Enhancing drought tolerance in C(4) crops

Adaptation to abiotic stresses is a quantitative trait controlled by many different genes. Enhancing the tolerance of crop plants to abiotic stresses such as drought has therefore proved to be somewhat elusive in terms of plant breeding. While many C(4) species have significant agronomic importance, most of the research effort on improving drought tolerance has focused on maize. Ideally, drought tolerance has to be achieved without penalties in yield potential. Possibilities for success in this regard are highlighted by studies on maize hybrids performed over the last 70 years that have demonstrated that yield potential and enhanced stress tolerance are associated traits. However, while our understanding of the molecular mechanisms that enable plants to tolerate drought has increased considerably in recent years, there have been relatively few applications of DNA marker technologies in practical C(4) breeding programmes for improved stress tolerance. Moreover, until recently, targeted approaches to drought tolerance have concentrated largely on shoot parameters, particularly those associated with photosynthesis and stay green phenotypes, rather than on root traits such as soil moisture capture for transpiration, root architecture, and improvement of effective use of water. These root traits are now increasingly considered as important targets for yield improvement in C(4) plants under drought stress. Similarly, the molecular mechanisms underpinning heterosis have considerable potential for exploitation in enhancing drought stress tolerance. While current evidence points to the crucial importance of root traits in drought tolerance in C(4) plants, shoot traits may also be important in maintaining high yields during drought.     

Advances in Drought Resistance of Rice

Water deficit is a serious environmental stress and the major constraint to rice productivity. Losses in rice yield due to water shortage probably exceed losses from all other causes combined and the extent of the yield loss depends on both the severity and duration of the water stress. Drought affects rice at morphological, physiological, and molecular levels such as delayed flowering, reduced dry matter accumulation and partitioning, and decreased photosynthetic capacity as a result of stomatal closure, metabolic limitations, and oxidative damage to chloroplasts. Small-statured rice plants with reduced leaf area and short growth duration are better able to tolerate drought stress, although the mechanisms are not yet fully understood. Increased water uptake by developing larger and deeper root systems, and the accumulation of osmolytes and osmoprotectants are other important mechanisms for drought resistance. Drought resistance in rice has been improved by using plant growth regulators and osmoprotectants. In addition, several enzymes have been found that act as antioxidants. Silicon has also improved drought resistance in rice by silicification of the root endodermis and improving water uptake. Seed priming improves germination and crop stand establishment under drought. Rice plants expressing HVA1, LEA proteins, MAP kinase, DREB and endo-1, 3-glucanase are better able to withstand drought stress. Polyamines and several enzymes act as antioxidants and reduce adverse effects of drought stress in rice. Drought resistance can be managed by developing and selecting drought-tolerant genotypes. Rice breeding and screening may be based on growth duration, root system, photosynthesis traits, stomatal frequency, specific leaf weight, leaf water potential, and yield in target environments. This review discusses recent developments in integrated approaches, such as genetics, breeding and resource management to increase rice yield and reduce water demand for rice production.     

Overexpression of HVA1 gene from barley generates tolerance to salinity and water stress in transgenic mulberry (Morus indica)

Late embryogenesis abundant (LEA) proteins are members of a large group of hydrophilic proteins found primarily in plants. The barley hva1 gene encodes a group 3 LEA protein and is induced by ABA and water deficit conditions. We report here the over expression of hva1 in mulberry under a constitutive promoter via Agrobacterium-mediated transformation. Molecular analysis of the transgenic plants revealed the stable integration and expression of the transgene in the transformants. Transgenic plants were subjected to simulated salinity and drought stress conditions to study the role of hva1 in conferring tolerance. The transgenic plants showed better cellular membrane stability (CMS), photosynthetic yield, less photo-oxidative damage and better water use efficiency as compared to the non-transgenic plants under both salinity and drought stress. Under salinity stress, transgenic plants show many fold increase in proline concentration than the non-transgenic plants and under water deficit conditions proline is accumulated only in the non-transgenic plants. Results also indicate that the production of HVA1 proteins helps in better performance of transgenic mulberry by protecting membrane stability of plasma membrane as well as chloroplastic membranes from injury under abiotic stress. Interestingly, it was observed that hva1 conferred different degrees of tolerance to the transgenic plants towards various stress conditions. Amongst the lines analysed for stress tolerance transgenic line ST8 was relatively more salt tolerant, ST30, ST31 more drought tolerant, and lines ST11 and ST6 responded well under both salinity and drought stress conditions as compared to the non-transgenic plants. Thus hva1 appears to confer a broad spectrum of tolerance under abiotic stress in mulberry.     

