Physiological and Biochemical Responses of Hexaploid and Tetraploid Wheat to Drought Stress

An experiment was conducted to investigate the physiological and biochemical responses of two hexaploids viz., C 306 (water stress tolerant) and Hira (water stress susceptible), and two tetraploids, HW 24 (Triticum dicoccum) and A 9‐30‐1 (Triticum durum) wheat genotypes to water stress under pot culture condition. Water stress was imposed for a uniform period of 10 days at 50, 60 and 70 days after sowing (DAS) and observations were recorded at 60, 70 and 80 DAS. Total dry matter and plant height were recorded at harvest. Water stress caused a decline in relative water content (RWC), chlorophyll and carotenoid content, membrane stability and nitrate reductase activity and increased accumulation of proline at all stages and abscisic acid (ABA) at 80 DAS in all the genotypes. Both the tetraploids showed a lower reduction in RWC and highest ABA accumulation under water stress. Among the hexaploids Hira showed the most decline in RWC and the lowest ABA accumulation. The tetraploids also showed comparatively higher carotenoid content and membrane stability, closely followed by C 306, while Hira showed the minimum response under water stress. Nitrate reductase activity and chlorophyll content under irrigated conditions were highest in Hira but under water stress the lowest per cent decline was observed in C 306, followed by HW 24, A 9‐30‐1, and Hira. Proline accumulation under water stress conditions was highest in hexaploids C 306 and Hira and lowest in tetraploids HW 24 and A 9‐30‐1. Tetraploids HW 24, followed by A 9‐30‐1 maintained higher plant height and total dry matter (TDM) under water stress and also showed a lower per cent decline under stress than hexaploids C 306 and Hira. From the results it is clear that proline accumulation did not contribute to better drought tolerance of tetraploids than hexaploids. It is also apparent that water stress tolerance is the result of the cumulative action of various physiological processes, and all the parameters/processes may not be positively associated with the drought tolerance of a particular tolerant genotype.     

Oxidative Stress and Antioxidants in Wheat Genotypes: Possible Mechanism of Water Stress Tolerance

The role of plant antioxidant systems in water stress tolerance was studied in three contrasting wheat genotypes. Water stress imposed at different stages after anthesis resulted in an increase in lipid peroxidation and a decrease in membrane stability and chlorophyll and carotenoid contents. The antioxidant enzymes ascorbate peroxidase, glutathione reductase and non-specific peroxidase also increased significantly under water stress. Genotype PBW 175, which had highest ascorbate peroxidase, glutathione reductase and peroxidase activity, had the lowest lipid peroxidation (malondialdehyde content) and highest membrane stability and contents of chlorophyll and carotenoids under water stress, while the susceptible genotype WH 542 exhibited the lowest antioxidant enzyme activity, membrane stability and contents of chlorophyll and carotenoids and the highest lipid peroxidation. Genotype HD 2402 showed intermediate behaviour. It seems that drought tolerance of PBW 175, as represented by higher membrane stability and chlorophyll and carotenoid contents and lower lipid peroxidation, is related to its higher antioxidant enzyme activity.     

Tolerance of Drought and Temperature Stress in Relation to Increased Antioxidant Enzyme Activity in Wheat

An experiment was conducted with three wheat genotypes differing in their sensitivity to moisture and/or temperature stress to study the relationship of the chloroplast antioxidant system to stress tolerance. Both moisture stress and temperature stress increased glutathione reductase and peroxidase and decreased membrane stab-iltty, chlorophyll content and chlorophyll stability index in all genotypes. Under moisture stress. DL 153–2 showed the highest membrane stabihty index, chlorophyll content, chlorophyll stability index, glutathione reductase activity and peroxidase activity. However, under elevated temperature conditions, HD 2285, and to a lesser extent DL 153–2, showed higher membrane stability, chlorophyll content and chlorophyll stability index and activities of glutathione reductase and peroxi-dase. Raj 3077, which is sensitive to both drought and temperature stress, showed the lowest membrane stability, chlorophyll content and chlorophyll stability index and glutathione reductase and perosidase activity under elevated temperature as well as drought conditions. Thus, it can be concluded that tolerance of the genotype to moisture and/or temperature stress is closely associated with its antioxidant enzyme system.     

