological mechanisms. The objective of this study is to demonstrate genetic variations in altered metabolic levels in chickpea varieties (tolerant and sensitive) grown under contrasting water regimes through ultrahigh‐performance liquid chromatography/high‐resolution mass spectrometry‐based untargeted metabolomic profiling. Chickpea plants were exposed to drought stress at the 3‐leaf stage for 25 days, and the leaves were harvested at 14 and 25 days after the imposition of drought stress. Stress produced significant reduction in chlorophyll content, F_v/F_m, relative water content, and shoot and root dry weight. Twenty known metabolites were identified as most important by 2 different methods including significant analysis of metabolites and partial least squares discriminant analysis. The most pronounced increase in accumulation due to drought stress was demonstrated for allantoin, l ‐proline, l ‐arginine, l ‐histidine, l ‐isoleucine, and tryptophan. Metabolites that showed a decreased level of accumulation under drought conditions were choline, phenylalanine, gamma‐aminobutyric acid, alanine, phenylalanine, tyrosine, glucosamine, guanine, and aspartic acid. Aminoacyl‐tRNA and plant secondary metabolite biosynthesis and amino acid metabolism or synthesis pathways were involved in producing genetic variation under drought conditions. Metabolic changes in light of drought conditions highlighted pools of metabolites that affect the metabolic and physiological adjustment in chickpea that reduced drought impacts.     

Morpho-physiological responses of sugar beet (Beta vulgaris L.) genotypes to drought stress

The identification of morpho-physiological traits related to drought tolerance and high yield potential is a challenge when selecting sugar beet genotypes with greater tolerance to water stress. In this paper, root morphological parameters, antioxidant systems, leaf relative water content (RWC) and H⁺-ATPase activity as key morpho-physiological traits involved in drought tolerance/susceptibility of sugar beet were studied. Genotypes showing a different drought tolerance index (DTI) but a similar yield potential, under moderate (?0.6 Mpa) and severe (?1.2 MPa) water stress, were selected and their morpho-physiological traits were investigated. The results showed a wide genetic variation in morpho-physiological parameters which demonstrated the different adaptive strategies under moderate and severe drought conditions in sugar beet. In particular, an efficient antioxidant system and redox signalling made some sugar beet genotypes more tolerant to drought stress. The alternative strategy of other genotypes was the reduction of root tissue density, which produced a less dense root system improving the axial hydraulic conductivity. These results could be considered as interesting challenge for a better understanding of the drought tolerance mechanisms in sugar beet.     

Effect of plant growth promoting <Emphasis Type="Italic">Pseudomonas</Emphasis> spp. on compatible solutes,antioxidant status and plant growth of maize under drought stress

Drought is one of the major abiotic stresses affecting yield of dryland crops. Rhizobacterial populations of stressed soils are adapted and tolerant to stress and can be screened for isolation of efficient stress adaptive/tolerant, plant growth promoting rhizobacterial (PGPR) strains that can be used as inoculants for crops grown in stressed ecosystems. The effect of inoculation of five drought tolerant plant growth promoting Pseudomonas spp. strains namely P. entomophila strain BV-P13, P. stutzeri strain GRFHAP-P14, P. putida strain GAP-P45, P. syringae strain GRFHYTP52, and P. monteilli strain WAPP53 on growth, osmoregulation and antioxidant status of maize seedlings under drought stress conditions was investigated. Drought stress induced by withholding irrigation had drastic effects on growth of maize seedlings. However seed bacterization of maize with Pseudomonas spp. strains improved plant biomass, relative water content, leaf water potential, root adhering soil/root tissue ratio, aggregate stability and mean weight diameter and decreased leaf water loss. The inoculated plants showed higher levels of proline, sugars, free amino acids under drought stress. However protein and starch content was reduced under drought stress conditions. Inoculation decreased electrolyte leakage compared to uninoculated seedlings under drought stress. As compared to uninoculated seedlings, inoculated seedlings showed significantly lower activities of antioxidant enzymes, ascorbate peroxidase (APX), catalase (CAT), glutathione peroxidase (GPX) under drought stress, indicating that inoculated seedlings felt less stress as compared to uninoculated seedlings. The strain GAP-P45 was found to be the best in terms of influencing growth and biochemical and physiological status of the seedlings under drought stress. The study reports the potential of rhizobacteria in alleviating drought stress effects in maize.     

