八个树种叶水力性状对水分条件的响应及其驱动因素

树木叶片的水力效率和安全性会对水分条件的改变做出一定的响应, 进而影响树木的生长和分布, 然而叶导水率(K_leaf)和叶水力脆弱性(P₅₀)对不同水分条件的响应模式及其影响因素尚不清楚。该研究选取了晋西北关帝山和黑茶山两种水分条件下的8种树种, 测量其水力性状、叶片导管和形态性状, 比较两地不同树种的K_leaf和P₅₀的变化, 分析叶片水力效率和安全性之间的权衡关系, 并探讨叶片水力性状在不同树种及水分条件下的响应模式及其驱动因素。结果表明: 对同一树种而言, 湿润的关帝山叶最大导水率(K_max)和P₅₀均高于干旱的黑茶山; 对同一地区而言, 从在高水分条件下生长的树种到在易干旱环境生长的树种, K_max和P₅₀均逐渐下降。K_max、P₅₀、膨压丧失点水势(TLP)之间均存在显著相关关系。两地叶片P₅₀与导管密度、导管塌陷预测值((t/b)³)、叶片厚度、比叶质量显著正相关, 与导管直径、叶面积显著负相关, 不同树种的K_leaf和P₅₀与叶导管性状的关系大于叶形态性状。同一树种的关帝山到黑茶山P₅₀变化量(δP₅₀)与比叶质量和叶干物质含量在两地的变化量显著正相关, 同一树种δP₅₀与叶形态性状变化量的关系大于与叶导管性状的。以上结果表明: 随着水分条件变差, 叶片水力效率降低, 水力安全性提高, 不同树种叶片水力效率与安全性之间存在一定的权衡关系, 不同树种叶水力性状的差别受叶导管性状影响的程度大于受叶形态性状的影响, 同一树种叶水力安全性对水分条件变化的响应主要依靠叶形态性状的驱动, 树木在提高自身叶水力安全的同时增加了叶构建的碳投资。     

Leaf hydraulic traits and their trade-offs for nine Chinese temperate tree species with different wood properties

Aims Trees with different wood properties display variations in xylem anatomy and leaf vein structure, which may influence tree water transport efficiency and water-use strategy, and consequently constrain tree survival, growth and distribution. However, the effects of wood properties on leaf hydraulic conductance and vulnerability and their potential trade-offs at leaf level are not well understood. Our aims were to examine variations in leaf hydraulic traits of trees with different wood properties and explore potential trade-offs between leaf hydraulic efficiency and safety.
Methods Nine tree species with different wood properties were selected for measuring the leaf hydraulic traits, including three diffuse-porous species (Populus davidiana, Tilia amurensis, Betula platyphylla), three ring-porous species (Quercus mongolica, Fraxinus mandshurica, Juglans mandshurica), and three non-porous species (Picea koraiensis, Pinus sylvestris var. mongolica, Pinus koraiensis). Four dominant and healthy trees per species were randomly selected. The hydraulic traits measured included leaf hydraulic conductance on leaf area (K_area) and dry mass (K_mass) basis, leaf hydraulic vulnerability (P₅₀), and leaf water potential at turgor loss point (TLP), while the leaf structural traits were leaf dry mass content (LDMC), leaf density (LD) and leaf mass per unit area (LMA).
Important findings The K_area, K_mass, and P₅₀ differed significantly among the tree species with different woody properties (p < 0.05). Both K_area and K_mass were the lowest for the non-porous trees, and did not differ significantly between the diffuse-porous and ring-porous trees. The ring-porous trees had the highest P₅₀ values, while the diffuse-porous and non-porous trees showed no significant differences in P₅₀. Both K_area and K_mass were negatively correlated with P₅₀ (p < 0.05) for all the trees, and the relationships for the diffuse-porous, ring-porous, and non-porous trees were fitted into linear, power, exponential functions, respectively. This indicates that significant trade-offs exist between leaf hydraulic efficiency and safety. The K_mass was correlated (p < 0.01) with TLP in a negative linear function for the diffuse- and ring-porous trees and in a negative exponential function for the non-porous trees. The P₅₀ increased with increasing TLP. These results suggest that apoplastic and symplastic drought resistance are strictly coordinated in order to protect living cells from approaching their critical water status under water stresses. The K_mass was negatively correlated (p < 0.01) with LDMC, LD, or LMA, while the P₅₀ was positively correlated with LDMC and LD; this suggests that variations in K_mass and P₅₀ are driven by similar changes in structural traits regardless of wood traits. We conclude that the tree tolerance to hydraulic dysfunction increases with increasing carbon investment in the leaf hydraulic system.     