Photosynthesis and antioxidative defense mechanisms in deciphering drought stress tolerance of crop plants

Crop plants are regularly exposed to an array of abiotic and biotic stresses, among them drought stress is a major environmental factor that shows adverse effects on plant growth and productivity. Because of this these factors are considered as hazardous for crop production. Drought stress elicits a plethora of responses in plants resulting in strict amendments in physiological, biochemical, and molecular processes. Photosynthesis is the most fundamental physiological process affected by drought due to a reduction in the CO₂ assimilation rate and disruption of primary photosynthetic reactions and pigments. Drought also expedites the generation of reactive oxygen species (ROS), triggering a cascade of antioxidative defense mechanisms, and affects many other metabolic processes as well as affecting gene expression. Details of the drought stress-induced changes, particularly in crop plants, are discussed in this review, with the major points: 1) leaf water potentials and water use efficiency in plants under drought stress; 2) increased production of ROS under drought leading to oxidative stress in plants and the role of ROS as signaling molecules; 3) molecular responses that lead to the enhanced expression of stress-inducible genes; 4) the decrease in photosynthesis leading to the decreased amount of assimilates, growth, and yield; 5) the antioxidant defense mechanisms comprising of enzymatic and non-enzymatic antioxidants and the other protective mechanisms; 6) progress made in identifying the drought stress tolerance mechanisms; 7) the production of transgenic crop plants with enhanced tolerance to drought stress.     

植物干旱生态生理的研究

随着科学研究的发展,涉猎的范畴越来越广,探索的问题越来越深,从而由一些单一的学科如物理学、化学、数学、生物学发展出许多边缘学科,如生物物理学、生物化学、生物     

Improving plant drought tolerance and growth under water limitation through combinatorial engineering of signalling networks

Agriculture is by far the biggest water consumer on our planet, accounting for 70 per cent of all freshwater withdrawals. Climate change and a growing world population increase pressure on agriculture to use water more efficiently (‘more crop per drop’). Water‐use efficiency (WUE) and drought tolerance of crops are complex traits that are determined by many physiological processes whose interplay is not well understood. Here, we describe a combinatorial engineering approach to optimize signalling networks involved in the control of stress tolerance. Screening a large population of combinatorially transformed plant lines, we identified a combination of calcium‐dependent protein kinase genes that confers enhanced drought stress tolerance and improved growth under water‐limiting conditions. Targeted introduction of this gene combination into plants increased plant survival under drought and enhanced growth under water‐limited conditions. Our work provides an efficient strategy for engineering complex signalling networks to improve plant performance under adverse environmental conditions, which does not depend on prior understanding of network function.     

植物干旱胁迫下水分代谢、碳饥饿与死亡机理

植物在生长发育过程中受众多环境因子共同作用。随着全球气候变化,气温升高、降水量下降等问题频繁出现。目前气象学家一致预测未来环境变暖会使干旱更加频繁剧烈,这一环境改变使植物死亡更加严重。植物在水分胁迫、特别是干旱胁迫条件下,体内水分代谢与碳代谢会发生失衡现象:光合速率降低、蒸腾速率降低,带来生长降低;为维持植物新陈代谢,植物呼吸作用必然下调。在长期干旱胁迫条件下植物体内碳水化合物储存发生失衡现象,这种失衡使植物陷入碳饥饿现象。另外,由于水分失衡而出现的木质部栓塞和空穴会进一步加剧水分运输障碍,而修复空穴则需要大量非结构性碳水化合物(NSC),这使植物陷入两难选择。总结了植物干旱胁迫下,碳饥饿与水分代谢、植物死亡关系的相关研究,对未来的研究方向和重点提出建议,以期对未来的植物死亡研究提供帮助。     

