Differences in osmotic adjustment,foliar abscisic acid dynamics,and stomatal regulation between an isohydric and anisohydric woody angiosperm during drought

Species are often classified along a continuum from isohydric to anisohydric, with isohydric species exhibiting tighter regulation of leaf water potential through stomatal closure in response to drought. We investigated plasticity in stomatal regulation in an isohydric (Eucalyptus camaldulensis) and an anisohydric (Acacia aptaneura) angiosperm species subject to repeated drying cycles. We also assessed foliar abscisic acid (ABA) content dynamics, aboveground/belowground biomass allocation and nonstructural carbohydrates. The anisohydric species exhibited large plasticity in the turgor loss point (Ψ_TLP), with plants subject to repeated drying exhibiting lower Ψ_TLP and correspondingly larger stomatal conductance at low water potential, compared to plants not previously exposed to drought. The anisohydric species exhibited a switch from ABA to water potential‐driven stomatal closure during drought, a response previously only reported for anisohydric gymnosperms. The isohydric species showed little osmotic adjustment, with no evidence of switching to water potential‐driven stomatal closure, but did exhibit increased root:shoot ratios. There were no differences in carbohydrate depletion between species. We conclude that a large range in Ψ_TLP and biphasic ABA dynamics are indicative of anisohydric species, and these traits are associated with exposure to low minimum foliar water potential, dense sapwood and large resistance to xylem embolism.     

Water-use strategies of six co-existing Mediterranean woody species during a summer drought

Drought stress is known to limit plant performance in Mediterranean-type ecosystems. We have investigated the dynamics of the hydraulics, gas exchange and morphology of six co-existing Mediterranean woody species growing under natural field conditions during a drought that continued during the entire summer. Based on the observed minimum leaf water potentials, our results suggest that the six co-existing species cover a range of plant hydraulic strategies, from isohydric to anisohydric. These differences are remarkable since the selected individuals grow within several meters of each other, sharing the same environment. Surprisingly, whatever the leaf water potentials were at the end of the dry period, stomatal conductance, photosynthesis and transpiration rates were relatively similar and low across species. This result contradicts the classic view that anisohydric species are able to maintain gas exchange for longer periods of time during drought stress. None of the plants showed the expected structural acclimation response to the increasing drought (reduction of leaf-to-sapwood area ratio), thereby rejecting the functional equilibrium hypothesis for our study system. Instead, three of the six species increased photosynthetic area at the branch level. The observed dissimilar patterns of gas exchange, hydraulics and morphology across species seem to be equally successful given that photosynthesis at the leaf level was maintained at similar rates over the whole dry period.     

Water use strategy affects avoidance of ozone stress by stomatal closure in Mediterranean trees—A modelling analysis

Both ozone (O₃) and drought can limit carbon fixation by forest trees. To cope with drought stress, plants have isohydric or anisohydric water use strategies. Ozone enters plant tissues through stomata. Therefore, stomatal closure can be interpreted as avoidance to O₃ stress. Here, we applied an optimization model of stomata involving water, CO₂, and O₃ flux to test whether isohydric and anisohydric strategies may affect avoidance of O₃ stress by stomatal closure in four Mediterranean tree species during drought. The data suggest that stomatal closure represents a response to avoid damage to the photosynthetic mechanisms under elevated O₃ depending on plant water use strategy. Under high-O₃ and well-watered conditions, isohydric species limited O₃ fluxes by stomatal closure, whereas anisohydric species activated a tolerance response and did not actively close stomata. Under both O₃ and drought stress, however, anisohydric species enhanced the capacity of avoidance by closing stomata to cope with the severe oxidative stress. In the late growing season, regardless of the water use strategy, the efficiency of O₃ stress avoidance decreased with leaf ageing. As a result, carbon assimilation rate was decreased by O₃ while stomata did not close enough to limit transpirational water losses.     

Drought limitations to leaf‐level gas exchange: results from a model linking stomatal optimization and cohesion–tension theory

We merge concepts from stomatal optimization theory and cohesion–tension theory to examine the dynamics of three mechanisms that are potentially limiting to leaf‐level gas exchange in trees during drought: (1) a ‘demand limitation’ driven by an assumption of optimal stomatal functioning; (2) ‘hydraulic limitation’ of water movement from the roots to the leaves; and (3) ‘non‐stomatal’ limitations imposed by declining leaf water status within the leaf. Model results suggest that species‐specific ‘economics’ of stomatal behaviour may play an important role in differentiating species along the continuum of isohydric to anisohydric behaviour; specifically, we show that non‐stomatal and demand limitations may reduce stomatal conductance and increase leaf water potential, promoting wide safety margins characteristic of isohydric species. We used model results to develop a diagnostic framework to identify the most likely limiting mechanism to stomatal functioning during drought and showed that many of those features were commonly observed in field observations of tree water use dynamics. Direct comparisons of modelled and measured stomatal conductance further indicated that non‐stomatal and demand limitations reproduced observed patterns of tree water use well for an isohydric species but that a hydraulic limitation likely applies in the case of an anisohydric species.     