散孔材和环孔材树种叶水分传输能力及其与抗旱性的关系

选取树龄相同的3种散孔材(杨树、梧桐和樱花)和3种环孔材(刺槐、合欢和白蜡)树种,用3种不同方法(解剖法、加压法和水容法)研究了其叶水力导度的差异及与抗旱性(PV曲线参数)的关系.结果显示:解剖法估算的最大叶水力导度高于加压法和水容法,加压法和水容法在6个树种中的5个上测定值完全一致,3种散孔材与环孔材树种的叶最大水力导度无显著差异.3种散孔材树种的饱和渗透势和膨压损失点渗透势与3种环孔材相比差异不大,但膨压损失点的相对含水量则低于环孔材树种,质外体含水量高于环孔材树种,导致其综合抗旱性指数也高于3种环孔材树种.研究表明,散孔材和环孔材树种的叶最大水力导度与其抗旱性之间并无显著相关关系.     

散孔材与环孔材树种枝干、叶水力学特性的比较研究

为揭示散孔材与环孔材树种树木水分生理特性的差异,选取了常见的3种散孔材落叶树种(毛白杨、法国梧桐和樱花)和3种环孔材落叶树种(刺槐、合欢和白蜡),研究了其枝干与叶水力学性质的差异及其协调性。结果表明:3种环孔材树种枝干横截面积基础上的最大比导水率(Ks-max)大于3种散孔材树种,但其木质部对空穴化的脆弱性(P50branch)高于散孔材树种,6种树木枝干的水分传输能力和抵抗空穴化能力之间存在一种相互制约的权衡关系。3种散孔材与3种环孔材树种的叶最大水力导度(Kl-max)和水力脆弱性(P50leaf)并无显著差异;对于3种散孔材树种,叶的水力脆弱性要高于枝干,但对3种环孔材树种而言,枝干的水力脆弱性要高于叶。6种树木枝干和叶的水力学性质(Kmax、P50)之间并无相关关系。这些结果表明:散孔材与环孔材树种的枝干水力学特性有明显差异,但叶水力学特性无差异;枝干与叶水力学性质之间是相互独立的。     

长白山阔叶树种木质部环孔和散孔结构特征的分化导致其水力学性状的显著差异

木质部的解剖结构特征对树木水分传输功能有重要的影响,阔叶树种木质部环孔和散孔结构特征的分化,很可能导致两个功能类群在水力学结构上存在显著差异,但是有关两个功能类群间细致的水力学性状的对比研究还较少,二者整枝水平的导水率及纹孔水平的细致结构差异尚未见报道.本试验以长白山阔叶红松林常见的3个环孔材和4个散孔材乔木树种为研究对象,对比了两个功能类群树种的整枝导水率(k_shoot)、枝条木质部栓塞脆弱性(p_50)等重要水力学相关生理功能特征,并分析了两个功能类群间的木质部组织水平和纹孔水平上的解剖结构特征差异.结果表明:与茎段导水率差异一致,环孔材树种的整枝导水率也显著高于散孔材,但枝条木质部气穴化抵抗力显著弱于散孔材,二者的差异反映了整枝水平上木质部导水效率和安全性之间的权衡关系,与两个功能类群的水力学生理特征存在显著差异一致,二者在最大导管长度、导管直径、纹孔开口面积、纹孔开口比例等光学和扫描电镜观测解剖结构特征上都存在显著差异;木质部解剖特征(组织水平、纹孔水平)和k_shoot、p_50等生理特征间,以及木质部不同解剖特征之间存在显著的相关,且两个功能类群遵循相同的规律,反映了木质部结构对水分传输功能的重要影响,而导水率和气穴化抵抗力对木质部对立的结构要求,体现了树木水分传输系统构建的生物物理局限性.     

气候过渡区环孔材和散孔材树种的水分利用特征

采用Granier树干液流监测系统, 于2014年6-9月份监测河南信阳鸡公山自然保护区内的枫香(散孔材)和栓皮栎(环孔材)水分利用特征及其对环境因子的响应。结果显示: 在7月份干旱天气和虫灾情况下, 导致环孔材树种叶面积大量减少, 7-9月份期间枫香和栓皮栎的日间平均树干液流密度值分别为33.1 g·m^–2·s^–1和24.8 g·m^–2·s^–1; 而6月份的数值分别为31.6 g·m^–2·s^–1和44.2 g·m^–2·s^–1。枫香和栓皮栎的树干液流密度与大气水汽压亏缺(VPD)呈对数函数关系, 决定系数R²分别为0.38和0.91。液流速率与lnVPD的斜率/ VPD =1下的液流速率, 枫香和栓皮栎分别为0.62和2.87; 此比值受栓皮栎的叶面积和水力导度的影响。枫香的实际蒸腾速率普遍比通过方程计算的预测值低。由于栓皮栎水分利用对环境的较高敏感性, 水分胁迫会导致水分利用下降从而影响生长速率, 进而减缓木材产出的时间。因此在气候变化背景下(极端干旱事件频繁发生), 需要根据不同的林业管理目标合理配置两种木材结构树种。     

