Ecophysiological responses of two closely related Magnoliaceae genera to seasonal changes in subtropical China

Aims Plants use a variety of hydraulic strategies to adapt to seasonal drought that differ by species and environmental conditions. The early-diverging Magnoliaceae family includes two closely related genera with contrasting leaf habits, Yulania (deciduous) and Michelia (evergreen), which naturally inhabit temperate and tropical regions, respectively. Here, we evaluate the hydraulic strategy of species from both genera that have been ex situ conserved in a subtropical region to determine how they respond to the novel cool–dry season climatic pattern.Methods We measured ecophysiological traits in five Michelia and five Yulania species conserved in the South China Botanical Garden in both wet and dry season conditions and monitored the whole-year sap flow for four of these species.Important findings We found that Magnoliaceae species that have been ex situ conserved in a subtropical climate did not suffer from excessive water stress due to the mild drought conditions of the dry season and the ecophysiological adjustments the species made to avoid this stress, which differed by leaf habit. Specifically, deciduous species completely shed their leaves during the dry season, while evergreen species decreased their turgor loss points, dry mass based photosynthetic rates, stomatal conductance and specific leaf areas (SLAs) compared to wet season measurements. In comparing the two distinct leaf habits during the wet season, the leathery-leaved evergreen species had higher leaf hydraulic conductance and leaf to sapwood area ratios than the papery-leaved deciduous species, while the deciduous species had greater hydraulic conductivity calculated on both a stem and leaf area basis, dry mass based photosynthetic rates, leaf nutrients, SLAs and stomatal sizes than the evergreen species. Interestingly, species from both genera maintained similar sap flow in the wet season. Both photosynthetically active radiation and vapour pressure deficit affected the diurnal patterns of sap flow in the wet season, while only vapour pressure deficit played a dominant role in the dry season. This study reveals contrasting hydraulic strategies in Yulania and Michelia species under subtropical seasonal conditions, and suggests that these ecophysiological adjustments might be affected more by leaf habit than seasonality, thus reflecting the divergent evolution of the two closely related genera. Furthermore, we show that Magnoliaceae species that are ex situ conserved in a subtropical climate are hydraulically sound, a finding that will inform future conservation efforts of this ancient family under the threat of climatic change.     

Correlated evolution of stem and leaf hydraulic traits in Pereskia (Cactaceae)

Recent studies have demonstrated significant correlations between stem and leaf hydraulic properties when comparing across species within ecological communities. This implies that these traits are co-evolving, but there have been few studies addressing plant water relations within an explicitly evolutionary framework. This study tests for correlated evolution among a suite of plant water-use traits and environmental parameters in seven species of Pereskia (Cactaceae), using phylogenetically independent contrasts. There were significant evolutionary correlations between leaf-specific xylem hydraulic conductivity, Huber Value, leaf stomatal pore index, leaf venation density and leaf size, but none of these traits appeared to be correlated with environmental water availability; only two water relations traits - mid-day leaf water potentials and photosynthetic water use efficiency - correlated with estimates of moisture regime. In Pereskia, it appears that many stem and leaf hydraulic properties thought to be critical to whole-plant water use have not evolved in response to habitat shifts in water availability. This may be because of the extremely conservative stomatal behavior and particular rooting strategy demonstrated by all Pereskia species investigated. These results highlight the need for a lineage-based approach to understand the relative roles of functional traits in ecological adaptation.     

Leaf morphology of 40 evergreen and deciduous broadleaved subtropical tree species and relationships to functional ecophysiological traits

