Arbuscular mycorrhizal symbiosis and abiotic stress in plants: A review

Abiotic stresses (such as salinity, drought, cold, heat, mineral deficiency and metals/metalloids) have become major threats to the global agricultural production. These stresses in isolation and/or combination control plant growth, development and productivity by causing physiological disorders, ion toxicity, and hormonal and nutritional imbalances. Some soil microorganisms like arbuscular mycorhizal fungi (AMF) inhabit the rhizosphere and develop a symbiotic relationship with the roots of most plant species. AMF can significantly improve resistance of host plants to varied biotic and abiotic stresses. Taking into account recent literature, this paper: (a) overviews major abiotic stresses and introduces the arbuscular mycorrhizae symbiosis (b) appraises the role and underlying major mechanisms of AMF in plant tolerance to major abiotic stresses including salinity, drought, temperature regimes (cold and heat), nutrient-deficiency, and metal/metalloids; (c) discusses major molecular mechanisms potentially involved in AMF-mediated plant-abiotic stress tolerance; and finally (d) highlights major aspects for future work in the current direction.     

干旱与盐胁迫对花生根际土壤细菌群落结构和花生产量的影响

以花生品种花育25号为试验材料,采用盆栽试验研究了开花期干旱和盐分胁迫对花生生长发育和荚果产量的影响,并运用高通量测序技术分析干旱、盐胁迫及旱盐双重胁迫下,花生根际土壤细菌群落结构的变化特征。结果表明: 不同胁迫处理花生根际土壤细菌群落均以变形菌门、放线菌门、Saccharibacteria、绿弯菌门、蓝藻菌门和酸杆菌门为主。干旱、盐胁迫及旱盐双重胁迫均不同程度降低了变形菌门和放线菌门的相对丰度,但显著提高了蓝藻菌门的含量,且旱盐双重胁迫较其单一胁迫引起的根际蓝藻菌门丰度变化更显著。土壤细菌功能预测分析显示,信号转导机制、防御机制及翻译后修饰、蛋白质周转和分子伴侣等相关功能在旱盐双重胁迫的细菌中活性更强,可能对花生生长及胁迫应答具有重要影响。统计学分析显示,开花期干旱、盐胁迫和旱盐双重胁迫严重影响花生生长发育,并显著降低产量。研究结果可为通过改良土壤微生物环境来提高植物胁迫耐受性提供参考。     

Genome-Wide Bisulphite Sequencing Uncovered the Contribution of DNA Methylation to Rice Short-Term Drought Memory Formation

Rice is the important crop for more than half of the world population. However, drought can have a devastating impact on rice growth and reduce yield drastically. Understanding the response of rice to drought stress is especially important for crop breeding. Previously we found that rice enhanced its tolerance to drought stresses via stress memory mechanisms. Numerous memory genes were identified to play important roles in the process. DNA methylation was reported to mediate tolerance via regulating gene expression and enhances the survival rate of rice encountering drought stress. However, how DNA methylation involved in stress memory is still not clear. In this study, genome-wide bisulphite sequencing at a single base resolution methylome profiling level was performed and analyzed in rice cultivar under recurrent drought stresses and recovery treatments. We found that rice drought stress memory-related differentially methylated regions (DMRs) showed dynamic and distinct patterns. The drought-memory DMRs may regulate Transposable elements and gene expression to cope with short-term repeated drought stresses. Our findings of drought-memory DMRs can explain mechanisms of rice drought stress memory in a new perspective on global methylome details. Using epigenetic markers to breed drought-resistant rice would become a feasible way in the future research.