Effect of Post-Anthesis Water Stress on Accumulation of Dry Matter, Carbon and Nitrogen and Their Partitioning in Wheat Varieties Differing in Drought Tolerance

In four wheat ( Triticum aestivum L.) cultivars of tall (C306 and Narmada) and dwarf (HD2329 and Kundan) type, post-anthesis water stress affected the dry matter accumulation in plant parts with respect to main shoot controls. HD2329 among the dwarf types and Narmada 112 among the tall types were more adversely affected by stress, with greater reductions in their biomass and grain yield. Of the tolerant types, C306 (tall) showed a marginal reduction while Kundan (dwarf) had no reduction in these parameters as a result of stress. The results also indicated a varietal response to carbon and nitrogen accumulation and their partitioning in the main shoot when subjected to post-anthesis stress. In the susceptible dwarf cultivar HD2329, and in the susceptible tall cultivar Narmada 112, carbon and nitrogen contents were reduced in the grains of stressed main shoots. Of the tolerant cultivars, the dwarf type Kundan was not affected by stress, while the tall type C306 registered an increase in carbon content and its partitioning to grain.     

Genotypic Variation in Wheat in Response to Water Stress and Abscisic Acid-Induced Accumulation of Osmolytes in Developing Grains

Two wheat genotypes differing in water stress sensitivity (C306, relatively tolerant; HD2329, relatively susceptible) were water stressed during early grain filling by withholding water for 7 days at watery‐ripe (WR) stage and examined for water status, abscisic acid (ABA) and osmolytes in grains as well as in flag leaf (FL). Both the genotypes differed significantly from each other in endogenous levels of ABA, proline, glycine betaine, total soluble sugars, reducing sugars, sucrose and potassium. The tolerant genotype showed higher content of ABA, proline, glycine betaine, total sugars, reducing sugars and had higher water content in its FL and grains than the susceptible genotype, which contained more of glycine beatine and potassium but had lower ABA and water content in its FL and grains. Although carbohydrates constituted the major amount of all the solutes, proline and glycine betaine increased manifold during stress. Exogenous application of 2 μm ABA at 5 days after anthesis to FL of stressed plants increased the endogenous content of ABA, accelerated the accumulation of osmolytes, improved the water status of FL and grains that resulted in higher grain weight, especially in the susceptible genotype. Differential response of contrasting wheat genotypes to water stress may be governed by ABA‐dependent solute accumulation in grains and FL.     

白刺对干旱胁迫的生理生化反应

研究白刺幼苗抗旱特性的生理学机制,期望为干旱农业生产提供理论依据。实验采用盆栽,用控制浇水的方法对不同强度的干旱胁迫梯度上对白刺幼苗叶片相对含水量、丙二醛、脯氨酸、可溶性蛋白质、叶绿素含量、超氧离子自由基产生速率和抗氧化酶活性的变化特点进行了研究。试验结果表明：在干旱胁迫下,白刺叶片相对含水量降低、脯氨酸含量呈增加趋势;可溶性蛋白质、叶绿素含量在轻度和中度干旱胁迫时上升,在重度干旱胁迫时显著下降;随着干旱胁迫的加剧,膜透性上升,丙二醛含量和超氧离子自由基产生速率增加。抗氧化系统酶活性测定结果表明,SOD、POD、CAT 3个酶能配合协同作用,降低膜脂过氧化程度,减少水分胁迫造成的伤害。     

Differential Responses of Antioxidative Defence System to Long‐Term Field Drought in Wheat (Triticum aestivum L.) Genotypes Differing in Drought Tolerance

Drought is a severe abiotic stress and the major constraint on wheat (Triticum aestivum L.) productivity world wide. Deciphering the mechanisms of drought tolerance is a challenging task because of the complexity of drought responses, environmental factors and their interactions. The objective of this study was to evaluate the ability of the antioxidative defence system in imparting tolerance against drought‐induced oxidative stress and yield loss in two wheat genotypes, when subjected to long‐term field drought. Drought resulted in an increase in H₂O₂ accumulation and lipid peroxidation and decrease in ascorbate level in roots and leaves at different plant developmental stages. Drought‐tolerant genotype having higher antioxidative enzymes activities, and ascorbate level was superior to that of sensitive genotype in maintaining lower H₂O₂ content and lipid peroxidation and higher growth, yield and yield components under water deficit. Various antioxidative enzymes showed positive correlation with ascorbate and negative with H₂O₂ content. In developing grains, antioxidative defence response was nearly similar among both the genotypes under control condition; however, sensitive genotype failed to modulate the activities of antioxidative enzymes according to the ROS rush under field drought. Poor capacity of the antioxidative defence system in vegetative and reproductive tissues of sensitive genotype seems to be responsible, at least partly, for reduced yield potential under water deficit.     

