Plant Morphological and Biochemical Responses to Field Water Deficits : II. Responses of Leaf Glycerolipid Composition in Cotton

The effects of water deficits on leaf glycerolipid composition were analyzed in two photoperiodic strains of field grown cotton (Gossypium kirsutum L.) that differ in sensitivity to drought. Leaves from plants grown under dryland conditions exhibited increased dry weight and specific leaf weight. The average midday leaf water potential in the dryland treatment decreased to −1.9 and −2.4 megapascals, respectively, for the T25 and T185 genotypes. Total leaf lipid content of plants exposed to dryland conditions was 5.9 and 7.5% of leaf dry weight for strain T25 and T185, respectively. The difference in leaf lipid content between these genotypes was caused by water deficits and was attributed to loss of both phospholipids and glycolipids in strain T25. There was no apparent loss of phospholipids due to water deficits in the T185 genotype; however, a significant loss of glycolipids was partially compensated by a 2-fold increase in triacylglycerol. No change in triacylglycerol was found between treatments in T25 leaves. Water deficit caused a significant decline in the relative degree of acylunsaturation in phospholipids and glycolipids from both genotypes; however, the double bond index for triacylglycerol increased in both genotypes. It is believed that the observed responses of leaf lipid composition to dryland conditions may be an additional criterion for characterization and selection of new drought-tolerant cotton genotypes.     

Integrated drought responses of black poplar: how important is phenotypic plasticity?

Climate change is expected to increase drought frequency and intensity which will threaten plant growth and survival. In such fluctuating environments, perennial plants respond with hydraulic and biomass adjustments, resulting in either tolerant or avoidant strategies. Plants' response to stress relies on their phenotypic plasticity. The goal of this study was to explore physiology of young Populus nigra in the context of a time‐limited and progressive water deficit in regard to their growth and stress response strategies. Fourteen French 1‐year‐old black poplar genotypes, geographically contrasted, were subjected to withholding water during 8 days until severe water stress. Water fluxes (i.e. leaf water potentials and stomatal conductance) were analyzed together with growth (i.e. radial and longitudinal branch growth, leaf senescence and leaf production). Phenotypic plasticity was calculated for each trait and response strategies to drought were deciphered for each genotype. Black poplar genotypes permanently were dealing with a continuum of adjusted water fluxes and growth between two extreme strategies, tolerance and avoidance. Branch growth, leaf number and leaf hydraulic potential traits had contrasted plasticities, allowing genotype characterization. The most tolerant genotype to water deficit, which maintained growth, had the lowest global phenotypic plasticity. Conversely, the most sensitive and avoidant genotype ceased growth until the season's end, had the highest plasticity level. All the remaining black poplar genotypes were close to avoidance with average levels of traits plasticity. These results underpinned the role of plasticity in black poplar response to drought and calls for its wider use into research on plants' responses to stress.     

Comparison of the Drought Stress Responses of Tolerant and Sensitive Wheat Cultivars During Grain Filling: Changes in Flag Leaf Photosynthetic Activity,ABA Levels,and Grain Yield

Water status parameters, flag leaf photosynthetic activity, abscisic acid (ABA) levels, grain yield, and storage protein contents were investigated in two drought-tolerant (Triticum aestivum L. cv. MV Emese and cv. Plainsman V) and two drought-sensitive (cvs. GK élet and Cappelle Desprez) wheat genotypes subjected to soil water deficit during grain filling to characterize physiological traits related to yield. The leaf water potential decreased earlier and at a higher rate in the sensitive than in the tolerant cultivars. The net CO₂ assimilation rate (P _N) in flag leaves during water deficit did not display a strict correlation with the drought sensitivity of the genotypes. The photosynthetic activity terminated earliest in the tolerant cv. Emese, and the senescence of flag leaves lasted 7 days longer in the sensitive Cappelle Desprez. Soil drought did not induce characteristic differences between sensitive and tolerant cultivars in chlorophyll a fluorescence parameters of flag leaves during post-anthesis. Changes in the effective quantum yield of PSII (Φ_PSII) and the photochemical quenching (qP) depended on the genotypes and not on the sensitivity of cultivars. In contrast, the levels of ABA in the kernels displayed typical fluctuations in the tolerant and in the sensitive cultivars. Tolerant genotypes exhibited an early maximum in the grain ABA content during drought and the sensitive cultivars maintained high ABA levels in the later stages of grain filling. In contrast with other genotypes, the grain number per ear did not decrease in Plainsman and the gliadin/glutenin ratio was higher than in the control in Emese during drought stress. A possible causal relationship between high ABA levels in the kernels during late stages of grain filling and a decreased grain yield was found in the sensitive cultivars during drought stress.     

