干旱对陆地生态系统生产力的影响

该文综述了干旱对陆地生态系统生产力的影响,分析了其影响机制,并总结了植被对干旱的响应与适应及其机理机制。干旱通过抑制光合作用来降低陆地生态系统总初级生产力,干旱还可以降低生态系统的自养呼吸和异养呼吸。同时干旱还可以通过影响其它干扰形式来间接影响陆地生态系统生产力,如增加火干扰的发生频率和强度,增加植物的死亡率,增加病虫害的发生等。在生态系统水平上干旱可以降低碳固定,减弱碳汇功能,甚至把生态系统从碳汇改变成碳源。目前生态系统水平上的干旱影响研究主要通过两种方法实现,一种是模型模拟,另一种就是大型模拟实验。作为陆地生态系统生产力的实现者,在干旱胁迫条件下,植物也会采取积极的适应策略以减弱干旱对生态系统生产力的影响,其适应策略主要分以下3种:在一些周期性发生干旱的地区,植物会调整生长期以避开干旱或通过休眠来减弱干旱所造成的伤害;还有一些植物会通过调节体内的代谢过程,改变一些生理特性来抵御干旱;而长期生活在干旱条件下的植物则通过进化来改变了自身的生理生化代谢过程,形成耐旱机制。目前,植物对干旱响应的分子学机制,以及生态系统水平上对干旱的响应和适应仍然是薄弱的领域,也必然成为未来研究的重点。     

Effects of extreme drought on terrestrial ecosystems: review and prospects

作为地球表层重要的组成部分, 陆地生态系统是人类生存和发展的重要场所。进入21世纪以来, 气候变化导致干旱事件发生的强度、频度和持续时间显著增加, 对陆地生态系统带来深远的影响, 严重制约甚至威胁人类社会的可持续发展。因此, 开展极端干旱对陆地生态系统影响的研究并评估其生态风险效应, 是当前全球变化领域研究的重点问题。该文从植物生理生态过程、生物地球化学循环、生物多样性以及生态系统结构和功能4个方面综述了极端干旱对陆地生态系统的影响, 并对当前的研究热点进行探讨, 深度剖析当前研究中存在的难点问题和未来可能的发展方向, 以期为未来开展干旱对陆地生态系统影响的观测与预测研究提供参考, 为在未来干旱影响下加强陆地生态系统风险评估和管理提供新思路。     

黄土高原植被日光诱导叶绿素荧光对气象干旱的响应

随着气候变化的加剧,干旱的频率、持续时间以及发生范围都越来越严重,探索植被光合对干旱的响应以及气象因子对植被光合的影响对于人们如何应对干旱具有重要意义。基于遥感的日光诱导叶绿素荧光(SIF)具有对干旱条件下区域植被光合作用进行早期监测和准确评估的潜力。本研究基于星载SIF和标准化降水蒸散发指数(SPEI)研究了黄土高原地区2001—2017年生长季内(4—10月)植被光合作用对干旱的响应关系及其受气象因子的影响程度。结果表明: 黄土高原地区植被生长季内SIF与SPEI呈显著正相关关系的区域占比为87.8%,其中,半干旱地区植被光合对干旱的响应较敏感,半湿润地区敏感性较低。不同类型植被光合对干旱的响应存在差异,草地对干旱响应的敏感性最高,响应最强的SPEI时间尺度为3～4个月;林地的敏感性最低,SPEI时间尺度为3～10个月。气象因子与SIF存在显著的相关关系,其中,温度和降雨是影响黄土高原植被光合的重要影响因子,光合有效辐射的影响模式与温度相似。黄土高原地区生长季内不同的气候和植被类型条件下,植被光合所受干旱及各气象因素的影响存在较大差异。     