Global variations in ecosystem‐scale isohydricity

Droughts are expected to become more frequent and more intense under climate change. Plant mortality rates and biomass declines in response to drought depend on stomatal and xylem flow regulation. Plants operate on a continuum of xylem and stomatal regulation strategies from very isohydric (strict regulation) to very anisohydric. Coexisting species may display a variety of isohydricity behaviors. As such, it can be difficult to predict how to model the degree of isohydricity at the ecosystem scale by aggregating studies of individual species. This is nonetheless essential for accurate prediction of ecosystem drought resilience. In this study, we define a metric for the degree of isohydricity at the ecosystem scale in analogy with a recent metric introduced at the species level. Using data from the AMSR‐E satellite, this metric is evaluated globally based on diurnal variations in microwave vegetation optical depth (VOD), which is directly related to leaf water potential. Areas with low annual mean radiation are found to be more anisohydric. Except for evergreen broadleaf forests in the tropics, which are very isohydric, and croplands, which are very anisohydric, land cover type is a poor predictor of ecosystem isohydricity, in accordance with previous species‐scale observations. It is therefore also a poor basis for parameterizing water stress response in land‐surface models. For taller ecosystems, canopy height is correlated with higher isohydricity (so that rainforests are mostly isohydric). Highly anisohydric areas show either high or low underlying water use efficiency. In seasonally dry locations, most ecosystems display a more isohydric response (increased stomatal regulation) during the dry season. In several seasonally dry tropical forests, this trend is reversed, as dry‐season leaf‐out appears to coincide with a shift toward more anisohydric strategies. The metric developed in this study allows for detailed investigations of spatial and temporal variations in plant water behavior.     

Risk-taking plants: Anisohydric behavior as a stress-resistance trait

Water scarcity is a critical limitation for agricultural systems. Two different water management strategies have evolved in plants: an isohydric strategy and an anisohydric strategy. Isohydric plants maintain a constant midday leaf water potential (Ψ_leaf) when water is abundant, as well as under drought conditions, by reducing stomatal conductance as necessary to limit transpiration. Anisohydric plants have more variable Ψ_leaf and keep their stomata open and photosynthetic rates high for longer periods, even in the presence of decreasing leaf water potential. This risk-taking behavior of anisohydric plants might be beneficial when water is abundant, as well as under moderately stressful conditions. However, under conditions of intense drought, this behavior might endanger the plant. We will discuss the advantages and disadvantages of these two water-usage strategies and their effects on the plant’s ability to tolerate abiotic and biotic stress. The involvement of plant tonoplast AQPs in this process will also be discussed.     

Relationships between stomatal behavior,xylem vulnerability to cavitation and leaf water relations in two cultivars of Vitis vinifera

Current understanding of physiological mechanisms governing stomatal behavior under water stress conditions is still incomplete and controversial. It has been proposed that coordination of stomatal kinetics with xylem vulnerability to cavitation [vulnerability curve (VC)] leads to different levels of isohydry/anisohydry in different plant species/cultivars. In this study, this hypothesis is tested in Vitis vinifera cultivars displaying contrasting stomatal behavior under drought stress. The cv Montepulciano (MP, near‐isohydric) and Sangiovese (SG, anisohydric) were compared in terms of stomatal response to leaf and stem water potential, as possibly correlated to different petiole hydraulic conductivity (k_petiole) and VC, as well as to leaf water relations parameters. MP leaves showed almost complete stomatal closure at higher leaf and stem water potentials than SG leaves. Moreover, MP petioles had higher maximum k_petiole and were more vulnerable to cavitation than SG. Water potential at the turgor loss point was higher in MP than in SG. In SG, the percentage reduction of stomatal conductance (PLg_s) under water stress was almost linearly correlated with corresponding percentage loss of k_petiole (PLC), while in MP PLg_s was less influenced by PLC. Our results suggest that V. vinifera near‐isohydric and anisohydric genotypes differ in terms of xylem vulnerability to cavitation as well as in terms of k_petiole, and that the coordination of these traits leads to their different stomatal responses under water stress conditions.     