Interaction between PGPR and PGR for water conservation and plant growth attributes under drought condition

Drought is one of the key restraints to agricultural productivity worldwide and is expected to increase further. Drought stress accompanied by reduction in precipitation pose major challenges to future food safety. Strategies should be develop to enhance drought tolerance in crops like chickpea and wheat, in order to enhance their growth and yield. Drought tolerance strategies are costly and time consuming however, recent studies specify that plant growth promoting rhizobacteria (PGPR) and plant growth regulators (PGRs) can help plants to withstand under harsh environmental condition and enable plants to cope with drought stress. PGPR can act as biofertilizer and bioenhancer for different legumes and non-legumes. The use of PGPR and symbiotic microorganisms, may be valuable in developing strategies to assist water conservation in plants. The use of PGPR has been confirmed to be an ecologically sound way of enhancing crop yields by facilitating plant growth through direct or indirect mechanism. The mechanisms of PGPR for water conservation include secretion of exopolysaccharides, biofilm formation, alternation in phytohormone content, improvement in sugar concentration, enhancing availability of micro- and macronutrients and changes in plant functional traits. Similarly, plant growth regulators (PGRs) are specially noticed in actively growing tissues under stress conditions and have been associated in the control of cell division, embryogenesis, root formation, fruit development and ripening, and reactions to biotic and abiotic stresses and upholding water conservation status in plants. Previous studies also suggest that plant metabolites interact with plant physiology under stress condition and impart drought tolerance. Metabolites like, sugars, amino acids, organic acid and polyols play a key role in drought tolerance of crop plants grown under stress condition. It is concluded from the present study that PGRs in combination with PGPR consortium can be an effective formulation to promote plant growth and maintenance of plant turgidity under drought stress. This review is a compilation of the effect of drought stress on crop plants and described interactions between PGPR/PGRs and plant development, knowledge of water conservation and stress release strategies of PGPR and PGRs and the role of plant metabolites in drought tolerance of crop plants. This review also bridges the gaps that summarizes the mechanism of action of PGPR for drought tolerance of crop plants and sustainability of agriculture and applicability of these beneficial rhizobacteria in different agro-ecosystems under drought stress.     

硅对干旱胁迫下玉米水分代谢的影响

利用盆栽试验研究了施硅(K2SiO3)对玉米植株水分代谢的影响.结果表明:施硅降低了干旱胁迫下玉米植株的气孔导度,降低了干旱胁迫早期到中期的蒸腾速率,保持了干旱胁迫后期较高的蒸腾速率,从而导致施硅玉米植株的叶片含水量和水势高于对照.由于植株的水分状况改善,施硅玉米植株生物量高于对照.硅增强玉米植株的抗旱性,而提高植株保水能力是硅提高抗旱性的重要原因.     

Effect of water stress and NaCl triggered changes on yield,physiology, biochemistry of broad bean (Vicia faba) plants and on quality of harvested pods

Salinity and drought are the most important abiotic stresses affecting crop yield. Broad bean was chosen as model plant for assessing the impact of salt stress and its interaction with drought in the field experiments. The factors examined in the experiments were the two irrigation rates (normal watering — NW with 3 L plant^?1 and drought — D) and three salinity rates imposed by foliar application (0, 50, 100 mg L^?1 NaCl). Highest NaCl level with normal water irrigation caused maximum reduction in plant height and production, which it was due to photosynthetic disturbances. Salt injuries were alleviated by increasing water stress. The control plants exposed to NaCl lost their ability over water control. The increased malondialdehyde (MDA) and H₂O₂ indicate the prevalence of oxidative stress due to salinity. The levels of proline and carbohydrates were higher under salinity alone than under simultaneous exposure to drought and NaCl. The protein concentration of immature and mature broad bean pods was more inhibited more by NaCl supply than by drought alone. The combination of drought and NaCl resulted in a significant increase in proteins, glucose, fructose and sucrose content. Overall, the ameliorative effect of drought under NaCl supply was quantified.     