Soil moisture determines the recovery time of ecosystems from drought

Recovery time, the time it takes for ecosystems to return to normal states after experiencing droughts, is critical for assessing the response of ecosystems to droughts; however, the spatial dominant factors determining recovery time are poorly understood. We identify the global patterns of terrestrial ecosystem recovery time based on remote sensed vegetation indices, analyse the affecting factors of recovery time using random forest regression model, and determine the spatial distribution of the dominant factors of recovery time based on partial correlation. The results show that the global average recovery time is approximately 3.3 months, and that the longest recovery time occurs in mid-latitude drylands. Analysis of affecting factors of recovery time suggests that the most important environmental factor affecting recovery time is soil moisture during the recovery period, followed by temperature and vapour pressure deficit (VPD). Recovery time shortens with increasing soil moisture and prolongs with increasing VPD; however, the response of recovery time to temperature is nonmonotonic, with colder or hotter temperatures leading to longer recovery time. Soil moisture dominates the drought recovery time over 58.4% of the assessed land area, mostly in the mid-latitudes. The concern is that soil moisture is projected to decline in more than 65% regions in the future, which will lengthen the drought recovery time and exacerbate drought impacts on terrestrial ecosystems, especially in southwestern United States, the Mediterranean region and southern Africa. Our research provides methodological insights for quantifying recovery time and spatially identifies dominant factors of recovery time, improving our understanding of ecosystem response to drought.     

Effects of drought on nutrient and ABA transport in Ricinus communis

We studied the effects of variations of water flux through the plant, of diurnal variation of water flux, and of variation of vapour pressure deficit at the leaf on compensation pressure in the Passioura-type pressure chamber, the composition of the xylem sap and leaf conductance in Ricinus communis. The diurnal pattern of compensation pressure showed stress relaxation during the night hours, while stress increased during the day, when water limitation increased. Thus compensation pressure was a good measure of the momentary water status of the root throughout the day and during drought. The bulk soil water content at which predawn compensation pressure and abscisic acid concentration in the xylem sap increased and leaf conductance decreased, was high when the water usage of the plant was high. For all xylem sap constituents analysed, variations in concentrations during the day were larger than changes in mean concentrations with drought. Mean concentrations of phosphate and the pH of the xylem sap declined with drought, while nitrate concentration remained constant. When the measurement leaf was exposed to a different VPD from the rest of the plant, leaf conductance declined by 400mmol m^?2 s^?1 when compensation pressure increased by 1 MPa in all treatments. The compensation pressure needed to keep the shoot turgid, leaf conductance and the abscisic acid concentration in the xylem were linearly related. This was also the case when the highly dynamic development of stress was taken into account.     

植物应对干旱胁迫的气孔调节

气孔是植物控制叶片与大气之间碳、水交换的重要门户,植物的生长和生存都依赖于叶片气孔对碳获取和水散失的调控.因此,气孔调节机理研究与气孔导度模型研发是精确模拟陆地生态系统碳、水循环过程不可或缺的内容.近年来,随着气候变化的加剧,干旱事件愈发频繁,对植物的存活、生长和分布产生深刻影响.为了深入理解植物碳-水耦合机理过程、预测全球变化下植物及群落的动态,开展植物应对干旱胁迫的气孔调节研究尤为重要.本文综述了植物在干旱胁迫条件下气孔调节机制和模型研究进展.首先阐述了植物气孔对干旱胁迫的主动调节与被动调节,讨论了气孔调节的演化过程,包括蕨类和石松类植物的被动水力调节、被子植物的主动调节和裸子植物的双重调节机制,认为裸子植物的气孔调节方式是植物进化过程中介于蕨类、石松类植物和被子植物之间的一种重要过渡类型.然后分析了气孔调节与水力调节的关系,讨论了“植物水势和气孔导度解耦”问题中存在的争议.之后介绍了基于水分利用效率假说和最大碳增益假说所建立的气孔导度优化模型的应用,并指出后者有更强的预测能力和应用前景.最后,为了有效减少植被对气候变化响应预测中的不确定性,提出了2个亟待开展的研究问题:将植物叶片的气孔调节功能研究由个体扩展到生态系统甚至更大尺度,改进陆地生态系统碳水循环机理模型;量化气孔调节的主动水力反馈过程,修正植物气孔功能水力模型.     