Improved salt tolerance and seed cotton yield in cotton (<Emphasis Type="Italic">Gossypium hirsutum</Emphasis> L.) by transformation with <Emphasis Type="Italic">betA</Emphasis> gene for glycinebetaine synthesis

Homozygous transgenic cotton (Gossypium hirsutum L.) plants that accumulated glycinebetaine (GB) in larger quantities were more tolerant to salt than wild-type (WT) plants. Four transgenic lines, namely 1, 3, 4, and 5, accumulated significantly higher levels of GB than WT plants did both before and after salt stress. At 175 and 275 mM NaCl, seeds of all the transgenic lines germinated earlier and recorded a higher final germination percentage, and the seedlings grew better, than those of the WT. Under salt stress, all the lines showed some characteristic features of salt tolerance, such as higher leaf relative water content (RWC), higher photosynthesis, better osmotic adjustment (OA), lower percentage of ion leakage, and lower peroxidation of the lipid membrane. Levels of endogenous GB in the transgenic plants were positively correlated with RWC and OA. The results indicate that GB in transgenic cotton plants not only maintains the integrity of cell membranes but also alleviates osmotic stress caused by high salinity. Lastly, the seed cotton yield of transgenic lines 4 and 5 was significantly higher than that of WT plants in saline soil. This research indicates that betA gene has the potential to improve crop’s salt tolerance in areas where salinity is limiting factors for agricultural productivity.     

Methyl Jasmonate‐Induced Alteration in Lipid Peroxidation,Antioxidative Defence System and Yield in Soybean Under Drought

Methyl jasmonate (MeJA), a plant‐signalling molecule, is involved in an array of plant development and the defence responses. This study was conducted to explore the role of exogenous MeJA application in alleviating the adversities of drought stress in soybean (Glycine max L. Merrill.). Soybean plants were grown under normal conditions until blooming and were then subjected to drought by withholding irrigation followed by foliar application of (50 μm ) MeJA. Drought stress substantially suppressed the yield and yield‐related traits, whereas it accelerated the membrane lipid peroxidation. Nonetheless, substantial increase in activities of enzymatic antioxidants (superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT)), proline, relative water contents (RWC) with simultaneous decrease in membrane lipid peroxidation was observed in MeJA‐treated plants under drought. These beneficial effects led to improvement in biological and grain yield, and harvest index under drought. Interestingly, MeJA application was also useful under well‐watered conditions. These results suggest the involvement of MeJA in improving the drought tolerance of soybean by modulating the membrane lipid peroxidation and antioxidant activities.     

外源一氧化氮对盐胁迫下烟草幼苗生理及抗氧化性的影响

为探究外源一氧化氮（NO）对盐胁迫下烟草幼苗的影响,为了解烟草抗盐机制和生产应用提供一定参考,以烟草品种K326为研究对象,使用硝普钠（sodium nitroprusside,SNP）作为外源一氧化氮供体,研究不同处理烟草幼苗的生理及抗氧化性。试验结果表明,盐胁迫会显著抑制幼苗的正常生长发育,通过使用不同浓度的SNP预处理后,烟草幼苗叶片含水率明显上升,根系活力上升,提高了胁迫环境下的叶绿素含量,渗透物质含量显著增加,幼苗体内的多种抗氧化酶含量显著升高,过氧化氢含量和MDA含量下降,降低了膜脂过氧化的程度,保护细胞结构不被破坏,降低叶片细胞透性。试验结果说明,外源一氧化氮可以有效提高烟草幼苗的耐盐性,并且具有明显的浓度效应,0.10mmol/L SNP预处理的效果最好,试验结果为提高烟草的耐盐性提供一定依据。     