     

Differential drought-induced modulation of ozone tolerance in winter wheat species

Recent reports challenge the widely accepted idea that drought may offer protection against ozone (O(3)) damage in plants. However, little is known about the impact of drought on the magnitude of O(3) tolerance in winter wheat species. Two winter wheat species with contrasting sensitivity to O(3) (O(3) tolerant, primitive wheat, T. turgidum ssp. durum; O(3) sensitive, modern wheat, T. aestivum L. cv. Xiaoyan 22) were exposed to O(3) (83ppb O(3), 7h d(-1)) and/or drought (42% soil water capacity) from flowering to grain maturity to assess drought-induced modulation of O(3) tolerance. Plant responses to stress treatments were assessed by determining in vivo biochemical parameters, gas exchange, chlorophyll a fluorescence, and grain yield. The primitive wheat demonstrated higher O(3) tolerance than the modern species, with the latter exhibiting higher drought tolerance than the former. This suggested that there was no cross-tolerance of the two stresses when applied separately in these species/cultivars of winter wheat. The primitive wheat lost O(3) tolerance, while the modern species showed improved tolerance to O(3) under combined drought and O(3) exposure. This indicated the existence of differential behaviour of the two wheat species between a single stress and the combination of the two stresses. The observed O(3) tolerance in the two wheat species was related to their magnitude of drought tolerance under a combination of drought and O(3) exposure. The results clearly demonstrate that O(3) tolerance of a drought-sensitive winter wheat species can be completely lost under combined drought and O(3) exposure.     

The shifting influence of drought and heat stress for crops in northeast Australia

Characterization of drought environment types (ETs) has proven useful for breeding crops for drought‐prone regions. Here, we consider how changes in climate and atmospheric carbon dioxide (CO₂) concentrations will affect drought ET frequencies in sorghum and wheat systems of northeast Australia. We also modify APSIM (the Agricultural Production Systems Simulator) to incorporate extreme heat effects on grain number and weight, and then evaluate changes in the occurrence of heat‐induced yield losses of more than 10%, as well as the co‐occurrence of drought and heat. More than six million simulations spanning representative locations, soil types, management systems, and 33 climate projections led to three key findings. First, the projected frequency of drought decreased slightly for most climate projections for both sorghum and wheat, but for different reasons. In sorghum, warming exacerbated drought stresses by raising the atmospheric vapor pressure deficit and reducing transpiration efficiency (TE), but an increase in TE due to elevated CO₂ more than offset these effects. In wheat, warming reduced drought stress during spring by hastening development through winter and reducing exposure to terminal drought. Elevated CO₂ increased TE but also raised radiation‐use efficiency and overall growth rates and water use, thereby offsetting much of the drought reduction from warming. Second, adding explicit effects of heat on grain number and grain size often switched projected yield impacts from positive to negative. Finally, although average yield losses associated with drought will remain generally higher than that for heat stress for the next half century, the relative importance of heat is steadily growing. This trend, as well as the likely high degree of genetic variability in heat tolerance, suggests that more emphasis on heat tolerance is warranted in breeding programs. At the same time, work on drought tolerance should continue with an emphasis on drought that co‐occurs with extreme heat.     

Improvement of heat and drought photosynthetic tolerance in wheat by overaccumulation of glycinebetaine

Within their natural habitat, crops are often subjected to drought and heat stress, which suppress crop growth and decrease crop production. Causing overaccumulation of glycinebetaine (GB) has been used to enhance the crop yield under stress. Here, we investigated the response of wheat (Triticum aestivum L.) photosynthesis to drought, heat stress and their combination with a transgenic wheat line (T6) overaccumulating GB and its wild-type (WT) Shi4185. Drought stress (DS) was imposed by controlling irrigation until the relative water content (RWC) of the flag leaves decreased to between 78 and 82%. Heat stress (HS) was applied by exposing wheat plants to 40°C for 4 h. A combination of drought and heat stress was applied by subjecting the drought-stressed plants to a heat stress as above. The results indicated that all stresses decreased photosynthesis, but the combination of drought and heat stress exacerbated the negative effects on photosynthesis more than exposure to drought or heat stress alone. Drought stress decreased the transpiration rate (Tr), stomatal conductance (Gs) and intercellular CO₂ concentration (Ci), while heat stress increased all of these; the deprivation of water was greater under drought stress than heat stress, but heat stress decreased the antioxidant enzyme activity to a greater extent. Overaccumulated GB could alleviate the decrease of photosynthesis caused by all stresses tested. These suggest that GB induces an increase of osmotic adjustments for drought tolerance, while its improvement of the antioxidative defense system including antioxidative enzymes and antioxidants may be more important for heat tolerance.     