天山雪莲冷调节蛋白基因siCOR转化烟草植株的抗旱性分析

冷调节蛋白(cold regulated proteins, CORPs)是植物在冷驯化下产生的特异性蛋白, 与植物的抗寒性密切相关。然而, 大量研究表明, 绝大多数植物冷诱导基因同样会响应水分胁迫。采用半定量RT-PCR分析天山雪莲(Sasussured involucrata)冷调节蛋白基因siCOR的表达, 结果表明siCOR是一个受干旱胁迫诱导表达的基因。为研究siCOR基因是否与抗旱性相关, 以siCOR转基因烟草为研究材料, 利用水分胁迫处理进行抗旱性分析。结果表明与野生型(wild-type, WT)相比, 转siCOR植株叶片萎蔫较迟且程度较轻, 复水后恢复快且较完全; 其叶片相对含水量和PSII相对量子产率的降低幅度、相对电导率和丙二醛含量的升高幅度均低于野生型烟草植株。采用PEG6000模拟干旱胁迫, 发现转siCOR植株T3代种子的萌发率较高, 主根生长的受抑制程度较野生型轻。以上结果表明, siCOR基因在植物对干旱胁迫的响应中起重要作用。     

Effect of arbuscular mycorrhizal inoculation on water status and photosynthesis of Populus cathayana males and females under water stress

Drought is one of the most serious environmental limitations for poplar growth. Although the ways in which plants deal with water stress and the effects of arbuscular mycorrhizal (AM) formation have been well documented, little is known about how the male and female plants of Populus cathayana respond to drought and AM formation. We also aimed to investigate the potential role of AM fungi in maintaining gender balance. We tested the impact of drought and AM formation on water status and photosynthesis. The results suggested that both sexes showed similar responses to water stress: drought decreased the growth of stem length (GSL), growth of ground diameter (GGD), relative water content (RWC), increased the relative electrolyte leakage (REL), and limited the photosynthesis and chlorophyll fluorescence indexes. However, the responses of the two sexes to drought and AM formation differed to some extent. AM formation had positive effects on RWC, photosynthesis and the intrinsic water use efficiency (WUEi) but negative effects on the REL of males and females, especially under drought. AM formation enhanced the maximum quantum yield of photosystem II (PSII) (Fv/Fm), the actual quantum yield of PSII (ΦPSII), non‐photochemical quenching (qN) and photochemical quenching (qP) under drought conditions, and had no significant effects under well‐watered conditions except on the qP of males. Principal component analysis showed that males were significantly more drought tolerant than females, and AM formation enhanced drought tolerance, particularly among males, which suggested that AM fungi are beneficial for ecological stability and for P. cathayana survival under drought conditions.     

Phloem exudate metabolic content reflects the response to water-deficit stress in pea plants (Pisum sativum L.)

Drought stress impacts the quality and yield of Pisum sativum. Here, we show how short periods of limited water availability during the vegetative stage of pea alters phloem sap content and how these changes are connected to strategies used by plants to cope with water deficit. We have investigated the metabolic content of phloem sap exudates and explored how this reflects P. sativum physiological and developmental responses to drought. Our data show that drought is accompanied by phloem-mediated redirection of the components that are necessary for cellular respiration and the proper maintenance of carbon/nitrogen balance during stress. The metabolic content of phloem sap reveals a shift from anabolic to catabolic processes as well as the developmental plasticity of P. sativum plants subjected to drought. Our study underlines the importance of phloem-mediated transport for plant adaptation to unfavourable environmental conditions. We also show that phloem exudate analysis can be used as a useful proxy to study stress responses in plants. We propose that the decrease in oleic acid content within phloem sap could be considered as a potential marker of early signalling events mediating drought response.     