We explored potential of morphological and anatomical leaf traits for predicting ecophysiological key functions in subtropical trees. We asked whether the ecophysiological parameters stomatal conductance and xylem cavitation vulnerability could be predicted from microscopy leaf traits. We investigated 21 deciduous and 19 evergreen subtropical tree species, using individuals of the same age and from the same environment in the Biodiversity‐Ecosystem Functioning experiment at Jiangxi (BEF‐China). Information‐theoretic linear model selection was used to identify the best combination of morphological and anatomical predictors for ecophysiological functions. Leaf anatomy and morphology strongly depended on leaf habit. Evergreen species tended to have thicker leaves, thicker spongy and palisade mesophyll, more palisade mesophyll layers and a thicker subepidermis. Over 50% of all evergreen species had leaves with multi‐layered palisade parenchyma, while only one deciduous species (Koelreuteria bipinnata) had this. Interactions with leaf habit were also included in best multi‐predictor models for stomatal conductance (g_s) and xylem cavitation vulnerability. In addition, maximum g_s was positively related to log ratio of palisade to spongy mesophyll thickness. Vapour pressure deficit (vpd) for maximum g_s increased with the log ratio of palisade to spongy mesophyll thickness in species having leaves with papillae. In contrast, maximum specific hydraulic conductivity and xylem pressure at which 50% loss of maximum specific xylem hydraulic conductivity occurred (Ψ₅₀) were best predicted by leaf habit and density of spongy parenchyma. Evergreen species had lower Ψ₅₀ values and lower maximum xylem hydraulic conductivities. As hydraulic leaf and wood characteristics were reflected in structural leaf traits, there is high potential for identifying further linkages between morphological and anatomical leaf traits and ecophysiological responses.     

Genetic variation of ecophysiological traits in two gynodioecious species of Schiedea (Caryophyllaceae)

Evolution of dimorphic breeding systems may involve changes in ecophysiological traits as well as floral morphology because of greater resource demands on females. Differences between related species suggest that ecophysiological traits should be heritable, and species with higher female frequencies should show greater sexual differentiation. We used modified partial diallel crossing designs to estimate narrow-sense heritabilities and genetic correlations of sex-specific ecophysiological and morphological traits in closely related gynodioecious Schiedea salicaria (13% females) and Schiedea adamantis (39% females). In S. salicaria, hermaphrodites and females differed in photosynthetic rate and specific leaf area (SLA). Narrow-sense heritabilities were significant for stomatal conductance, SLA and inflorescence number in hermaphrodites, and for SLA and inflorescence number in females. Schiedea adamantis had no sexual dimorphism in measured traits; stomatal conductance, stem number and inflorescence number were heritable in females, and stem number was heritable in hermaphrodites. In both species, significant genetic correlations of traits between sexes were rare, indicating that traits can evolve independently in response to sex-differential selection. Significant genetic correlations were detected between certain traits within sexes of both species. Low heritability of some ecophysiological traits may reflect low additive genetic variability or high phenotypic plasticity in these traits.     

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.     

Are lianas more drought-tolerant than trees? A test for the role of hydraulic architecture and other stem and leaf traits

Lianas are an important component of Neotropical forests, where evidence suggests that they are increasing in abundance and biomass. Lianas are especially abundant in seasonally dry tropical forests, and as such it has been hypothesized that they are better adapted to drought, or that they are at an advantage under the higher light conditions in these forests. However, the physiological and morphological characteristics that allow lianas to capitalize more on seasonal forest conditions compared to trees are poorly understood. Here, we evaluate how saplings of 21 tree and liana species from a seasonal tropical forest in Panama differ in cavitation resistance (P ₅₀) and maximum hydraulic conductivity (K _h), and how saplings of 24 tree and liana species differ in four photosynthetic leaf traits (e.g., maximum assimilation and stomatal conductance) and six morphological leaf and stem traits (e.g., wood density, maximum vessel length, and specific leaf area). At the sapling stage, lianas had a lower cavitation resistance than trees, implying lower drought tolerance, and they tended to have a higher potential hydraulic conductivity. In contrast to studies focusing on adult trees and lianas, we found no clear differences in morphological and photosynthetic traits between the life forms. Possibly, lianas and trees are functionally different at later ontogenetic stages, with lianas having deeper root systems than trees, or experience their main growth advantage during wet periods, when they are less vulnerable to cavitation and can achieve high conductivity. This study shows, however, that the hydraulic characteristics and functional traits that we examined do not explain differences in liana and tree distributions in seasonal forests.     

The effects of intervessel pit characteristics on xylem hydraulic efficiency and photosynthesis in hemiepiphytic and non‐hemiepiphytic Ficus species