Trade-offs between plant leaf hydraulic and economic traits

Leaf is the most important organ for carbon-water coupling of a plant because it is the primary medium for photosynthesis. It also acts as the hydraulic bottleneck and safety valve against hydraulic catastrophic dysfunctions. The leaf economics spectrum, which reflects the balance between investments and returns of leaf economic traits, provides a useful framework for examining species strategies as shaped by their evolutionary history. Changes in leaf hydraulic traits will influence leaf economic traits as well as plant survival and growth. Exploring trade-offs between leaf hydraulic and economic traits is thus of significance for modeling carbon-water relations, understanding the mechanisms of water/carbon investments, and extending the leaf economic spectrum. In this review, we first examined the trade-offs between leaf hydraulic and economic traits. Specially, we analyzed the relationships between leaf hydraulic conductivity and hydraulic vulnerability, water potential at the turgor loss point, water capacitance, safety margin, and leaf morphological, structural and functional traits. We then discussed potential mechanisms regulating leaf hydraulic and economic traits from leaf morphology, anatomy, venation, and stomatal functions. Finally, we proposed future research to: (1) develop an integrated whole-plant economics spectrum, including carbon-nitrogen-water resources and root-stem-leaf hydraulic transport system that will help revealing ecophysiological mechanisms of plant structure-functional coupling, carbon sequestration and water use; (2) explore a generalized trade-offs among leaf hydraulic safety, hydraulic efficiency and carbon fixation efficiency to advance our understanding of the relationships between biophysical structure and physiological metabolism in plant leaf construction under drought stress; and (3) explore the carbon-water metabolic relationship and coupling of water transport and growth rate for the metabolic theory and predictions at community scale.     

Stomatal kinetics and photosynthetic gas exchange along a continuum of isohydric to anisohydric regulation of plant water status

Species' differences in the stringency of stomatal control of plant water potential represent a continuum of isohydric to anisohydric behaviours. However, little is known about how quasi‐steady‐state stomatal regulation of water potential may relate to dynamic behaviour of stomata and photosynthetic gas exchange in species operating at different positions along this continuum. Here, we evaluated kinetics of light‐induced stomatal opening, activation of photosynthesis and features of quasi‐steady‐state photosynthetic gas exchange in 10 woody species selected to represent different degrees of anisohydry. Based on a previously developed proxy for the degree of anisohydry, species' leaf water potentials at turgor loss, we found consistent trends in photosynthetic gas exchange traits across a spectrum of isohydry to anisohydry. More anisohydric species had faster kinetics of stomatal opening and activation of photosynthesis, and these kinetics were closely coordinated within species. Quasi‐steady‐state stomatal conductance and measures of photosynthetic capacity and performance were also greater in more anisohydric species. Intrinsic water‐use efficiency estimated from leaf gas exchange and stable carbon isotope ratios was lowest in the most anisohydric species. In comparisons between gas exchange traits, species rankings were highly consistent, leading to species‐independent scaling relationships over the range of isohydry to anisohydry observed.     

Aquaporins and plant transpiration

Although transpiration and aquaporins have long been identified as two key components influencing plant water status, it is only recently that their relations have been investigated in detail. The present review first examines the various facets of aquaporin function in stomatal guard cells and shows that it involves transport of water but also of other molecules such as carbon dioxide and hydrogen peroxide. At the whole plant level, changes in tissue hydraulics mediated by root and shoot aquaporins can indirectly impact plant transpiration. Recent studies also point to a feedback effect of transpiration on aquaporin function. These mechanisms may contribute to the difference between isohydric and anisohydric stomatal regulation of leaf water status. The contribution of aquaporins to transpiration control goes far beyond the issue of water transport during stomatal movements and involves emerging cellular and long‐distance signalling mechanisms which ultimately act on plant growth.     

Responses of two semiarid conifer tree species to reduced precipitation and warming reveal new perspectives for stomatal regulation

Relatively anisohydric species are predicted to be more predisposed to hydraulic failure than relatively isohydric species, as they operate with narrower hydraulic safety margins. We subjected co‐occurring anisohydric Juniperus monosperma and isohydric Pinus edulis trees to warming, reduced precipitation, or both, and measured their gas exchange and hydraulic responses. We found that reductions in stomatal conductance and assimilation by heat and drought were more frequent during relatively moist periods, but these effects were not exacerbated in the combined heat and drought treatment. Counter to expectations, both species exhibited similar g_s temporal dynamics in response to drought. Further, whereas P. edulis exhibited chronic embolism, J. monosperma showed very little embolism due to its conservative stomatal regulation and maintenance of xylem water potential above the embolism entry point. This tight stomatal control and low levels of embolism experienced by juniper refuted the notion that very low water potentials during drought are associated with loose stomatal control and with the hypothesis that anisohydric species are more prone to hydraulic failure than isohydric species. Because direct association of stomatal behaviour with embolism resistance can be misleading, we advocate consideration of stomatal behaviour relative to embolism resistance for classifying species drought response strategies.     