Ectopic expression of ADP ribosylation factor 1 (SaARF1) from smooth cordgrass (Spartina alterniflora Loisel) confers drought and salt tolerance in transgenic rice and Arabidopsis

Salinity and drought are two very important abiotic stressors that negatively impact the growth and yield of all sensitive crop plants. Genes from halophytes have been shown to be useful to engineer crop plants that can survive under adverse soil and water conditions. The present report establishes, for the first time, the physiological role of a class one ADP ribosylation factor gene (SaARF1) from the halophyte Spartina alterniflora (smooth cordgrass) in imparting salinity and drought stress tolerance when expressed in both monocot (rice) and dicot (Arabidopsis) systems. The Arabidopsis and rice plants overexpressing ARF1 are many-fold more tolerant to salt and drought than wild-type (WT) plants. The transgenics exhibited improved growth and productivity relative to WT through tissue tolerance by maintaining higher relative water content and membrane stability, and higher photosynthetic yield by retaining higher chlorophyll concentration and fluorescence under stress conditions compared to WT. These findings indicated that genes from halophyte resources can be useful to engineer and improve salt and drought stress tolerance in both monocot and dicot plants.     

Increase of glycinebetaine synthesis improves drought tolerance in cotton

The tolerance to drought stress of the homozygous transgenic cotton (Gossypium hirsutum L.) plants with enhanced glycinebetaine (GB) accumulation was investigated at three development stages. Among the five transgenic lines investigated, lines 1, 3, 4, and 5 accumulated significantly higher levels of GB than the wild-type (WT) plants either before or after drought stress, and the transgenic plants were more tolerant to drought stress than the wild-type counterparts from young seedlings to flowering plants. Under drought stress conditions, transgenic lines 1, 3, 4, and 5 had higher relative water content, increased photosynthesis, better osmotic adjustment (OA), a lower percentage of ion leakage, and less lipid membrane peroxidation than WT plants. The GB levels in transgenic plants were positively correlated with drought tolerance under water stress. The results suggested that GB may not only protect the integrity of the cell membrane from drought stress damage, but also be involved in OA in transgenic cotton plants. Most importantly, the seedcotton yield of transgenic line 4 was significantly greater than that of WT plants after drought stress, which is of great value in cotton production.     

The effect of drought stress on the sex ratio variation ofSilene otites

Under dry environmental conditions the sex ratio of many dioecious plants is male-biased, which is usually explained by the higher susceptibility of females to drought stress. We investigated if spatio-temporal variation in the sex ratio ofSilene otites could be explained by the higher sensitivity of female plants to drought stress as compared to males. Long-term field observations, however, did not support this hypothesis. The sex ratio in 34 patches at the study site in Central Germany changed from slightly female biased in 1994 to strongly male-biased in 1997 and 1998. The interannual change in the proportion of plants that were female was positively correlated with the number of days with soil-water deficit in the late summer, suggesting higher mortality in males than in females under drought stress. In two closely studied patches, mortality in males was also higher than in females, although this difference could not be related to drought stress. These field observations were supported by an experiment with potted plants in two climate chambers, in which male mortality was higher during a three-week period without water supply. We conclude that the often reported male bias in patches ofS. otites is not caused by sexual differences in the sensitivity to drought stress. Field data in this study, however, suggest that maleS. otites plants flower earlier than females, which causes a shift in sex ratio to more male bias among flowering plants.     

植物气孔对全球环境变化的响应及其调控防御机制

气孔是植物与环境发生联系的重要门户,控制着植物与外界的气体和水分交换。本文针对全球大气CO2浓度升高、气候变暖、干旱加剧等环境问题,分析了气孔对全球水循环、碳循环的重要贡献。系统总结了气孔的形态发育和生理功能对大气高CO2浓度、干旱、土壤盐渍化、病虫害等的响应及其调控防御机制。综述了脱落酸（ABA）、Ca2＋、H2O2、一氧化氮（NO）和光信号调控气孔运动的分子机制。从理论和实践两方面,提出了通过调控气孔运动协调CO2同化和水分散失的矛盾,在不影响光合效率的前提下提高水分利用率等未来的研究方向。