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.     

Assessing the drought tolerance variability in Mediterranean alfalfa (Medicago sativa L.) genotypes under arid conditions

This work investigated the variability in drought tolerance under arid conditions of Mediterranean alfalfa genotypes with the overall aim to assess the main criteria that are associated with the relative tolerance and to discover the most tolerant ecotypes. For this, 16 alfalfa genotypes originating from seven countries of the Mediterranean basin were tested in an experimental station in south of Tunisia. The trial was conducted under two irrigation treatments. The first was normally irrigated by providing an amount of water corresponding to the potential evapotranspiration of the crop, and in the second with water deficit which was applied by stopping the irrigation during 8 weeks in summer. A significant decrease was observed under water deficit for biomass production, leaf stem ratio and water use efficiency. The sensitive index, in stress treatment, varies between 13.8% and 46.2% for dry yield. Results showed that some genotypes exhibited high-forage yield potential even in the presence of stress, mainly Amerist, Sardi10 and Siriver. Proline accumulation in leaves was greater in water-stressed plants, while the K⁺ osmo-regulatory role was not definite. High biomass production, accumulation of proline and constancy of K⁺ in leaves are the most important criteria for tolerant alfalfa.     

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.     

Detrimental effect of rust infection on the water relations of bean

Bean plants (Phaseolus vulgaris L.) infected with the rust Uromyces phaseoli became unusually susceptible to drought as sporulation occurred. Under the conditions used (1,300 ft-c, 27 C, and 55% relative humidity) such plants wilted at soil water potentials greater than −1 bar, whereas healthy plants did not wilt until the soil water potential fell below −3.4 bars. Determinations of leaf water and osmotic potentials showed that an alteration in leaf osmotic potential was not responsible for the wilting of diseased plants. When diffusive resistance was measured as a function of decreasing leaf water content, the resistance of healthy leaves increased to 50 sec cm⁻¹ by the time relative water content decreased to 70%, whereas the resistance of diseased leaves remained less than 8 sec cm⁻¹ down to 50% relative water content. Apparently, water vapor loss through cuticle damaged by the sporulation process, together with the reduction in root to shoot ratio which occurs in diseased plants, upset the water economy of the diseased plant under mild drought conditions.     

Impact of drought stress on growth and quality of miscanthus for biofuel production

Miscanthus has a high potential as a biomass feedstock for biofuel production. Drought tolerance is an important breeding goal in miscanthus as water deficit is a common abiotic stress and crop irrigation is in most cases uneconomical. Drought may not only severely reduce biomass yields, but also affect biomass quality for biofuel production as cell wall remodeling is a common plant response to abiotic stresses. The quality and plant weight of 50 diverse miscanthus genotypes were evaluated under control and drought conditions (28 days no water) in a glasshouse experiment. Overall, drought treatment decreased plant weight by 45%. Drought tolerance – as defined by maintenance of plant weight – varied extensively among the tested miscanthus genotypes and ranged from 30% to 110%. Biomass composition was drastically altered due to drought stress, with large reductions in cell wall and cellulose content and a substantial increase in hemicellulosic polysaccharides. Stress had only a small effect on lignin content. Cell wall structural rigidity was also affected by drought conditions; substantially higher cellulose conversion rates were observed upon enzymatic saccharification of drought‐treated samples with respect to controls. Both cell wall composition and the extent of cell wall plasticity under drought varied extensively among all genotypes, but only weak correlations were found with the level of drought tolerance, suggesting their independent genetic control. High drought tolerance and biomass quality can thus potentially be advanced simultaneously. The extensive genotypic variation found for most traits in the evaluated miscanthus germplasm provides ample scope for breeding of drought‐tolerant varieties that are able to produce substantial yields of high‐quality biomass under water deficit conditions. The higher degradability of drought‐treated samples makes miscanthus an interesting crop for the production of second‐generation biofuels in marginal soils.     