Effect of Water Stress on Physiological Attributes and their Relationship with Growth and Yield of Wheat Cultivars at Different Stages

The effects of water stress on physiological attributes of drought‐sensitive (Kalyansona) and drought‐tolerant (C‐306) wheat cultivars were studied in a pot experiment. Water stress was imposed by withholding irrigation at boot and anthesis stages. Leaf water potential, leaf osmotic potential and leaf turgor potential (measured with pressure chamber and osmometer), as well as leaf diffusive resistance, leaf transpiration rate and leaf‐to‐air transpiration gradient (measured with a steady‐state porometer) were measured diurnally. Growth and yield parameters were recorded after harvesting of the crop. Triplicate data were analysed using a completely randomized design and correlations amongst these parameters were computed. Water stress was found to reduce diurnal leaf water potential and leaf osmotic potential in both the genotypes but leaf osmotic potential was significantly higher in the drought‐tolerant cultivar C‐306 than in the drought‐sensitive cultivar Kalyansona. Positive turgor was recorded in both the genotypes under water stress and non‐stress conditions. Water‐stressed plants showed significantly lower turgor potential than control plants. In diurnal observations, water‐stressed plants exhibited significantly higher leaf diffusive resistance in both genotypes at both stages. The diffusive resistance of C‐306 was predominantly higher than that of Kalyansona. Water stress decreased leaf transpiration rate at both stages but the reduction was higher at the anthesis stage. The leaf‐to‐air temperature gradient was much higher in C‐306 than in Kalyansona at the boot stage but at the anthesis stage genotypic variation was non‐significant. The capacity to maintain cooler foliage was lower at the anthesis stage than at the boot stage in both the cultivars. Shoot dry weight, number of grains, test weight, grain yield, biological yield and harvest index decreased to a greater extent when water stress was imposed at the anthesis stage, while imposition of water stress at the boot stage caused a greater reduction in plant height and number of tillers. Similarly, water stress caused a smaller reduction in growth, yield and yield attributes in C‐306 than in Kalyansona. In general, the correlation coefficient of grain and biological yield with water potential and its components was positive and highly significant. Similarly, turgor potential was also correlated positively and significantly with grain yield at both the stages, but with biological yield it was significant only at the anthesis stage. A negative and significant correlation was obtained for diffusive resistance and leaf‐to‐air temperature gradient with grain yield at the boot and anthesis stages. The rate of transpiration was also positively and significantly correlated to grain and biological yields at both the stages. Amongst the yield attributes, number of leaves and number of tillers were positively correlated at the anthesis stage, whereas leaf area and shoot dry weight were significantly correlated with grain and biological yields at both the stages.     

Functional mechanisms of drought tolerance in maize through phenotyping and genotyping under well watered and water stressed conditions

Developing tolerant genotypes is crucial for stabilizing maize productivity under drought stress conditions as it is one of the most important abiotic stresses affecting crop yields. Twenty seven genotypes of maize (Zea mays L.) were evaluated for drought tolerance for three seasons under well watered and water stressed conditions to identify interactions amongst various tolerance traits and grain yield as well as their association with SSR markers. The study revealed considerable genetic diversity and significant variations for genotypes, environment and genotype × environment interactions for all the traits. The ranking of genotypes based on drought susceptibility index for morpho-physiological traits was similar to that based on grain yield and principal component analysis. Analysis of trait – trait and trait – yield associations indicated significant positive correlations amongst the water relations traits of relative water content (RWC), leaf water potential and osmotic potential as well as of RWC with grain yield under water stressed condition. Molecular analysis using 40 SSRs revealed 32 as polymorphic and 62 unique alleles were detected across 27 genotypes. Cluster analysis resulted in categorization of the genotypes into five distinct groups which was similar to that using principal component analysis. Based on overall performance across seasons tolerant and susceptible genotypes were identified for eventual utilization in breeding programs as well as for QTL identification. The marker-trait association analysis revealed significant associations between few SSR markers with water relations as well as yield contributing traits under water stressed conditions. These associations highlight the importance of functional mechanisms of intrinsic tolerance and cumulative traits for drought tolerance in maize.     