Xylem vulnerability to cavitation and hydraulic efficiency are directly linked to fine‐scale bordered pit features in water‐conducting cells of vascular plants. However, it is unclear how pit characteristics influence water transport and carbon economy in tropical species. The primary aim of this study was to evaluate functional implications of changes in pit characteristics for water relations and photosynthetic traits in tropical Ficus species with different growth forms (i.e. hemiepiphytic and non‐hemiepiphytic) grown under common conditions. Intervessel pit characteristics were measured using scanning electron microscopy in five hemiepiphytic and five non‐hemiepiphytic Ficus species to determine whether these traits were related to hydraulics, leaf photosynthesis, stomatal conductance and wood density. Ficus species varied greatly in intervessel pit structure, hydraulic conductivity and leaf physiology, and clear differences were observed between the two growth forms. The area and diameter of pit aperture were negatively correlated with sapwood‐specific hydraulic conductivity, mass‐based net assimilation rate, stomatal conductance (g_s), intercellular CO₂ concentration (C_i) and the petiole vessel lumen diameters (D_v), but positively correlated with wood density. Pit morphology was only negatively correlated with sapwood‐ and leaf‐specific hydraulic conductivity and D_v. Pit density was positively correlated with g_s, C_i and D_v, but negatively with intrinsic leaf water‐use efficiency. Pit and pit aperture shape were not significantly correlated with any of the physiological traits. These findings indicate a significant role of pit characteristics in xylem water transport, carbon assimilation and ecophysiological adaptation of Ficus species in tropical rain forests.     

陕北水蚀风蚀交错带柠条适应不同生境的形态和生理可塑性

以陕北水蚀风蚀交错带4个不同坡位与土壤质地生境[沟底+坝淤绵沙土(A)、梁坡+红黄土(B)、坡顶+绵沙土(C)、坡顶+风沙土盖绵沙土(D)]中生长的柠条为研究对象,研究了其生长、光合和水力性状的可塑性变化,以揭示柠条对不同生境的适应机制。结果表明:(1)生境D和生境A样地1~3m和3m土层的平均含水量明显高于生境B和C;(2)与生境B和C相比,生境A和D中生长的柠条冠幅、株高和新枝长显著增加,叶厚度减小,正午叶水势、净光合速率、气孔导度和蒸腾速率亦显著增加,但不同生境中柠条枝比导水率、Huber值和栓塞程度无显著差异;(3)柠条各测定指标中,新枝长、净光合速率、气孔导度和蒸腾速率表现出较大的可塑性。研究认为,柠条可能主要通过新枝长或光合生理特征的改变来适应不同的水分生境。     

Hydraulic and photosynthetic characteristics differ between co-generic tree and liana species: a case study of Millettia and Gnetum in tropical forest

木质藤本是热带森林的重要组成部分, 显著影响森林的结构和功能。已有研究发现木质藤本与乔木的水力结构存在显著差异: 木质藤本的缠绕或攀缘茎细小, 但其木质部具有粗大的长导管, 输水效率高, 抗栓塞能力低。为降低基因型差异对比较结果的影响, 该研究选取热带崖豆藤属(Millettia)和买麻藤属(Gnetum)的乔木和木质藤本, 比较同属内不同生长型植物的水力和光合性状的差异, 分析水分传导效率与抗栓塞能力之间以及水力与光合性状之间的相关关系。结果发现: (1)崖豆藤属植物水力性状的种间差异大, 与生活型和需光性有关。耐阴的木质藤本反而具有较低的水分传导效率和较高的抗栓塞能力。(2)买麻藤属植物是裸子植物较为进化的类群(具有导管和阔叶), 其乔木的水分传导效率很低, 但是其木质藤本的水分传导效率高于其他阳生性的被子植物。(3)不论乔木还是木质藤本, 水分传导的有效性与安全性在枝条和叶片水平上均没有显著的权衡关系。(4)与同属乔木相比, 木质藤本的叶片较枝条的抗栓塞能力更强, 在旱季具有更高的最大净光合速率和气孔导度, 支持了木质藤本的“旱季生长优势假说”。该研究揭示了热带木质藤本水力性状的多样性和重要性, 为阐明环境变化对这一重要植物类群的影响, 需要对它们的水力特征进行更广泛的研究。     