A meta-analysis on morphological,physiological and biochemical responses of plants with PGPR inoculation under drought stress

Plant growth-promoting rhizobacteria (PGPR) can help plants to resist drought stress. However, the mechanisms of how PGPR inoculation affect plant status under drought remain incompletely understood. We performed a meta-analysis of plant response to PGPR inoculation by compiling data from 57 PGPR-inoculation studies, including 2, 387 paired observations on morphological, physiological and biochemical parameters under drought and well-watered conditions. We compare the PGPR effect on plants performances among different groups of controls and treatments. Our results reveal that PGPR enables plants to restore themselves from drought-stressed to near a well-watered state, and that C4 plants recover better from drought stress than C3 plants. Furthermore, PGPR is more effective underdrought than well-watered conditions in increasing plant biomass, enhancing photosynthesis and inhibiting oxidant damage, and the responses of C4 plants to the PGPR effect was stronger than that of C3 plants under drought conditions. Additionally, PGPR belonging to different taxa and PGPR with different functional traits have varying degrees of drought-resistance effects on plants. These results are important to improve our understanding of the PGPR beneficial effects on enhanced drought-resistance of plants.     

Adjustments of water use efficiency by stomatal regulation during drought and recovery in the drought-adapted Vitis hybrid Richter-110 (V. berlandieri x V. rupestris)

The hybrid Richter-110 (Vitis berlandieri x Vitis rupestris) (R-110) has the reputation of being a genotype strongly adapted to drought. A study was performed with plants of R-110 subjected to water withholding followed by re-watering. The goal was to analyze how stomatal conductance (g(s)) is regulated with respect to different physiological variables under water stress and recovery, as well as how water stress affects adjustments of water use efficiency (WUE) at the leaf level. Water stress induced a substantial stomatal closure and an increase in WUE, which persisted many days after re-watering. The g(s) during water stress was mainly related to the content of ABA in the xylem and partly related to plant hydraulic conductivity but not to leaf water potential. By contrast, low g(s) during re-watering did not correlate with ABA contents and was only related to a sustained decreased hydraulic conductivity. In addition to a complex physiological regulation of stomatal closure, g(s) and rate of transpiration (E) were strongly affected by leaf-to-air vapor pressure deficit (VPD) in a way dependent of the treatment. Interestingly, E increased with increasing VPD in control plants, but decreased with increasing VPD in severely stressed plants. All together, the fine stomatal regulation in R-110 resulted in very high WUE at the leaf level. This genotype is revealed to be very interesting for further studies on the physiological mechanisms leading to regulation of stomatal responsiveness and WUE in response to drought.     

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.     

Measuring canopy loss and climatic thresholds from an extreme drought along a fivefold precipitation gradient across Texas

Globally, trees are increasingly dying from extreme drought, a trend that is expected to increase with climate change. Loss of trees has significant ecological, biophysical, and biogeochemical consequences. In 2011, a record drought caused widespread tree mortality in Texas. Using remotely sensed imagery, we quantified canopy loss during and after the drought across the state at 30‐m spatial resolution, from the eastern pine/hardwood forests to the western shrublands, a region that includes the boundaries of many species ranges. Canopy loss observations in ~200 multitemporal fine‐scale orthophotos (1‐m) were used to train coarser Landsat imagery (30‐m) to create 30‐m binary statewide canopy loss maps. We found that canopy loss occurred across all major ecoregions of Texas, with an average loss of 9.5%. The drought had the highest impact in post oak woodlands, pinyon‐juniper shrublands and Ashe juniper woodlands. Focusing on a 100‐km by ~1,000‐km transect spanning the State's fivefold east–west precipitation gradient (~1,500 to ~300 mm), we compared spatially explicit 2011 climatic anomalies to our canopy loss maps. Much of the canopy loss occurred in areas that passed specific climatic thresholds: warm season anomalies in mean temperature (+1.6°C) and vapor pressure deficit (VPD, +0.66 kPa), annual percent deviation in precipitation (?38%), and 2011 difference between precipitation and potential evapotranspiration (?1,206 mm). Although similarly low precipitation occurred during the landmark 1950s drought, the VPD and temperature anomalies observed in 2011 were even greater. Furthermore, future climate data under the representative concentration pathway 8.5 trajectory project that average values will surpass the 2011 VPD anomaly during the 2070–2099 period and the temperature anomaly during the 2040–2099 period. Identifying vulnerable ecological systems to drought stress and climate thresholds associated with canopy loss will aid in predicting how forests will respond to a changing climate and how ecological landscapes will change in the near term.     

