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。     

干旱胁迫植物个体生理响应及其生态模型预测研究进展

全球气候突变导致干旱事件频发,进而易引发严重的植物衰退甚至死亡,聚焦植物尤其是树木死亡的生理学机制并期望基于此评估及预测气候变化导致植物死亡风险已成为热点话题。植物通过调整内在生理代谢过程,例如通过调节渗透物质的含量,来平衡渗透势、维持细胞膨压、调节植物激素的信号水平,诱导植物气孔开放程度降低,有利于植物保存水分、调控植物水通道蛋白的表达,进而保持体内水分稳定并对干旱胁迫做出快速响应。这些生理过程中的每一环调节都为了确保水分运输的效率和安全性,增加植物抗旱性以及生态系统稳定性。植物的抗旱性不仅体现在生理代谢方面的调节,还表现在植物水力特性与解剖结构间相辅相成。当植物改变水力特性的同时,其茎叶会在解剖结构上做出调整以满足植物在干旱环境下水分供需平衡,从而降低植物蒸腾水分散失、增强细胞储水并提高生存能力。植物应对水分胁迫的策略通常与水分消耗和碳获取之间的平衡有关,明晰植物水分消耗与光合碳获取间存在平衡关系的性状特征便于更好地理解植物的水分利用策略。然而,植物表现出的任意单一性状特征的强弱都无法代表整个植物适应逆境的优劣,未来只有通过将植物更多性状特征进行相互关联,以具有代表植物水力功能、结...     

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。     

高寒草甸植物群落功能多样性对不同生长期干旱的响应机制

植物对干旱胁迫的响应表现为各功能性状的差异化表达。全球气候变化下,青藏高原地区降水格局发生改变,高寒草甸群落功能性状及功能多样性在不同生长期干旱事件下的响应机制对加深高寒草甸适应气候变化认知具有重要意义。以藏北高寒草甸为研究对象,设置截雨棚模拟生长季前期（ED）、中期（MD）和非生长季时期（ND）干旱事件,通过观测群落物种功能性状,分析高寒草甸群落功能多样性对不同生长期干旱的响应机制。结果表明：（1）叶片功能性状对干旱存在差异响应,表现为叶片小而厚且寿命长,同化速率降低,并受氮元素限制加剧;（2）生长季前期干旱对高寒草甸群落功能性状的影响最强,生长季中期干旱次之,非生长季干旱的影响最弱;（3）生长季干旱处理显著改变了群落的功能多样性,ED处理下功能分散度指数（FDiv）和功能分异度指数（FDis）显著降低（P<0.05）,而Rao二次熵指数（RaoQ）显著升高（P<0.05）,MD处理下功能均匀度指数（FEve）显著降低（P<0.05）;（4）相关性分析得出,群落功能性状与功能多样性对干旱的响应之间存在着联系。本研究发现高寒草甸植物功能性状与群落功能多样性对生长季前期和中期干旱存在差异化响应,指示着高寒草甸植物群落在响应不同时期干旱时可能采取不同的生存策略,即对生长季前期干旱采用耐旱策略、对生长季中期干旱采用避旱策略。探讨了高寒草甸植物群落功能多样性对不同生长时期干旱胁迫的响应机制,为预测未来季节性干旱事件对青藏高原高寒草甸植物功能性状、群落特征和功能多样性的影响提供科学依据。     

木本植物木质部的冻融栓塞应对研究进展

冻融栓塞在中高纬度地区木本植物中普遍存在。抗冻融栓塞能力对在寒冷环境中木本植物的生长和安全越冬十分关键, 这直接决定植物分布范围。冻融栓塞是由于冰中气体溶解度低, 木质部水分在低温下冷冻, 使之前水中溶解的气体逸出到导管中, 随后木质部中的冰融化又使气泡扩张而引发的栓塞现象。木质部解剖结构的差异会影响植物的抗冻融栓塞能力, 植物还可以通过调节木质部正压、代谢耗能等方式主动修复冻融栓塞, 也可通过增加树液溶质含量等逃避冷冻, 以减少低温损伤。然而, 与干旱栓塞相比, 目前对木质部冻融栓塞的形成以及植物响应和调节机制的理解不足。为此, 该文首先综述了木质部冻融栓塞的形成机制和植物的逃避、忍耐、修复等3种冻融栓塞的应对策略, 然后总结了木质部抗低温胁迫能力的生理表现、影响因子和评价指标, 并在此基础上讨论了低温抗性、干旱抗性和水力效率之间的多元权衡关系, 最后提出今后该领域中的5个优先研究问题: (1)不同植物冰冻的最低温度阈值; (2)是否存在应对低温胁迫的水力脆弱性分割机制; (3)冻融栓塞修复与代谢消耗的关系; (4)低温抗性、干旱抗性和水力效率之间的权衡关系; (5)抗冻融栓塞性状是否能够纳入经济性状谱系。     