木兰科常绿与落叶物种叶片构建策略的差异

为探究木兰科(Magnoliaceae)常绿与落叶物种叶片构建的生理生态策略,选取黄山木兰(Yulaniacylindrica)、玉兰(Y.denudata)和鸡公山玉兰(Y. jigongshanensis) 3种落叶物种,以及荷花玉兰(Magnolia grandiflora)、含笑花(Michelia figo)、石碌含笑(M. shiluensis) 3种常绿物种,对其叶片构建成本和叶片寿命相关的性状进行比较。结果表明,木兰科3落叶种的单位叶片面积成本(CCarea)显著低于3常绿种,但落叶和常绿物种的叶片质量成本(CCmass)差异不显著。落叶物种的叶氮、磷含量(Nmass,Pmass)和比叶面积(SLA)均显著高于常绿物种,而叶片寿命(LLS)显著低于常绿物种。CCarea与LLS呈显著正相关,Nmass、Pmass和SLA均与LLS呈显著负相关。这说明木兰科玉兰属落叶物种单位面积叶片构建成本小于常绿物种;落叶物种叶片寿命短,但采取低成本构建策略,提高比叶面积获得更多光资源,增加营养积累,也揭示了玉兰属落叶物种适应北亚热带较短的生长季和较低水热条件的生理生态策略。     

Ecophysiological significance of leaf traits in Cypripedium and Paphiopedilum

There is a close phylogenetic relationship between Paphiopedilum and Cypripedium, but these two genera diverge considerably in terms of their leaf traits. To understand the evolution and the ecophysiological significance of leaf traits, we investigated the leaf traits of three Paphiopedilum species and three Cypripedium species in southwestern China. Cypripedium tibeticum and C. flavum showed a significantly higher light-saturated photosynthetic rate (P(max) ), stomatal conductance (g(s) ), photosynthetic nitrogen utilization efficiency (PNUE) and specific leaf area (SLA), but lower ratio of leaf carbon to nitrogen content (C/N) and leaf construction cost (CC) than Paphiopedilum. These leaf traits of Cypripedium suggest its high resource use efficiency and high growth rate reflecting adaptation to a short growing period and abundant soil nutrients and water in alpine habitats. Conversely, the low P(max) , g(s) , PNUE, SLA and the ratio of chlorophyll a to chlorophyll b (Chl a/b), but high leaf nitrogen investment in light-harvesting component (P(L) ), CC and C/N in Paphiopedilum indicate its adaptation to a low light, nutrient-poor and limited soil water habitats in karst areas. As a sympatric species of Paphiopedilum, although C. lentiginosum retained the phylogenetic leaf traits of Cypripedium, such as high mass-based light-saturated photosynthetic rate (P(max-M) ), g(s) and PNUE, it had some similar leaf traits to those of Paphiopedilum, such as low mesophyll conductance (g(m) ) and Chl a/b, and high P(L) , which reflected an adaptation to the same habitat. Our results show that the evolution of the leaf traits of Paphiopedilum and Cypripedium are shaped by both phylogeny and environment.     

Leaf traits in parental and hybrid species of Sorbus (Rosaceae)

? Premise of the study: Knowledge of functional leaf traits can provide important insights into the processes structuring plant communities. In the genus Sorbus, the generation of taxonomic novelty through reticulate evolution that gives rise to new microspecies is believed to be driven primarily by a series of interspecific hybridizations among closely related taxa. We tested hypotheses for dispersion of intermediacy across the leaf traits in Sorbus hybrids and for trait linkages with leaf area and specific leaf area. ? Methods: Here, we measured and compared the whole complex of growth, vascular, and ecophysiological leaf traits among parental (Sorbus aria, Sorbus aucuparia, Sorbus chamaemespilus) and natural hybrid (Sorbus montisalpae, Sorbus zuzanae) species growing under field conditions. A recently developed atomic force microscopy technique, PeakForce quantitative nanomechanical mapping, was used to characterize the topography of cell wall surfaces of tracheary elements and to map the reduced Young's modulus of elasticity. ? Key results: Intermediacy was associated predominantly with leaf growth traits, whereas vascular and ecophysiological traits were mainly parental-like and transgressive phenotypes. Larger-leaf species tended to have lower modulus of elasticity values for midrib tracheary element cell walls. Leaves with a biomass investment related to a higher specific leaf area had a lower density. Leaf area- and length-normalized theoretical hydraulic conductivity was related to leaf thickness. ? Conclusions: For the whole complex of examined leaf traits, hybrid microspecies were mosaics of parental-like, intermediate, and transgressive phenotypes. The high proportion of transgressive character expressions found in Sorbus hybrids implies that generation of extreme traits through transgressive segregation played a key role in the speciation process.     