Plant drought tolerance assessment for re-vegetation in heterogeneous karst landscapes of southwestern China

A better understanding of plant water relations is needed for evaluating the suitability of plant species to site-specific reforestation programs in the heterogeneous karst landscapes in southwestern China that are characterized by temporary water deficit. During both wet and dry periods, leaf water potentials of 65 plant species from five different growth forms were studied at three representative sites (forest, shrubland and grassland), to compare their adaptive strategies against water stress and assess their suitability for reforestation programs. Herbs showed the highest predawn and midday water potentials and smallest diurnal ranges of water potential values at all the three sites, indicating that they follow water stress avoidance strategies. During the dry period, evergreen shrubs showed low water potentials, the largest diurnal ranges and highest soluble sugar contents. This indicates that they have a tolerance strategy responding to water stress. Deciduous shrubs and trees still showed relatively large diurnal ranges of water potential values and high soluble sugar contents, and did not shed leaves when experiencing the lowest midday water potentials during the dry period. They shed leaves only later in the dry winter period when even more serious drought was experienced. Their strategies seem to include both tolerance and avoidance mechanisms. Evergreen trees revealed relatively low water potentials with smallest diurnal range water potentials at the shrubland site, especially during the dry period, which indicated their weak ability to tolerate severe water stress. Increasing degradation of the vegetation clearly impacts negatively plant water relations. Using the ranges of leaf water potentials, the relative suitability of the plants for reforestation could be evaluated.     

Physiological response of tomatoes at drought,heat and their combination followed by recovery

In nature, crops encounter a combination of abiotic stresses that severely limit yield. Our aim was to dynamically expose the changes of tomatoes' physiological parameters to drought, heat and their combination and thereby clarify the relationship between the responses to single and combined stress. We studied the effect of single and combined drought and heat stresses on the shoot and root of two tomato cultivars (Sufen No.14 as CV1; Jinlingmeiyu as CV2). After being exposed to combined stress for 6 days, the dry weight of shoot and root significantly decreased. The F_q′/F_m′ (quantum yield of photosystem II) was significantly lower in CV1 upon drought and combined stress and in CV2 subjected to combined stress (between days 4 and 6) compared to control. The relative water content during combined stress was significantly lower than control from day 4 to recovery day 2. On days 3 and 6, the water loss rate significantly increased under heat stress and decreased at drought and combined stress, respectively. The combined stress caused severe damages on photosystem II and chloroplast ultrastructure. The root activity after stress recovered even though drought significantly increased the activity from day 2 to day 6. Combined stress result in complex responses during tomato growth. The CV1 was more heat tolerant than CV2, but there was no varietal difference at drought and combined stress. This study contributes to the understanding of the underlying physiological response mechanism of plant to combined stress and crop improvement by providing valuable information for abiotic stress‐tolerant tomato breeding.     

Regulation of citrus responses to the combined action of drought and high temperatures depends on the severity of water deprivation

Plants grown in natural environment are regularly subjected to different combinations of abiotic stresses. Recent studies revealed that citrus plants subjected to a combination of severe drought and high temperatures displayed specific physiological, hormonal, molecular and metabolic responses. In the present study, we have performed a long‐term experiment combining moderate drought and heat in Cleopatra mandarin to evaluate the impact of the stress‐sequence, intensity and duration. Our results support previous observation of high sensitivity of Cleopatra mandarin to abiotic stresses that include high temperatures. In this sense, a combination of drought and heat stress negatively impacts Cleopatra seedlings independently of the drought intensity. However, some responses to combined drought and heat depend on drought intensity, especially those involved in stomatal regulation. The intricate natural environment, abiotic stress combinations and global climatic changes increase the complexity of studying plant responses to stress factors in the laboratory. Consequently, new experimental approaches taking in consideration different stress combinations should be implemented to study the viability of Cleopatra mandarin as a rootstock in a rapidly changing environment.     