Emerging tools,concepts and ideas to track the modulator genes underlying plant drought adaptive traits: An overview

Crop vulnerability to multiple abiotic stresses is increasing at an alarming rate in the current global climate change scenario, especially drought. Crop improvement for adaptive adjustments to accomplish stress tolerance requires a comprehensive understanding of the key contributory processes. This requires the identification and careful analysis of the critical morpho-physiological plant attributes and their genetic control. In this review we try to discuss the crucial traits underlying drought tolerance and the various modes followed to understand their molecular level regulation. Plant stress biology is progressing into new dimensions and a conscious attempt has been made to traverse through the various approaches and checkpoints that would be relevant to tackle drought stress limitations for sustainable crop production.     

Comparative physiological analysis in the tolerance to salinity and drought individual and combination in two cotton genotypes with contrasting salt tolerance

Soil salinity and drought are the two most common and frequently co‐occurring abiotic stresses limiting cotton growth and productivity. However, physiological mechanisms of tolerance to such condition remain elusive. Greenhouse pot experiments were performed to study genotypic differences in response to single drought (4% soil moisture; D) and salinity (200 mM NaCl; S) stress and combined stresses (D + S) using two cotton genotypes Zhongmian 23 (salt‐tolerant) and Zhongmian 41 (salt‐sensitive). Our results showed that drought and salinity stresses, alone or in combination, caused significant reduction in plant growth, chlorophyll content and photosynthesis in the two cotton genotypes, with the largest impact visible under combined stress. Interestingly, Zhongmian 23 was more tolerant than Zhongmian 41 under the three stresses and displayed higher plant dry weight, photosynthesis and antioxidant enzymes activities such as superoxide dismutase (SOD), peroxidase (POD) catalase (CAT) and ascorbate peroxidase (APX) activities compared to control, while those parameters were significantly decreased in salt‐stresses Zhongmian 41 compared to control. Moreover, Na⁺/K⁺‐ATPase activity was more enhanced in Zhongmian 23 than in Zhongmian 41 under salinity stress. However, under single drought stress and D + S stress no significant differences were observed between the two genotypes. No significant differences were detected in Ca²⁺/Mg²⁺‐ATPase activity in Zhongmian 41, while in Zhongmian 23 it was increased under salinity stress. Furthermore, Zhongmian 23 accumulated more soluble sugar, glycine‐betaine and K⁺, but less Na⁺ under the three stresses compared with Zhongmian 41. Obvious changes in leaf and root tips cell ultrastructure was observed in the two cotton genotypes. However, Zhongmian 23 was less affected than Zhongmian 41 especially under salinity stress. These results give a novel insight into the mechanisms of single and combined effects of drought and salinity stresses on cotton genotypes.     

羊草叶片气体交换参数对温度和土壤水分的响应

 采用生长箱控制的方法研究了羊草(Leymus chinensis)幼苗叶片光合参数对5个温度和5个水分梯度的响应和适应。结果表明：轻度、中度土壤干旱并没有限制羊草叶片的生长,对气体交换参数亦无显著影响,反映了羊草幼苗对土壤水分胁迫的较高耐性。叶片生物量以26 ℃时最大,其它依次为23 ℃、20 ℃、29 ℃和32 ℃。温度升高使气孔导度和蒸腾速率增加, 却使光合速率和水分利用效率降低。水分和温度对叶片生物量、光合速率、气孔导度和蒸腾速率存在显著的交互作用,表明高温加强了干旱对叶片生长和气体交换的影响, 降低了羊草对土壤干旱的适应能力。高温和干旱的交互作用将显著减少我国半干旱地区草原的羊草生产力。     

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.     