Strategies of a Bornean tropical rainforest water use as a function of rainfall regime: isohydric or anisohydric?

Although Bornean tropical rainforests are among the moistest biomes in the world, they sporadically experience periods of water stress. The observations indicate that these ecosystems tend to have little regulation of water use, despite episodes of relatively severe drought. This water-use behaviour is often referred to as anisohydric behaviour, as opposed to isohydric plants that regulate stomatal movement to prevent hydraulic failure. Although it is generally thought that anisohydric behaviour is an adaptation to more drought-prone habitats, we show that anisohydric plants may also be more favoured than isohydric plants under very moist environments where there is little risk of hydraulic failure. To explore this subject, we examined the advantages of isohydric and anisohydric species as a function of the hydroclimatic environment using a stochastic model of soil moisture and carbon assimilation dynamics parameterized by field observations. The results showed that under very moist conditions, anisohydric species tend to have higher productivity than isohydric plants, despite the fact that the two plant types show almost the same drought-induced mortality. As precipitation decreases, the mortality of anisohydric plants drastically increases whereas that of isohydric plants remains relatively constant and low; in these conditions, isohydric plants surpass anisohydric plants in their productivity.     

Differences in hydraulic architecture account for near-isohydric and anisohydric behaviour of two field-grown Vitis vinifera L. cultivars during drought

A comparative study on stomatal control under water deficit was conducted on grapevines of the cultivars Grenache, of Mediterranean origin, and Syrah of mesic origin, grown near Montpellier, France and Geisenheim, Germany. Syrah maintained similar maximum stomatal conductance (g_max) and maximum leaf photosynthesis (A_max) values than Grenache at lower predawn leaf water potentials, Ψ_leaf, throughout the season. The Ψ_leaf of Syrah decreased strongly during the day and was lower in stressed than in watered plants, showing anisohydric stomatal behaviour. In contrast, Grenache showed isohydric stomatal behaviour in which Ψ_leaf did not drop significantly below the minimum Ψ_leaf of watered plants. When g was plotted versus leaf specific hydraulic conductance, K_l, incorporating leaf transpiration rate and whole‐plant water potential gradients, previous differences between varieties disappeared both on a seasonal and diurnal scale. This suggested that isohydric and anisohydric behaviour could be regulated by hydraulic conductance. Pressure‐flow measurements on excised organs from plants not previously stressed revealed that Grenache had a two‐ to three‐fold larger hydraulic conductance per unit path length (K_h) and a four‐ to six‐fold larger leaf area specific conductivity (LSC) in leaf petioles than Syrah. Differences between internodes were only apparent for LSC and were much smaller. Cavitation detected as ultrasound acoustic emissions on air‐dried shoots showed higher rates for Grenache than Syrah during the early phases of the dry‐down. It is hypothesized that the differences in water‐conducting capacity of stems and especially petioles may be at the origin of the near‐isohydric and anisohydric behaviour of g.     

植物气孔气态失水与SPAC系统液态供水的相互调节作用研究进展

讨论了植物气孔气态失水与SPAC系统液态供水相互作用研究领域的一些重要现象和行为.当植物水力信号和化学信号共同作用促进气孔对叶水势的调节时,植物对叶水势的调节表现为等水行为.气孔对环境湿度变化响应的反馈机制可用来解释土壤干旱条件下气孔和光合的午休现象,以及气孔导度和水流导度之间的相关关系;而气孔对环境湿度变化响应的前馈机制,则可用来解释气孔导度对大气 叶片间水汽饱和差的滞后反应.植物最大限度地利用木质部传输水分的策略,要求气孔快速响应以避免木质部过度气穴化和短时间内将气穴逆转的相应机制.     