Plant hydraulics and photosynthesis of 34 woody species from different successional stages of subtropical forests

It is important to understand the ecophysiological characters of plants when exploring mechanisms underlying species substitution in the process of plant succession. In the present study, we selected 34 woody species from different stages of secondary succession in subtropical forests of southern China, and measured their hydraulic conductivity, gas exchange rates, leaf nutrients and drought‐tolerance traits such as xylem resistance to cavitation, turgor loss point and carbon isotope ratio. Principal component analysis revealed that early‐, mid‐ and late‐successional species were significantly separated along axis 1, which was strongly associated with hydraulic‐photosynthetic coordination. In contrast to species distributed in late‐successional forest, early‐successional species had the highest hydraulic conductivity, net photosynthetic rates, photosynthetic nitrogen and phosphorus use efficiencies, but had the lowest photosynthetic water‐use efficiency. However, changes of the measured drought‐tolerance traits of the 34 species along the succession did not demonstrate a clear trend – no significant correlations between these traits and plant successional stages were found. Moreover, the trade‐off between hydraulic efficiency and safety was not identified. Taken together, our results suggested that hydraulic efficiency and photosynthetic function, rather than drought tolerance, play an important role in species distributions along plant succession in subtropical forests.     

Differential leaf expansion can enable hydraulic acclimation to sun and shade

Although leaf size is one of the most responsive plant traits to environmental change, the functional benefits of large versus small leaves remain unclear. We hypothesized that modification of leaf size within species resulting from differences in irradiance can allow leaves to acclimate to different photosynthetic or evaporative conditions while maintaining an efficient balance between hydraulic supply (vein density) and evaporative demand. To test this, we compared the function and anatomy of leaf hydraulic systems in the leaves of a woody angiosperm (Toona ciliata M. Roem.) grown under high and low irradiance in controlled conditions. Our results confirm that in this species, differential leaf expansion regulates the density of veins and stomata such that leaf hydraulic conductance and stomatal conductance remain proportional. A broader sample of field-grown tree species suggested that differences in leaf venation and stomatal traits induced by sun and shade were not regulated by leaf size in all cases. Our results, however, suggest that leaf size plasticity can provide an efficient way for plants to acclimate hydraulic and stomatal conductances to the contrasting evaporative conditions of sun and shade.     

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.     

植物叶片水力与经济性状权衡关系的研究进展

叶片既是植物光合产物形成的主要场所, 又是整株植物的水力瓶颈、应对灾难性水力失调的安全阀门, 是植物碳水耦合权衡的重要器官。叶经济型谱反映了叶片经济性状“投资-收益”的权衡, 为验证植物进化过程中形成的物种对策提供了适用的理论框架。叶片水力性状变化会影响叶片经济性状及植物存活和生长。因此, 探索植物叶片水力与经济性状的权衡关系, 对建立植物碳-水耦合模型、揭示植物水-碳投资机理、扩展植物性状型谱等均有重要意义。该文首先综述了叶片水力性状、经济性状及两者之间的权衡关系, 分析了叶片导水率与水力脆弱性、失膨点水势、水容、安全阈值等水力性状以及与叶片的形态、结构和气体交换功能性状之间的关系。然后, 从叶片形态、解剖和叶脉网络结构以及气孔功能方面探讨了叶片水力性状与经济性状的调节机制。最后, 提出今后应加强三方面的研究: (1)探索建立植物根-茎-叶水力输导系统的碳-氮-水资源的整株经济型谱, 以揭示植物功能结构耦合、高效固碳用水的生理生态学机制; (2)探索叶片水力安全、水力效率和固碳效率之间的普适性权衡关系, 以深入理解抗旱植物叶片构建的生物物理结构与生理代谢的关系; (3)探索个体水平碳水代谢关系、水分运输与生长速率的耦合, 为代谢推演理论和植物群落尺度预测提供基础。     

Stem and leaf hydraulics of congeneric tree species from adjacent tropical savanna and forest ecosystems