干旱条件下AM真菌对植物生长和土壤水稳定性团聚体的影响

采用分室培养系统,模拟正常水分和干旱胁迫两种环境条件,探讨不同丛枝菌根真菌(arbuscular mycorrhizal fungi,AMF)对紫花苜蓿(Medicago sativa L.)生长和土壤水稳性团聚体的影响.试验条件下,Glomus intraradices对苜蓿根系的侵染率均显著高于Acaulospora scrobiculata和Diversispora spurcum接种处理.正常水分条件下,供试AM真菌均能显著提高植株生物量及磷浓度.干旱胁迫显著抑制了植株生长和菌根共生体发育,总体上菌根共生体对植株生长没有明显影响,接种D.spurcum甚至趋于降低植株生物量;同时,仅有G.intraradices显著提高了植株磷浓度.AM真菌主要影响到＞2mm的水稳性团聚体数量,以G.intraradices作用效果最为显著.在菌丝室中,G.intraradices显著提高了总球囊霉素含量.研究表明AM真菌对土壤大团聚体形成具有积极作用,而菌根效应因土壤水分条件和不同菌种而异,干旱胁迫下仅有G.intraradices对土壤结构和植物生长表现出显著积极作用.在应用菌根技术治理退化土壤时,需要选用抗逆性强共生效率高的菌株,对于不同AM真菌抗逆性差异的生物学与遗传学基础尚需进一步研究.     

Plant diversity does not buffer drought effects on early‐stage litter mass loss rates and microbial properties

Human activities are decreasing biodiversity and changing the climate worldwide. Both global change drivers have been shown to affect ecosystem functioning, but they may also act in concert in a non‐additive way. We studied early‐stage litter mass loss rates and soil microbial properties (basal respiration and microbial biomass) during the summer season in response to plant species richness and summer drought in a large grassland biodiversity experiment, the Jena Experiment, Germany. In line with our expectations, decreasing plant diversity and summer drought decreased litter mass loss rates and soil microbial properties. In contrast to our hypotheses, however, this was only true for mass loss of standard litter (wheat straw) used in all plots, and not for plant community‐specific litter mass loss. We found no interactive effects between global change drivers, that is, drought reduced litter mass loss rates and soil microbial properties irrespective of plant diversity. High mass loss rates of plant community‐specific litter and low responsiveness to drought relative to the standard litter indicate that soil microbial communities were adapted to decomposing community‐specific plant litter material including lower susceptibility to dry conditions during summer months. Moreover, higher microbial enzymatic diversity at high plant diversity may have caused elevated mass loss of standard litter. Our results indicate that plant diversity loss and summer drought independently impede soil processes. However, soil decomposer communities may be highly adapted to decomposing plant community‐specific litter material, even in situations of environmental stress. Results of standard litter mass loss moreover suggest that decomposer communities under diverse plant communities are able to cope with a greater variety of plant inputs possibly making them less responsive to biotic changes.     

Soil microbial moisture dependences and responses to drying–rewetting: The legacy of 18 years drought