木本植物抗旱机理研究进展

干旱是主要的环境胁迫因子之一,严重影响植物的分布与生长发育。研究和探索旱生植物的抗旱机理已成为众多研究者关注的焦点。本文综述了部分抗旱木本植物根、茎、叶等与干旱环境相适应的结构特征,分析了干旱胁迫下,植物自身的渗透调节、抗氧化酶系统、内源激素变化、抗旱蛋白对干旱胁迫的响应机理,并概述了抗旱相关基因的研究进展。     

岩溶木本植物对干旱的生理生态适应

受全球气候变暖和季风气候影响,西南岩溶区年降水量及其在季节间的分配发生明显变化,无雨期频率和持续时间增加,且基岩风化严重,基质储水能力差,致使岩溶木本植物面临的季节性和地质性干旱加剧。该文通过参考相关文献分析结构性状和生理调节探讨岩溶木本植物如何适应地质性和季节性干旱。结果表明岩溶木本植物应对干旱的策略与其他干旱、半干旱区的植物大体一致,主要有抗旱和避旱两种策略:抗旱性植物一般具有比叶面积小、叶肉多汁、储水组织发达、细胞液浓度高等适应干旱的特征,可通过增加木材密度、增强木质部导管的抗栓塞性和提高水分利用效率以适应干旱; 避旱植物则可通过小而密的气孔和叶脉、发达的表皮毛、栅栏组织和维管束鞘等结构特征减少水分丧失,并可通过落叶、深根吸收深层水源和脱落酸(ABA)介导提早关闭气孔以适应干旱。虽然关于岩溶植物形态结构和生理调节对干旱适应机制的研究取得了一定进展,但仍然存在一些亟待解决的问题,例如:深入研究岩溶地区基岩水分状况及其对植物的贡献; 加强岩溶木本植物根系结构和生物量分配、树木构型及根际微生物与木本植物干旱适应的协同关系研究; 同时探索如何将岩溶植物生态适应研究成果应用于生产实践中,科学指导石漠化治理与生态修复。     

木本植物木质部的冻融栓塞应对研究进展

冻融栓塞在中高纬度地区木本植物中普遍存在。抗冻融栓塞能力对在寒冷环境中木本植物的生长和安全越冬十分关键,这直接决定植物分布范围。冻融栓塞是由于冰中气体溶解度低,木质部水分在低温下冷冻,使之前水中溶解的气体逸出到导管中,随后木质部中的冰融化又使气泡扩张而引发的栓塞现象。木质部解剖结构的差异会影响植物的抗冻融栓塞能力,植物还可以通过调节木质部正压、代谢耗能等方式主动修复冻融栓塞,也可通过增加树液溶质含量等逃避冷冻,以减少低温损伤。然而,与干旱栓塞相比,目前对木质部冻融栓塞的形成以及植物响应和调节机制的理解不足。为此,该文首先综述了木质部冻融栓塞的形成机制和植物的逃避、忍耐、修复等3种冻融栓塞的应对策略,然后总结了木质部抗低温胁迫能力的生理表现、影响因子和评价指标,并在此基础上讨论了低温抗性、干旱抗性和水力效率之间的多元权衡关系,最后提出今后该领域中的5个优先研究问题:(1)不同植物冰冻的最低温度阈值;(2)是否存在应对低温胁迫的水力脆弱性分割机制;(3)冻融栓塞修复与代谢消耗的关系;(4)低温抗性、干旱抗性和水力效率之间的权衡关系;(5)抗冻融栓塞性状是否能够纳入经济性状谱系。     