Plant responses to drought and rewatering

Plants would be more vulnerable to water stress and thereafter rewatering or a cycled water environmental change, which occur more frequently under climatic change conditions in terms of the prediction scenarios. Effects of water stress on plants alone have been well-documented in many reports. However, the combined responses to drought and rewatering and its mechanism are relatively scant. As we know, plant growth, photosynthesis and stomatal aperture may be limited under water deficit, which would be regulated by physical and chemical signals. Under severe drought, while peroxidation may be provoked, the relevant antioxidant metabolism would be involved to annihilate the damage of reactive oxygen species. As rewatering, the recoveries of plant growth and photosynthesis would appear immediately through growing new plant parts, re-opening the stomata, and decreasing peroxidation; the recovery extents (reversely: pre-drought limitation) due to rewatering strongly depend on pre-drought intensity, duration and species. Understanding how plants respond to episodic drought and watering pulse and the underlying mechanism is remarkably helpful to implement vegetation management practices in climatic changing.Key words: drought stress, peroxidation, photosynthesis, relative growth rate, pre-drought limitation, rewatering, signals, stomatal conductanceUnder the climatic changing context, drought has been, and is becoming an acute problem most constraining plant growth, terrestrial ecosystem productivity, in many regions all over the world, particularly in arid and semi-arid area.^1–3 Based on the fourth assessment report by IPCC, global surface average temperature will have a 1.1–6.4°C range increase by the end of this century.³ It is indicated that a warming above 3°C would eliminate thoroughly fixed carbon function of global terrestrial vegetation, shift a net carbon source. With global warming, it is expected that water deficit would be escalated by increasing evapotranspiration, increasing the frequency and intensity of drought with an increase from 1% to 30% in extreme drought land area by 2100;³ which would offset the beneficial effect from the elevated CO₂ concentration, further limiting the structure and function of the terrestrial ecosystem. The global climate models may forecast the precipitation regimes including its distribution and amount, but the complicated responses of terrestrial ecosystem to climate change may adversely affect the predict accuracy.^1,4Plant would response to water stress by dramatically complex mechanisms from genetic molecular express, biochemical metabolism through individual plant physiological processes to ecosystem levels^2,5,6 which may mainly includes six aspects: (1) drought escape via completing plant life cycle before severe water deficit. E.g., earlier flowering in annuals species before the onset of severe drough;⁷ (2) drought avoidance via enhancing capacity of getting water. E.g., developing root systems or conserving it such as reduction of stomata and leaf area/canopy cover;^8,9 (3) drought tolerance mainly via improving osmotic adjustment ability and increasing cell wall elasticity to maintain tissue turgidity;¹⁰ (4) drought resistance via altering metabolic path for life survives under severe stress (e.g., increased antioxidant metabolism);^11,12 (5) drought abandon by removing a part of individual, e.g., shedding elder leaves under water stress;² (6) drought-prone biochemical-physiological traits for plant evolution under long-term drought condition via genetic mutation and genetic modification.^13–15 The processes may be involved in multi-aspects simultaneously in responses of plants to drought stress and thereafter rewatering.In the field context, there is always interval occurrence in drought and/or rewetting events, particular under climatic change conditions predicting more frequent drought and flooding events.³ The water cycle change may greatly impact plant growth, photosynthesis and many key metabolic functions, thereby ecosystem productivity and agricultural achievement.^5,16–18 Actually, sporadic precipitation would become a critical issue for maintaining ecosystem structural stability and even it''s surviving in arid and semi-arid area. For example, a small rainfall pulse can induce a rapid response in a desert ecosystem, which quickly triggers plant growth so that the plants can survive.¹⁹ Thus, to highlight how plant and terrestrial ecosystem cope with adverse abnormal climatic change variables is, and always will be crucial research issue in practical management of plant growth and vegetation productivity. Here, we try to provide a brief insight into how plant responses to the pre-drought and rewatering in terms of the plant growth, gas exchange and key related-physiological processes such as reactive oxygen species (ROS) metabolism. Finally a regulation path schematic is presented to try to explain the involved processes.     

Can fungal endophytes fast-track plant adaptations to climate change?