Improving water use efficiency by changing hydraulic and stomatal characteristics in soybean exposed to drought: the involvement of nitric oxide

A variety of cellular responses is needed to ensure the plants survival during drought, but little is known about the signaling mechanisms involved in this process. Soybean cultivars (EMBRAPA 48 and BR 16, tolerant and sensitive to drought, respectively) were exposed to the following treatments: control conditions (plants in field capacity), drought (20% of available water in the soil), sodium nitroprusside (SNP) treatment (plants irrigated and treated with 100-µM SNP [SNP–nitric oxide (NO) donor molecule], and Drought + SNP (plants subjected to drought and SNP treatment). Plants remained in these conditions until the reproductive stage and were evaluated for physiological (photosynthetic pigments, chlorophyll a fluorescence and gas exchange rates), hydraulic (water potential, osmotic potential and leaf hydraulic conductivity) and morpho-anatomical traits (biomass, venation density and stomatal characterization). Exposure to water deficit considerably reduced water potential in both cultivars and resulted in decrease in photosynthesis and biomass accumulation. The addition of the NO donor attenuated these damaging effects of water deficit and increased the tolerance index of both cultivars. The results showed that NO was able to reduce plant's water loss, while maintaining their biomass production through alteration in stomatal characteristics, hydraulic conductivity and the biomass distribution pattern. These hydraulic and morpho-anatomical alterations allowed the plants to obtain, transport and lose less water to the atmosphere, even in water deficit conditions.     

Protective Mechanism Triggered by Pigeonpea Plants Exposed to Water Deficit: Modifications Linked to Paraheliotropism,Stomatal Characteristics and Antioxidant Enzymes

Drought is the main challenge in modern agriculture. The development and utilization of genotypes with a protective mechanism against drought may be crucial for mitigating the impacts on pigeonpea metabolism resulting from water deficit. This study aimed to determine if leaf characteristics and antioxidant enzymes have protective roles and possible benefits to growth by comparing the physiological, anatomical, biochemical and morphological responses of two different pigeonpea genotypes under water deficit. The experiment was randomized with four treatments, two genotypes (FL and I43, sensitive and tolerant to drought, respectively) and water conditions (control and water deficit). Water deficit was induced by suspension of irrigation for six days. Water deficit promoted negative interferences in both genotypes, but I43 genotype had a protective mechanism, reducing drought impacts. Leaf inclination (LI), elevations of 337% and 373%, was detected in the FL and I43 genotypes, respectively. Stomatal characteristics, FL and I43 genotypes, had reductions of 22% and 10%, respectively, in stomatal density (SD) on the abaxial face in comparison with the control. However, for stomatal functionality (SF) on the abaxial face, the FL and I43 genotypes had contrasting behaviours with a decrease and an increase of 7% and 3%, respectively, compared to the control. Antioxidant enzymes, FL and I43 genotypes, had increased superoxide dismutase (SOD) activities of 71% and 154%, respectively. For catalase (CAT), the water deficit resulted in increases of 55% and 415% for the genotypes, respectively. The tolerant genotype (I43) increased LI (paraheliotropism), minimizing the incidence of solar radiation in the leaves and water losses by transpiration. Fewer negative effects on stomatal characteristics (SD and SF) and the consequent maintenance of gas exchange (net photosynthetic rate and transpiration rate) guaranteed the influx of carbon dioxide in the leaf and mitigated the effects of water deficit on I43. In addition, the tolerant genotype (I43) had lower oxidative stress during the water deficit resulting from the greatly increased activities of antioxidant enzymes (SOD, CAT and APX). Our results clearly revealed that biochemical, physiological and anatomical modifications are intrinsically connected with a protective mechanism, mitigating the impacts caused by water limitation on growth of pigeonpea plants.