Coordination and trade‐offs between leaf and stem hydraulic traits and stomatal regulation along a spectrum of isohydry to anisohydry

The degree of plant iso/anisohydry, a widely used framework for classifying species‐specific hydraulic strategies, integrates multiple components of the whole‐plant hydraulic pathway. However, little is known about how it associates with coordination of functional and structural traits within and across different organs. We examined stem and leaf hydraulic capacitance and conductivity/conductance, stem xylem anatomical features, stomatal regulation of daily minimum leaf and stem water potential (Ψ), and the kinetics of stomatal responses to vapour pressure deficit (VPD) in six diverse woody species differing markedly in their degree of iso/anisohydry. At the stem level, more anisohydric species had higher wood density and lower native capacitance and conductivity. Like stems, leaves of more anisohydric species had lower hydraulic conductance; however, unlike stems, their leaves had higher native capacitance at their daily minimum values of leaf Ψ. Moreover, rates of VPD‐induced stomatal closure were related to intrinsic rather than native leaf capacitance and were not associated with species' degree of iso/anisohydry. Our results suggest a trade‐off between hydraulic storage and efficiency in the leaf, but a coordination between hydraulic storage and efficiency in the stem along a spectrum of plant iso/anisohydry.     

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.     

Regulation and acclimation of leaf gas exchange in a piñon–juniper woodland exposed to three different precipitation regimes

Leaf gas‐exchange regulation plays a central role in the ability of trees to survive drought, but forecasting the future response of gas exchange to prolonged drought is hampered by our lack of knowledge regarding potential acclimation. To investigate whether leaf gas‐exchange rates and sensitivity to drought acclimate to precipitation regimes, we measured the seasonal variations of leaf gas exchange in a mature piñon–juniper Pinus edulis–Juniperus monosperma woodland after 3 years of precipitation manipulation. We compared trees receiving ambient precipitation with those in an irrigated treatment (+30% of ambient precipitation) and a partial rainfall exclusion (?45%). Treatments significantly affected leaf water potential, stomatal conductance and photosynthesis for both isohydric piñon and anisohydric juniper. Leaf gas exchange acclimated to the precipitation regimes in both species. Maximum gas‐exchange rates under well‐watered conditions, leaf‐specific hydraulic conductance and leaf water potential at zero photosynthetic assimilation all decreased with decreasing precipitation. Despite their distinct drought resistance and stomatal regulation strategies, both species experienced hydraulic limitation on leaf gas exchange when precipitation decreased, leading to an intraspecific trade‐off between maximum photosynthetic assimilation and resistance of photosynthesis to drought. This response will be most detrimental to the carbon balance of piñon under predicted increases in aridity in the southwestern USA.     

Mechanisms of plant survival and mortality during drought: why do some plants survive while others succumb to drought?

Severe droughts have been associated with regional-scale forest mortality worldwide. Climate change is expected to exacerbate regional mortality events; however, prediction remains difficult because the physiological mechanisms underlying drought survival and mortality are poorly understood. We developed a hydraulically based theory considering carbon balance and insect resistance that allowed development and examination of hypotheses regarding survival and mortality. Multiple mechanisms may cause mortality during drought. A common mechanism for plants with isohydric regulation of water status results from avoidance of drought-induced hydraulic failure via stomatal closure, resulting in carbon starvation and a cascade of downstream effects such as reduced resistance to biotic agents. Mortality by hydraulic failure per se may occur for isohydric seedlings or trees near their maximum height. Although anisohydric plants are relatively drought-tolerant, they are predisposed to hydraulic failure because they operate with narrower hydraulic safety margins during drought. Elevated temperatures should exacerbate carbon starvation and hydraulic failure. Biotic agents may amplify and be amplified by drought-induced plant stress. Wet multidecadal climate oscillations may increase plant susceptibility to drought-induced mortality by stimulating shifts in hydraulic architecture, effectively predisposing plants to water stress. Climate warming and increased frequency of extreme events will probably cause increased regional mortality episodes. Isohydric and anisohydric water potential regulation may partition species between survival and mortality, and, as such, incorporating this hydraulic framework may be effective for modeling plant survival and mortality under future climate conditions.     

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.     

叶片结构的水力学特性对植物生理功能影响的研究进展

叶片是植物进行光合、呼吸、蒸腾作用的主要器官, 早期的研究主要集中于水分在叶片中的运输路径, 而对叶脉结构及其生态学意义研究甚少。近年来关于叶片叶脉结构、气孔结构的功能及叶片水力学特性的意义研究已经成为植物生理生态的研究热点。该文综述了叶脉的结构性状的指标(叶脉密度、直径、间距等), 叶片水力学结构特性对植物生长、水分运输、气体交换、光合作用等生理功能的影响, 及其与植物对干旱适应性之间的关系。叶脉结构是决定叶片生理功能的基础, 因此在未来的工作中应分析比较不同种类植物叶脉结构形态与导水、光合、呼吸、同化作用之间的关系, 建立植物茎干-枝-叶系统水力传导的机理性模型, 用以探索不同植物功能结构和高效用水生理生态学机制, 据此评估不同种类植物在未来气候情景下的地位。