植物水分调节对策:等水与非等水行为

水分是植物存活、生长和分布过程中的必需资源,阐明植物对干旱的应对和调节机制,是植物生理生态学和全球变化生态学的重要研究命题。植物对不同气候与土壤水分条件的长期适应会形成由一整套相关联的性状组成的水分调节策略,其中等水和非等水调节行为是两种典型的水分调节对策。区分并阐明植物的水分调节对策及其机制,不但在干旱地区植物育种、植被修复等实践中有广泛的应用前景,而且可为构建更精确的植被动态模型和预测气候变化情景下植被分布提供科学基础。该文首先阐述了等水和非等水调节行为的定义及3种定量分类方法:(1)基于气孔导度与叶水势的关系;(2)基于气孔导度与水汽压亏缺的关系;(3)基于黎明前叶水势与中午叶水势的关系。之后,从水力和碳经济性状两个方面比较分析了两种水分调节对策植物的种间差异。综合分析植物水分调节机制发现,水力信号与化学信号的相互作用是植物水分调节行为的主控因素。最后提出3个亟待开展研究的问题:(1)针对不同地区开展植物水分关系相关性状的测定,寻求可靠且普适的植物水分调节对策分类方法。(2)探索植物水分调节对策与水力、形态、结构、功能等性状之间的关联性,为改进植被动态模型提供可靠的参数。(3)加深理解不同时空尺度上植物水分调节过程,揭示植物对环境胁迫(尤其是干旱)的响应和适应机制。     

干旱胁迫对连翘幼苗非结构性碳分配和水力特性的影响

为了探究长期干旱胁迫下连翘不同器官的非结构性碳水化合物(NSC)含量与水力特性的协调及响应机制。以连续3年不同水分条件处理后的连翘幼苗为研究对象,设置3个水分处理(适宜供水、中度干旱胁迫和重度干旱胁迫),研究长期干旱胁迫后连翘幼苗的光合特性、生物量的分配、NSC各组分含量、水力特性的变化及其碳水两者之间的相关关系。结果表明:(1)适宜供水、中度干旱、重度干旱胁迫下,枝条的栓塞程度分别为30.7%、41.8%和42.3%,枝条导水率分别为0.95、0.71、0.65 kg m^-1 s^-1 MPa^-1。(2)重度干旱胁迫显著降低了净光合速率、蒸腾速率、气孔导度、水分利用效率。(3)重度干旱胁迫导致地上和粗根生物量显著降低,细根生物量和根冠比显著增加。此外,各器官的NSC含量显著降低,其根系NSC消耗量最高,根系的可溶性总糖和淀粉含量显著降低,枝条的可溶性总糖、葡萄糖和蔗糖含量增加了12.9%、31.1%和45.7%,而淀粉含量降低了40.7%。(4)枝条栓塞程度和导水率与可溶性总糖、淀粉、蔗糖和葡萄糖含量显著相关,其栓塞程度与可溶性总糖、葡萄糖和蔗糖呈正相关,而与淀粉呈负相关(P ＜ 0.01)。综上所述,干旱导致连翘枝条木质部的栓塞程度增加,导水率、光合作用和水分运输效率均显著降低,但连翘通过提高枝条内可溶性总糖、葡萄糖、蔗糖含量和降低淀粉、NSC含量以提高植物在干旱条件下的存活机率及旱后水分恢复能力,研究为半干旱区连翘培育和经营提供理论依据。     

不同水分条件下两个树种木质部栓塞对P素添加的响应

 在两种水分供给(干旱胁迫和适宜水分,土壤含水量分别为田间持水量的30%～40%和70%～80%)下,研究了耐旱树种元宝枫(Acer truncatum)和 中生树种女贞(Ligustrum lucidum )木质部栓塞(以导水率(Percentage loss of hydraulic conductivity, PLC)损失程度衡量)对P素添加的 响应。结果发现,两个树种PLC的日变化均呈现出先上升后降低的规律,表明木质部栓塞的形成与恢复是植物体的一种平常事件;除适宜水分条 件的女贞外,P素可以显著提高元宝枫和遭受干旱胁迫时女贞的PLC;两种水分条件下,干旱胁迫时元宝枫木质部栓塞明显高于适宜水分供给时 。女贞的PLC在两种水分状况下无显著差异;树种间,干旱胁迫促进了元宝枫木质部的栓塞形成,明显高于同等水分条件下的女贞。该研究结果 证实了“木质部限流耐旱假设”。     