     

Are ABA,ethylene or their interaction involved in the response of leaf growth to soil water deficit? An analysis using naturally occurring variation or genetic transformation of ABA production in maize

The role of abscisic acid (ABA) and its possible interaction with ethylene in mediating leaf elongation response to soil water deficit are a matter of controversy. To address this question, we used a set of maize genotypes with various levels of ABA either due to natural variability or to genetic transformation targeted on NCED/VP14, a key enzyme of ABA synthesis. The transgenic lines yielded less strong phenotypes than available mutants, making it possible to use them under normal growing conditions. We focused on leaf elongation during night periods in order to avoid the confounding effect of ABA on leaf water status. Our results suggest that over a wide range, internal ABA level (measured in both leaf extracts or xylem sap) has no clear effect on leaf elongation response to soil water deficit, except in the case of an antisense line presenting the strongest reduction in ABA accumulation that showed a slight maintenance of leaf elongation during water deficit. Leaf ethylene production rate was variable and not related to water deficit except in the ABA-deficient transgenic lines where it was increased by water deficit on average but not systematically. Moreover, variability in ethylene production rate was not linked to variability in elongation rate. Our results thus suggest that neither ABA nor ethylene seems to play a major role in the control of leaf elongation response to soil water deficit.     

Responses of potato genotypes to drought. II. Leaf area index, growth and yield

Canopy expansion, growth and yield were examined in 19 genotypes of potato (Solanurn tuberosum L.) grown either with irrigation or droughted from the time of plant emergence. In the irrigated treatment, genotypes differed both in the maximum leaf area index (L_ai) achieved and in the duration the canopy was maintained. Drought reduced both the rate of canopy expansion and the maximum L_ai achieved. In the droughted treatment, biomass production was correlated with leaf area duration. The relation between the effect of drought on the growth of individual leaves and biomass or yield is examined. It is concluded that selection for early leaf appearance combined with the ability to sustain leaf growth with increasing soil moisture deficit (SMD) would improve productivity in the presence of drought.     

Stomatal and nonstomatal regulation of water use in cotton, corn, and sorghum

Stomata of corn (Zea mays L.) and sorghum (Sorghum bicolor L.) responded to changes in leaf water potential during the vegetative growth phase. During reproductive growth, leaf resistances were minimal and stomata were no longer sensitive to bulk leaf water status even when leaf water potentials approached −27 bars. Stomata of corn, cotton (Gossypium hirsutum L.), and sorghum appear to respond to changes in the humidity deficit between the leaf and air and in this manner, regulated transpirational flux to some degree. Distinct differences in water transport efficiency were observed in the three species. Under nonlimiting soil water conditions, sorghum exhibited the greatest efficiency of water transport while under limiting soil moisture conditions, cotton appeared most efficient. Corn was the least efficient with respect to nonstomatal regulation of water use. Differences in drought tolerance among the three species are partially dependent on stomatal regulation of water loss, but efficiency of the water transport system may be more related to drought adaptation. This is particularly important since stomata of all three species did not respond to bulk leaf water status during a large portion of the growing season.     

Impact of drought on productivity and water use efficiency in 29 genotypes of Populus deltoides x Populus nigra

We examined the relationships among productivity, water use efficiency (WUE) and drought tolerance in 29 genotypes of Populus x euramericana (Populus deltoides x Populus nigra), and investigated whether some leaf traits could be used as predictors for productivity, WUE and drought tolerance. At Orléans, France, drought was induced on one field plot by withholding water, while a second plot remained irrigated and was used as a control. Recorded variables included stem traits (e.g. biomass) and leaf structural (e.g. leaf area) and functional traits [e.g. intrinsic water use efficiency (Wi) and carbon isotope discrimination (Delta)]. Productivity and Delta displayed large genotypic variability and were not correlated. Delta scaled negatively with Wi and positively with stomatal conductance under moderate drought, suggesting that the diversity for Delta was mainly driven by stomatal conductance. Most of the productive genotypes displayed a low level of drought tolerance (i.e. a large reduction of biomass), while the less productive genotypes presented a large range of drought tolerance. The ability to increase WUE in response to water deficit was necessary but not sufficient to explain the genotypic diversity of drought tolerance.     