Leaf and stem functional traits related to plant water relations were studied for six congeneric species pairs, each composed of one tree species typical of savanna habitats and another typical of adjacent forest habitats, to determine whether there were intrinsic differences in plant hydraulics between these two functional types. Only individuals growing in savanna habitats were studied. Most stem traits, including wood density, the xylem water potential at 50% loss of hydraulic conductivity, sapwood area specific conductivity, and leaf area specific conductivity did not differ significantly between savanna and forest species. However, maximum leaf hydraulic conductance (K _leaf) and leaf capacitance tended to be higher in savanna species. Predawn leaf water potential and leaf mass per area were also higher in savanna species in all congeneric pairs. Hydraulic vulnerability curves of stems and leaves indicated that leaves were more vulnerable to drought-induced cavitation than terminal branches regardless of genus. The midday K _leaf values estimated from leaf vulnerability curves were very low implying that daily embolism repair may occur in leaves. An electric circuit analog model predicted that, compared to forest species, savanna species took longer for their leaf water potentials to drop from predawn values to values corresponding to 50% loss of K _leaf or to the turgor loss points, suggesting that savanna species were more buffered from changes in leaf water potential. The results of this study suggest that the relative success of savanna over forest species in savanna is related in part to their ability to cope with drought, which is determined more by leaf than by stem hydraulic traits. Variation among genera accounted for a large proportion of the total variance in most traits, which indicates that, despite different selective pressures in savanna and forest habitats, phylogeny has a stronger effect than habitat in determining most hydraulic traits.     

黄土高原油松和刺槐叶片光合生理适应性比较

以黄土高原地区由南向北分布的杨凌、永寿、富县、安塞、米脂、神木等县为研究地点，研究不同地区油松和刺槐的光合特性与叶结构性状间的关系.结果表明，不同地区油松针叶和刺槐叶片的净光合速率（P_n）、光合氮利用效率（PNUE）、水分利用效率（WUE）、比叶质量（LMA）、氮含量（N_mass）和叶绿素相对含量（Chl）差异均达极显著水平（P＜0.001）,说明不同地区油松和刺槐的光合能力和叶结构性状参数差异很大.由南向北，油松的P_n、WUE和PNUE呈略微增加趋势，而刺槐则呈显著降低趋势，表明油松在干旱生境下仍能维持较高的光合能力，而刺槐光合能力明显受到抑制；油松和刺槐的LMA均呈略微上升趋势，而N_mass和Chl均呈略微下降趋势，且刺槐的变化幅度高于油松，说明油松从生理代谢和叶结构性状上对干旱环境的适应能力均强于刺槐.相关分析表明，不同地区油松和刺槐的LMA与N_mass整体上呈极显著负相关；P_n、PNUE与LMA、N_mass相关不显著，与Chl呈极显著正相关；WUE与LMA呈显著负相关（P＜0.05），与N_mass呈显著正相关.     

Stem hydraulic traits and leaf water-stress tolerance are co-ordinated with the leaf phenology of angiosperm trees in an Asian tropical dry karst forest

Background and Aims

The co-occurring of evergreen and deciduous angiosperm trees in Asian tropical dry forests on karst substrates suggests the existence of different water-use strategies among species. In this study it is hypothesized that the co-occurring evergreen and deciduous trees differ in stem hydraulic traits and leaf water relationships, and there will be correlated evolution in drought tolerance between leaves and stems.

Methods

A comparison was made of stem hydraulic conductivity, vulnerability curves, wood anatomy, leaf life span, leaf pressure–volume characteristics and photosynthetic capacity of six evergreen and six deciduous tree species co-occurring in a tropical dry karst forest in south-west China. The correlated evolution of leaf and stem traits was examined using both traditional and phylogenetic independent contrasts correlations.

Key Results

It was found that the deciduous trees had higher stem hydraulic efficiency, greater hydraulically weighted vessel diameter (D_h) and higher mass-based photosynthetic rate (A_m); while the evergreen species had greater xylem-cavitation resistance, lower leaf turgor-loss point water potential (π₀) and higher bulk modulus of elasticity. There were evolutionary correlations between leaf life span and stem hydraulic efficiency, A_m, and dry season π₀. Xylem-cavitation resistance was evolutionarily correlated with stem hydraulic efficiency, D_h, as well as dry season π₀. Both wood density and leaf density were closely correlated with leaf water-stress tolerance and A_m.

Conclusions

The results reveal the clear distinctions in stem hydraulic traits and leaf water-stress tolerance between the co-occurring evergreen and deciduous angiosperm trees in an Asian dry karst forest. A novel pattern was demonstrated linking leaf longevity with stem hydraulic efficiency and leaf water-stress tolerance. The results show the correlated evolution in drought tolerance between stems and leaves.Key words: Tropical dry forest, karst, leaf habit, hydraulic conductivity, cavitation resistance, leaf water-stress tolerance, wood density, leaf density, phylogenetic independent contrasts     

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.