外源氯化钙对盐胁迫下西兰花抗氧化酶系统及离子吸收的影响

摘要：以美国“格福”与日本“山水”两种耐盐性不同的西兰花为试验材料,采用基质营养液栽培,研究了外源氯化钙对盐胁迫下西兰花鲜重、干重、MDA、抗氧化酶活性及离子吸收的影响。结果表明：外源钙的添加,可显著促进西兰花的生长,减轻盐胁迫,增加地上部、地下部干重及鲜重。外源钙处理明显降低了西兰花叶片MDA含量,减轻盐胁迫对叶片膜脂过氧化程度。外源钙处理对西兰花SOD、POD、CAT活性的影响因品种耐盐性不同表现不一。外源钙处理可减少西兰花地上部、地下部Na+的积累,增加K+、Ca2+　的浓度。外源钙处理对盐敏感品种的效果要好于耐盐品种。最佳的钙浓度为10mmol/L。     

Genotypic variability in plant water status of French bean under drought stress

Seven genotypes of French bean (Phaseolus vulgaris L.) were evaluated under semi-controlled conditions at the Bangabandhu Sheikh Mujibur Rahman Agricultural University, Bangladesh to analyze genotypic variability in leaf water status under water stress. The plants were grown under two moisture regimes, viz. 80% field capacity (FC) and 50% FC throughout the growing season. The genotypes showed significant variation in water relation traits. Genotypes BB24 and BB43 maintained higher relative water content (RWC), but lower turgid weight/dry weight ratio (TW/DW) and water uptake capacity (WUC). When drought susceptibility index (DSI) among the genotypes was considered, BB24 was found the most tolerant to drought and BB04 was the most susceptible one. A close positive relationship between leaf TW/DW and DSI under drought was recorded (R ² = 0.627). Leaf TW/DW was decreased considerably due to water stress by 10% in genotype BB24 followed by BB43 (13%), and both BARI bushbean-2 and BB04 (19%). Stomatal aperture and whole plant transpiration rate were found minimal in the BB24 and BB43 compared to that of BB04 and BARI bushbean-2. Considering these water relation traits, genotypes BB24 and BB43 may be considered as relatively tolerant to tissue dehydration. The study also revealed that the TW/DW, WUC, stomatal aperture, and whole plant transpiration rate was negatively and significantly associated with yield; however, the RWC was positively correlated with yield under water stress conditions.     

Inter‐relationship between photosynthetic efficiency, Δ13C,antioxidant activity and sugarcane yield under drought stress in field conditions

Drought is one of the major climatic factors that reduce crop yields in cultivated areas around the world, and studies on physiological responses may help in the selection of drought tolerant genotypes. Thus, this work aimed to correlate gas exchange, photosynthetic efficiency, carbon isotope discrimination (Δ¹³C) and antioxidant activity in sugarcane varieties submitted to water stress under field conditions. Six sugarcane varieties were submitted to drought stress in three development stages: tillering, intense growth and ripening. In all varieties, the photosynthetic apparatus was severely affected by drought, with a reduction in photosynthetic rate and chlorophyll content. During the tillering stage, reductions in gas exchange and increase in Δ¹³C and bundle sheath leakiness (φ) were observed. In the intense growth stage, water stress caused increases in leaf temperature, intrinsic water use efficiency, antioxidant enzyme activity and lipid peroxidation, and reductions in stomatal conductance and transpiration in the RB72454, RB855113 and RB855536 varieties. However, the RB92579 variety maintained a better physiological homeostasis at all development stages and presented higher stalks yielding when submitted to drought. This work suggests that screen sugarcane genotype to drought stress should be performed during the intense growth stage, when plants are more sensitive to drought conditions.     

硅对盐胁迫下黄瓜叶片膜脂过氧化和活性氧清除系统的影响

研究了水培条件下硅对盐胁迫时黄瓜幼苗膜脂过氧化和活性氧清除系统的影响。结果表明,加硅使盐胁迫下黄瓜幼苗叶片保护酶(SOD,AsA-POD,G-POD)的活性显著升高,抗坏血酸(AsA)和谷胱甘肽(GSH)的含量明显增加,黄瓜叶片的丙二醛(MDA)的含量显著下降。加硅可显著提高盐胁迫条件下黄瓜叶片相对含水量,降低叶片电解质渗漏率。因此,加硅可减轻盐胁迫对黄瓜幼苗的伤害,硅参与了植物的代谢或生理活动。     