荒漠河岸林植物木质部导水与栓塞特征及其对干旱胁迫的响应

 以同处于干旱区的塔里木河下游(铁干里克)和黑河下游(乌兰图格)断面为研究区, 比较了荒漠河岸林主要建群种胡杨(Populus euphratica)、柽柳(Tamarix spp.)、疏叶骆驼刺(Alhagi sparsifolia)和花花柴(Karelinia caspia)在长期遭受不同干旱胁迫下的根、枝条木质部导水力和栓塞化程度的变化特征, 并分析了木质部导水对干旱胁迫的响应及适应策略。结果表明: 1) 黑河下游荒漠河岸林植物的导水能力显著高于塔里木河下游, 其中柽柳、胡杨、疏叶骆驼刺和花花柴根木质部的初始比导率(K_s0)分别高11.97、6.74、7.10和3.73倍, 枝条的K_s0分别高9.48、3.65、2.07和1.88倍, 地下水埋深导致的干旱胁迫程度不同是诱发荒漠植物导水能力差异的根本原因; 2)柽柳耐干旱能力最强, 适应范围较宽, 而花花柴、疏叶骆驼刺的耐旱性相对较弱, 适生范围较窄, 这可能与植物的根系分布有关; 3)干旱胁迫较轻时, 枝条木质部是荒漠河岸林植物水分传输的主要阻力部位, 干旱胁迫严重时, 根木质部是限制植株水流的最大阻碍部位; 4)荒漠河岸林植物主要通过调节枝条木质部的水流阻力来适应干旱胁迫, 且其适应策略与干旱胁迫程度有关, 干旱胁迫轻时, 植物通过限制枝条木质部水流来协调整株植物的均匀生长; 干旱胁迫严重时, 植物通过牺牲劣势枝条、增强优势枝条水流来提高植株整体生存的机会。     

Plant resistance to drought depends on timely stomatal closure

Stomata play a significant role in the Earth's water and carbon cycles, by regulating gaseous exchanges between the plant and the atmosphere. Under drought conditions, stomatal control of transpiration has long been thought to be closely coordinated with the decrease in hydraulic capacity (hydraulic failure due to xylem embolism). We tested this hypothesis by coupling a meta‐analysis of functional traits related to the stomatal response to drought and embolism resistance with simulations from a soil–plant hydraulic model. We report here a previously unreported phenomenon: the existence of an absolute limit by which stomata closure must occur to avoid rapid death in drought conditions. The water potential causing stomatal closure and the xylem pressure at the onset of embolism formation were equal for only a small number of species, and the difference between these two traits (i.e. safety margins) increased continuously with increasing embolism resistance. Our findings demonstrate the need to revise current views about the functional coordination between stomata and hydraulic traits and provide a mechanistic framework for modeling plant mortality under drought conditions.     

Ecological relevance of minimum seasonal water potentials

The minimum seasonal water potential a plant experiences, ψ_min, provides an important measure of plant water status, as it reflects the maximum water deficit that leaves and xylem must tolerate to maintain physiological activity. ψ_min also acts as a selective force on xylem structure which, in turn, generates correlations between ψ_min and numerous hydraulic traits. This review focuses on the ecological relevance of ψ_min as a reflection of overall plant hydraulic strategy. The focus is on plant functional strategies with respect to soil drought, but we conclude with preliminary findings on the role of atmospheric drought.     

Reproductive water supply is prioritized during drought in tomato

Reproductive success largely defines the fitness of plant species. Understanding how heat and drought affect plant reproduction is thus key to predicting future plant fitness under rising global temperatures. Recent work suggests reproductive tissues are highly vulnerable to water stress in perennial plants where reproductive sacrifice could preserve plant survival. However, most crop species are annuals where such a strategy would theoretically reduce fitness. We examined the reproductive strategy of tomato (Solanum lycopersicum var. Rheinlands Ruhm) to determine whether water supply to fruits is prioritized above vegetative tissues during drought. Using optical methods, we mapped xylem cavitation and tissue shrinkage in vegetative and reproductive organs during dehydration to determine the priority of water flow under acute water stress. Stems and peduncles of tomato showed significantly greater xylem cavitation resistance than leaves. This maintenance of intact water supply enabled tomato fruit to continue to expand during acute water stress, utilizing xylem water made available by tissue collapse and early cavitation of leaves. Here, tomato plants prioritize water supply to reproductive tissues, maintaining fruit development under drought conditions. These results emphasize the critical role of water transport in shaping life history and suggest a broad relevance of hydraulic prioritization in plant ecology.     