Rapid climate change threatens plant communities. While many studies address the impact of climate change on plants and mechanisms of their resilience to climate stressors, the role of the plant microbiome in aiding plants' adaptation to climate change has been less investigated. We argue here that fungal endophytes, an important constituent of the plant microbiome, may be key to the ability of plants to adapt to climatic stressors. The rapid adaptive response of endophytes coupled with their ability to ‘transfer’ resistance to their hosts may fast-track plants' adaptation to climate change. We briefly review the importance of Class 3 fungal endophytes of terrestrial plants and discuss how they may accelerate adaptations to climate change in crops and natural plant communities and call for efforts directed at improving the understanding of fungal endophyte-facilitated plant health. Such information could aid in devising improved strategies for mitigating climate change effects on plant communities.     

Programmed proteome response for drought avoidance/tolerance in the root of a C(3) xerophyte (wild watermelon) under water deficits

Water availability is a critical determinant for the growth and ecological distribution of terrestrial plants. Although some xerophytes are unique regarding their highly developed root architecture and the successful adaptation to arid environments, virtually nothing is known about the molecular mechanisms underlying this adaptation. Here, we report physiological and molecular responses of wild watermelon (Citrullus lanatus sp.), which exhibits extraordinarily high drought resistance. At the early stage of drought stress, root development of wild watermelon was significantly enhanced compared with that of the irrigated plants, indicating the activation of a drought avoidance mechanism for absorbing water from deep soil layers. Consistent with this observation, comparative proteome analysis revealed that many proteins induced in the early stage of drought stress are involved in root morphogenesis and carbon/nitrogen metabolism, which may contribute to the drought avoidance via the enhancement of root growth. On the other hand, lignin synthesis-related proteins and molecular chaperones, which may function in the enhancement of physical desiccation tolerance and maintenance of protein integrity, respectively, were induced mostly at the later stage of drought stress. Our findings suggest that this xerophyte switches survival strategies from drought avoidance to drought tolerance during the progression of drought stress, by regulating its root proteome in a temporally programmed manner. This study provides new insights into the complex molecular networks within plant roots involved in the adaptation to adverse environments.     

木本植物水力系统对干旱胁迫的响应机制

干旱导致树木死亡对生态系统功能和碳平衡有重大影响。植物水分运输系统失调是引发树木死亡的主要机制。然而, 树木对干旱胁迫响应的多维性和复杂性, 使人们对植物水分运输系统在极端干旱条件下的响应以及植物死亡机理的认识还不清楚。该文首先评述衡量植物抗旱性的指标, 着重介绍可以综合评价植物干旱抗性特征的新参数——气孔安全阈值(SSM)。SSM越高, 表明气孔和水力性状之间的协调性越强, 木质部栓塞的可能性越低, 水力策略越保守。然后, 阐述木本植物应对干旱胁迫的一般响应过程。之后, 分别综述植物不同器官(叶、茎和根)对干旱胁迫的响应机制。植物达到死亡临界阈值的概率和时间, 取决于相关生理和形态学特征的相互作用。最后, 介绍木本植物水力恢复机制, 并提出3个亟待开展的研究问题: (1)改进叶片水分运输(木质部和木质部外水力导度)的测量方法, 量化4种不同途径的叶肉水分运输的相对贡献; (2)量化叶片表皮通透性变化, 以便更好地理解植物水分利用策略; (3)深入研究树木水碳耦合机制, 将个体结构和生理特征与群落/景观格局和过程相关联, 以便更好地评估和监测干旱诱导树木死亡的风险。     

植物应对干旱胁迫的阶段性策略

干旱是影响植物生存、生长和分布的最重要的非生物胁迫之一,全球暖干化将加剧干旱胁迫.植物对干旱胁迫的响应和适应机制一直是
学术研究的热点领域.本文综述了植物应对干旱胁迫的生长和生理响应,在已有的研究结果基础上,提出了植物应对干旱胁迫的阶段性响应策略.从干旱开始到干旱致死,植物经历了干旱开始-轻度干旱-中度干旱-严重干旱-极端干旱5个阶段,分别对应着应激响应-主动适应-被动适应3种响应方式和适应机制.不同阶段中植物抗旱机制的核心任务不同.最后提出了研究植物阶段性响应策略需要解决的关键科学问题及研究方向.