春季水分胁迫对佛甲草生长的影响

为调查佛甲草对水分适应能力,采用盆栽试验方法,研究了水分胁迫及复水对佛甲草植株的生长状况、组织的相对含水量（RWC）、叶绿素含量、相对质膜透性、脯氨酸含量、根系活力和丙二醛等生理生化指标的变化。研究结果表明,在水分胁迫下,佛甲草植株生长受到一定程度抑制;在严重水分胁迫下组织相对含水量明显下降;叶绿素含量呈先升后降的趋势;细胞膜透性则表现为持续升高;受水分胁迫的影响,脯氨酸含量及丙二醛含量随水分胁迫强度的增强而增加,随胁迫时间的延长而降低,受胁迫越重,含量越高。复水后,生长情况得到明显改善,各种生理生化指标逐渐恢复到正常状态。     

干旱胁迫对蒲公英渗透调节物质、酶保护系统及质膜水孔蛋白PIP2-3基因表达的影响

为了揭示蒲公英在干旱胁迫下渗透调节和酶保护系统的作用机理以及质膜水孔蛋白PIP2-3基因的表达特征,采用盆栽控水试验,研究了不同干旱胁迫RWC=(90±5)%、RWC=(75±5)%、RWC=(50±5)%下蒲公英渗透调节物质、酶保护系统以及质膜水孔蛋白基因的相关表达。结果表明:随着干旱胁迫的加重,蒲公英叶片中的可溶性糖、可溶性蛋白、游离脯氨酸的含量显著增加,超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)的活性显著提高,在轻度胁迫(MS)中呈持续上升的趋势,重度胁迫(SS)中则呈现先升高后下降的趋势(可溶性糖除外);质膜水孔蛋白PIP2-3基因的相对表达量上调,轻度胁迫(MS)显著高于重度胁迫(SS)。复水7 d后,渗透调节物质的积累以及SOD的含量下降,但仍显著高于CK,POD、CAT的含量以及质膜水孔蛋白PIP2-3基因的相对表达量恢复正常,说明在干旱胁迫条件下蒲公英通过增加渗透调节物质的积累,提高酶保护系统的活性来增强植株的抗逆性,上调质膜水孔蛋白基因的相对表达量来加强水分运输能力。     

离体小麦叶片干旱胁迫过程中水分生理性状的变化

１９９０,１９９１和１９９３三个年度应用不同基因型,供水,密度和施氮水平在不同生育期对小麦叶片防体失水过程中叶水势,自由水和束缚水量含量及相对电导率等水分生理性状指标变化进行了同步测定分析,研究发现,随离体叶片水分胁迫增强,离体叶片的偿速率的变化与其它水分生理性状的变化基本一致,研究还表明,在失水过程中束缚水含量增加,且基因型间存在差异,可能影响离体叶片失水过程。     

Interactive Effects of Sudden and Gradual Drought Stress and Foliar‐applied Glycinebetaine on Growth,Water Relations,Osmolyte Accumulation and Antioxidant Defence System in Two Maize Cultivars Differing in Drought Tolerance

Influence of sudden and gradual drought stress (DS) and foliar‐applied glycinebetaine (GB) on growth, water relations, osmolyte accumulation and antioxidant defence system were investigated in the plants of two maize (Zea mays L.) cultivars, that is, drought‐tolerant Shaandan 9 (S₉) and drought‐sensitive Shaandan 911 (S₉₁₁). Sudden DS caused less accumulation of GB and free proline, but a more accumulation of malondialdehyde (MDA), which resulted in a greater reduction in leaf relative water content (RWC) and dry matter (DM) in both cultivars compared with the gradual DS. Exogenous GB application caused a rise in DM, RWC, contents of GB and free proline as well as the activities of SOD, CAT and POD along with a decline in MDA content to various extent in both cultivars under both types of DS. A more pronounced effectiveness of GB application was observed in S₉₁₁ than that in S₉ under the same type of DS. It seemed that the more serious damage of DS was on maize plants, and the better positive role of GB was observed in terms of mitigating the adverse effects of DS. From this study, it was possible to propose that hardening for drought resistance by gradual DS treatment and GB application are effective to make plants robust to thrive under water‐deficit conditions.