夏玉米对土壤水分持续减少的响应及其转折点阈值分析

玉米是世界三大粮食作物之一,玉米生产在中国粮食安全与畜牧业发展中具有举足轻重的作用。干旱是夏玉米生产最主要气象灾害,及时准确地获取干旱信息对夏玉米安全生产至关重要。以夏玉米郑单958品种为材料,设置充分供水和拔节期开始土壤水分持续减少两种水分处理,研究夏玉米对土壤水分持续减少的响应及其转折点阈值,为夏玉米干旱识别与监测提供依据。结果表明,土壤水分持续减少10d后生理指标开始陆续受到胁迫,20d后生物量积累受到抑制,30d左右形态特征开始受到胁迫。夏玉米生理指标中最先受到胁迫的是顶端第1片完全展开叶的含水量和水势,生物量积累指标中为茎生物量,形态指标中为叶数。夏玉米顶端第1片完全展开叶的含水量或水势、茎生物量和叶数开始受到土壤干旱过程胁迫的时间阈值分别为11、21、27d,水分亏缺阈值分别为34、66、86mm,土壤相对湿度阈值分别为64%、56%和52%。表明夏玉米对土壤干旱过程的响应首先表现为生理特征变化、其次为生物量积累变化、最后为形态特征变化。研究结果可为客观辨识夏玉米干旱的发生发展及监测预警提供参考。     

Photosynthetic parameters and leaf water potential of five common bean genotypes under mild water deficit

The leaf water potential, gas exchange and chlorophyll fluorescence were evaluated in five common bean (Phaseolus vulgaris) genotypes A222, A320, BAT477, Carioca and Ouro Negro subjected to moderate water deficit. At the maximum water deficit (10 d of water withholding), the leaf water potential of genotypes A320 and A222 was higher (−0.35 and −0.50 MPa) when compared to the other genotypes (−0.67 to −0.77 MPa). The stomatal conductance and net photosynthetic rate were significantly reduced in all genotypes due to the water deficit. The greater reduction in stomatal conductance of A320 under drought resulted in high intrinsic water use efficiency. Mild water deficit affected the photochemical apparatus in bean genotypes probably by down-regulation since plants did not show photoinhibition. The photochemical apparatus of A222 and A320 genotypes was more sensitive to drought stress, showing reduced apparent electron transport even after the recovery of plant water status. On the other hand, even after 10 d of water withholding, the maximum efficiency of photosystem 2 was not affected, what suggest efficiency of the photoprotection mechanisms.     

Screening for candidate bacterial biocontrol agents against soilborne fungal plant pathogens

Previous studies relating root systems and drought tolerance in oil palm focused mainly on biomass. Yet, total root length (TRL), total root surface area (TRS), and root distribution in the soil better determine water uptake. These morphological traits were studied on 3 oil palm genotypes displaying a contrasting drought tolerance. A new concept of potential root water extraction ratio (PRER) was developed using measured half-distances between roots and some assumptions about the distance of water migration from soil to root. PRER was determined in conjunction with soil moisture extraction efficiency (SMEE). The presumed tolerant genotype (T) had higher TRL, TRS and PRER than the susceptible genotype (S), whilst the performance of the control genotype (I) was intermediate. Surprisingly, during a period of moderate water deficit, T had a lower SMEE than S, which was interpreted successfully with PRER, as the result of a better access to a large volume of soil and of a slower drying out of the soil around the roots. PRER appears as a helpful indicator for comparing or ranking genotypes, and for addressing better the complexity of the genetic variability of drought tolerance.     

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.