Climate change will alter precipitation patterns with consequences for soil C cycling. An understanding of how fluctuating soil moisture affects microbial processes is therefore critical to predict responses to future global change. We investigated how long‐term experimental field drought influences microbial tolerance to lower moisture levels (“resistance”) and ability to recover when rewetted after drought (“resilience”), using soils from a heathland which had been subjected to experimental precipitation reduction during the summer for 18 years. We tested whether drought could induce increased resistance, resilience, and changes in the balance between respiration and bacterial growth during perturbation events, by following a two‐tiered approach. We first evaluated the effects of the long‐term summer drought on microbial community functioning to drought and drying–rewetting (D/RW), and second tested the ability to alter resistance and resilience through additional perturbation cycles. A history of summer drought in the field selected for increased resilience but not resistance, suggesting that rewetting after drought, rather than low moisture levels during drought, was the selective pressure shaping the microbial community functions. Laboratory D/RW cycles also selected for communities with a higher resilience rather than increased resistance. The ratio of respiration to bacterial growth during D/RW perturbation was lower for the field drought‐exposed communities and decreased for both field treatments during the D/RW cycles. This suggests that cycles of D/RW also structure microbial communities to respond quickly and efficiently to rewetting after drought. Our findings imply that microbial communities can adapt to changing climatic conditions and that this might slow the rate of soil C loss predicted to be induced by future cyclic drought.     

增强UV-B辐射和干旱对春小麦光合作用及其生长的影响

在室外盆栽条件下研究了UV-B辐射和土壤干旱对春小麦 '和尚头'生长和光合作用的影响.结果显示:(1)干旱、UV-B辐射、干旱+UV-B(复合)处理均可使叶片类黄酮含量增加,且干旱+UV-B处理增加显著高于其他处理(P<0.05).UV-B辐射和干旱单独处理均能显著降低叶片光合色素含量,但UV-B辐射的副作用大于干旱,复合处理对光合色素的影响介于UV-B和干旱之间.(2)各处理间的光合速率日均值大小次序为:对照>UV-B+干旱>UV-B>干旱;增强UV-B对净光合速率的抑制作用大于干旱,而UV-B+干旱处理的抑制作用较二者单独处理有所减轻.(3)UV-B辐射和干旱单独处理后总生物量比对照减少15%,且抑制作用为:干旱>UV-B>复合处理; UV-B辐射和干旱胁迫不但影响春小麦的生物量,而且影响小穗特征和产量.研究表明,UV-B辐射和干旱之间存在交互作用,说明一种胁迫可以减缓(轻)另外一种胁迫对春小麦的抑制作用.     

Role of AP2/EREBP transcription factors in gene regulation during abiotic stress.

Crop plants are exposed to many types of abiotic stress during their life cycle. Water deficit derived from drought, low temperature or high salt concentration in the soil, is one of the most common environmental stresses that affects growth and development of plants through alterations in metabolism and gene expression. Adaptation to these conditions may involve passive tolerance or active homeostatic mechanisms for maintaining water balance. Active responses occur at different levels in the plant and may represent a concomitant protection against other types of stress such as pathogen attack. Many morphological and physiological adaptations to water stress are under the control of the plant hormone abscisic acid and involve specific activation of target genes that in one way or another protect cells against water deficit or participate in the regulation of the drought response. Here, we discuss recent advances in our understanding of drought adaptation mediated by specific changes in gene expression and the role of AP2/EREBP nuclear factors in these processes.     

Methods and concepts in quantifying resistance to drought, salt and freezing, abiotic stresses that affect plant water status

The abiotic stresses of drought, salinity and freezing are linked by the fact that they all decrease the availability of water to plant cells. This decreased availability of water is quantified as a decrease in water potential. Plants resist low water potential and related stresses by modifying water uptake and loss to avoid low water potential, accumulating solutes and modifying the properties of cell walls to avoid the dehydration induced by low water potential and using protective proteins and mechanisms to tolerate reduced water content by preventing or repairing cell damage. Salt stress also alters plant ion homeostasis, and under many conditions this may be the predominant factor affecting plant performance. Our emphasis is on experiments that quantify resistance to realistic and reproducible low water potential (drought), salt and freezing stresses while being suitable for genetic studies where a large number of lines must be analyzed. Detailed protocols for the use of polyethylene glycol-infused agar plates to impose low water potential stress, assay of salt tolerance based on root elongation, quantification of freezing tolerance and the use of electrolyte leakage experiments to quantify cellular damage induced by freezing and low water potential are also presented.