A global analysis of plant recovery performance from water stress

Plant post‐drought recovery performance is essential to predict shifts in ecosystem dynamics and production during frequent climate change‐driven drought events. Yet, it is not clear how post‐drought recovery is related to evolutionary and geographic variations in plants. In this study, we generated a global data set of post‐drought recovery performance in 140 plant species from published studies. We quantified the plant post‐drought recovery performance by calculating a recovery index for multiple plant physiological and hydraulic parameters, including leaf water potential, net photosynthetic rate, leaf hydraulic conductance and shoot biomass. The magnitude of recovery among four plant functional types (deciduous angiosperms, evergreen angiosperms, gymnosperms, and crops), two plant growth forms (shrubs and trees), two water management strategies (isohydric and anisohydric), four xylem porosity types (diffuse, ring, semi‐ring and tracheid), and four major biomes (dry sclerophyll forest, boreal forest, temperate forest and tropical/subtropical forest) were compared. We found the inability to completely recover immediately after severe water stress is ubiquitous across all plant functional types and growth forms, while the rate and magnitude of post‐drought recovery varied greatly across different plant taxonomic categories and geographic ranges. In general, plant hydraulic architecture, leaf anatomy and physiology affect plants’ propensity towards recovery, and reflect evolutionary consequences of plant adaptation to their habitat. Due to the essential role of plant functional traits in regulating carbon storage in each biome, a better understanding plant post‐drought recovery performance could improve our predictions on ecosystem productivity in a rapidly changing climate.     

Aridity-dependent sequence of water potentials for stomatal closure and hydraulic dysfunctions in woody plants

The sequence of physiological events during drought strongly impacts plants' overall performance. Here, we synthesized the global data of stomatal and hydraulic traits in leaves and stems of 202 woody species to evaluate variations in the water potentials for key physiological events and their sequence along the climatic gradient. We found that the seasonal minimum water potential, turgor loss point, stomatal closure point, and leaf and stem xylem vulnerability to embolism were intercorrelated and decreased with aridity, indicating that water stress drives trait co-selection. In xeric regions, the seasonal minimum water potential occurred at lower water potential than turgor loss point, and the subsequent stomatal closure delayed embolism formation. In mesic regions, however, the seasonal minimum water potential did not pose a threat to the physiological functions, and stomatal closure occurred even at slightly more negative water potential than embolism. Our study demonstrates that the sequence of water potentials for physiological dysfunctions of woody plants varies with aridity, that is, xeric species adopt a more conservative sequence to prevent severe tissue damage through tighter stomatal regulation (isohydric strategy) and higher embolism resistance, while mesic species adopt a riskier sequence via looser stomatal regulation (anisohydric strategy) to maximize carbon uptake at the cost of hydraulic safety. Integrating both aridity-dependent sequence of water potentials for physiological dysfunctions and gap between these key traits into the hydraulic framework of process-based vegetation models would improve the prediction of woody plants' responses to drought under global climate change.     

Co-ordination between xylem anatomy,plant architecture and leaf functional traits in response to abiotic and biotic drivers in a nurse cushion plant

Background and AimsPlants in dry Mediterranean mountains experience a double climatic stress: at low elevations, high temperatures coincide with water shortage during summer, while at high elevations temperature decreases and water availability increases. Cushion plants often act as nurses by improving the microclimate underneath their canopies, hosting beneficiary species that may reciprocally modify their benefactors’ microenvironment. We assess how the nurse cushion plant Arenaria tetraquetra subsp. amabilis adjusts its hydraulic system to face these complex abiotic and biotic constraints.MethodsWe evaluated intra-specific variation and co-ordination of stem xylem anatomy, leaf functional traits and plant architecture in response to elevation, aspect and the presence of beneficiary species in four A. tetraquetra subsp. amabilis populations in the Sierra Nevada mountains, southern Spain.Key ResultsXylem anatomical and plant architectural traits were the most responsive to environmental conditions, showing the highest mutual co-ordination. Cushions were more compact and had smaller, more isolated conductive vessels in the southern than in the northern aspect, which allow minimization of the negative impacts of more intense drought. Only vessel size, leaf mass per area and terminal branch length varied with elevation. Nurse cushions co-ordinated plant architecture and xylem traits, having higher canopy compactness, fewer leaves per branch and fewer, more isolated vessels than non-nurse cushions, which reflects the negative effects of beneficiary plants on nurse water status. In non-nurse cushions, plant architecture co-ordinated with leaf traits instead. The interacting effects of aspect and elevation on xylem traits showed that stress due to frost at high elevation constrained xylem anatomy in the north, whereas stress due to drought had a parallel effect in the south.ConclusionsTrait co-ordination was weaker under more demanding environmental conditions, which agrees with the hypothesis that trait independence allows plants to better optimize different functions, probably entailing higher adjustment potential against future environmental changes.