Differential responses of the enzymes involved in proline biosynthesis and degradation in drought tolerant and sensitive cotton genotypes during drought stress and recovery

The relative water content (RWC), free proline levels and the activities of enzymes involved in proline metabolism were studied in drought tolerant (Ca/H 680) and drought sensitive (Ca/H 148) genotypes of cotton (Gossypium hirsutum L.) during induction of water stress and posterior recovery. Water stress caused a significant increase in proline levels and P5CS activity in leaves of both tolerant and sensitive genotypes, whereas the activity of P5CR increased minimally and the activity of OAT remains unchanged. The activity of PDH decreased under drought stress in both the genotypes. The leaf of tolerant genotype maintained higher RWC, photosynthetic activity and proline levels, as well as higher P5CS and P5CR activities under water stress than that of drought sensitive genotype. The drought induced proline levels and activities of P5CS and P5CR declined and tend to be equal to their respective controls, during recovery, whereas the PDH activity tends to increase. These results indicate that induction of proline levels by up regulation of P5CS and down regulation of PDH may be involved in the development of drought tolerance in cotton.     

QTLs linked to leaf epicuticular wax,physio-morphological and plant production traits under drought stress in rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

Drought stress is the major constraint to rice (Oryza sativa L.) production and yield stability in rainfed ecosystems. Identifying genomic regions contributing to drought resistance will help to develop rice cultivars suitable for rainfed regions through marker-assisted breeding. Quantitative trait loci (QTLs) linked to leaf epicuticular wax, physio-morphological and plant production traits under water stress and irrigated conditions were mapped in a doubled haploid (DH) line population from the cross CT9993-5-10-1-M/IR62266-42-6-2. The DH lines were subjected to water stress during anthesis. The DH lines showed significant variation for epicuticular wax (EW), physio-morphological and plant production traits under stress and irrigated conditions. A total of 19 QTLs were identified for the various traits under drought stress and irrigated conditions in the field, which individually explained 9.6%–65.6% of the phenotypic variation. A region EM15_10-ME8_4-R1394A-G2132 on chromosome 8 was identified for leaf EW and rate of water loss i.e., time taken to reach 70% RWC from excised leaves in rice lines subjected to drought stress. A large effect QTL (65.6%) was detected on chromosome 2 for harvest index under stress. QTLs identified for EW, rate of water loss from excised leaves and harvest index under stress in this study co-located with QTLs linked to shoot and root-related drought resistance traits in these rice lines and might be useful for rainfed rice improvement.     

Growth,photosynthesis and water relations as affected by different drought regimes and subsequent recovery in <Emphasis Type="Italic">Medicago laciniata</Emphasis> (L.) populations

The aims of this study were to investigate the effects of water deficit and recovery on growth, photosynthesis and water relations in four Medicago laciniata populations from saharian (Ml-90), inferior arid (Ml-204), superior arid (Ml-306) and semi-arid (Ml-376) Tunisian regions. After 28 d of sowing with ample irrigation, the plants were subjected to 4 water regimes: optimal irrigation (100% of field capacity, FC), moderate drought (75% FC), severe drought (35% FC) and rewatering (plants submitted to 35% FC during 7 d, afterwards the plants were rewatered to 100% FC). Harvest was carried out after 28 d of treatments. The drought tolerance in M. laciniata populations was found to be increased particularly with increasing temperatures of collection site of the population. The Ml-204 and Ml-90 populations used mainly physiological strategies for survival under moderate water shortage. Higher severe drought tolerance in both signaled populations would be related to their lower photosynthesis metabolic impairment, relatively higher leaf RWC and greater osmotic potential decrease. The results suggest that plants with low values of leaf features are likely to maintain higher leaf RWC under sever drought. The largest decrease of osmotic potential was found associated with the solute accumulations such as proline and K⁺.