小兴安岭不同耐荫性树种枝叶性状变异及权衡

随着植株生长,不同耐荫性树种枝叶性状对资源的响应策略存在差异。探究不同耐荫性树种在不同径级间枝叶性状变异及其相关关系,对理解植物功能性状种内和种间变异以及植物对资源的响应策略具有重要意义。以黑龙江凉水国家级自然保护区阔叶红松（Pinus koraiensis）林中不同径级（小树、中等树和大树）的喜光树种（白桦Betula platyphylla、枫桦Betula costata）和耐荫树种（春榆Ulmus japonica、紫椴Tilia amurensis、色木槭Acer mono）为研究对象,测定其出叶强度、枝横截面积、枝干重、单叶面积、总叶面积、总叶干重共6个枝叶性状。利用单因素方差（LSD）分析检验不同耐荫性树种在不同径级间其枝、叶性状是否存在显著差异;以标准化主轴估计（SMA）对不同耐荫性树种枝叶性状间相关关系进行分析。结果表明：除枝干重外,耐荫树种枝叶性状均大于喜光树种;不同耐荫性树种大树枝横截面积均最大,中等树叶面积均最小;喜光树种不同径级间枝干重、总叶干重和出叶强度无显著差异,而耐荫树种均存在显著差异。随径级增大,不同耐荫性树种枝横截面积与叶面积均呈正异速生长关系,且除耐荫树种枝横截面积与单叶面积外,均存在共同斜率;不同耐荫性树种出叶强度与单叶面积存在共同斜率为-0.51和0.47,小树和中等树枝横截面积与总叶面积和总叶干重的异速生长指数均与1存在显著差异,而大树与1无显著差异。结果表明：不同耐荫性树种在不同径级间对资源的获取策略存在差异,不同耐荫性树种小树阶段均表现出资源获取策略,大树阶段则表现出资源保守型策略。与耐荫树种相比,喜光树种表现为快速生长策略,而随着径级增大不同耐荫性树种这种差异逐渐消失,本研究结果为森林演替过程中种内和种间的相互作用提供了一个新的视角。     

我国温带山地森林48种常见树种叶片重量-出叶强度的关系

叶片是植物的主要光合器官, 其质量与数量的权衡关系体现植物对环境的适应策略。在全球气候变化的背景下, 研究叶片质量与数量关系有助于理解植物对环境变化的响应趋势。该研究应用标准化主轴回归方法, 探讨了我国温带山地森林中48个常见树种的单叶干重与出叶强度的权衡关系。结果表明, 所有物种以及落叶阔叶林、常绿和落叶阔叶树种、单叶以及亚冠层阔叶树种的单叶干重与出叶强度表现为异速生长关系; 针叶林、针阔混交林、常绿及落叶针叶树种、复叶以及冠层阔叶树种则表现为等速生长关系。研究结果表明, 叶大小和出叶强度并无恒定的权衡关系。     

东北东部14个温带树种树干呼吸的种内种间变异

采用红外气体分析仪—局部通量测定法于2009年6 — 10月原位测定了东北东部山区14个温带森林主要组成树种的树干表面CO2通量（RW）,旨在量化种内和种间RW的差异,探索RW随树木直径和季节的变化规律,为深入理解温带森林生态系统碳循环过程、构建和校验其模型提供基础数据。测定的树种包括：散孔材阔叶树（白桦、枫桦、山杨、紫椴、五角槭）、环孔材和半环孔材阔叶树（春榆、黄菠萝、胡桃楸、蒙古栎、水曲柳）、针叶树（兴安落叶松、红松、红皮云杉、樟子松）。结果表明：树种、月份及其交互作用均显著地影响RW（P < 0.001）。测定期间平均RW波动在1.32（黄菠萝）— 3.12 μmol CO2?m-2?s-1（兴安落叶松）之间,其中环孔材和半环孔材阔叶树的平均RW高于散孔材阔叶树,而针叶树的平均RW变异较大。各个树种RW的平均值均呈单峰型季节变化模式,其中7月份最高、10月份最低。RW种内的绝对变异（标准误,SE）波动在0.11 — 0.29 μmol CO2?m-2?s-1之间,相对变异（变异系数,CV）波动在61%（红皮云杉）— 89%（白桦）之间。各个树种的RW均有随胸径（DBH）增大而增加的趋势,但两者之间的回归模型及其决定系数因树种而异。除了春榆和水曲柳之外（P > 0.05）,其它树种的RW与DBH之间的回归关系显著（P < 0.05）,表明DBH可作为某些树种RW预测和上推的一个简便实用的指标。这些结果强调了在树干呼吸能力的比较和上推估测时应充分考虑其种内种间差异的重要性。     

清凉峰不同海拔木本植物小枝内叶大小-数量权衡关系

权衡关系是生活史对策理论的基础, 叶大小-数量的权衡关系对理解叶大小进化具有重要的意义。该研究以单叶面积和单叶片干重表示叶大小, 用小枝干重和小枝茎干重表示小枝大小, 采用标准化主轴估计(standardized major axis estimation, SMA)和系统独立比较分析(phylogenetically independent contrast analysis, PIC)的方法, 对浙江省清凉峰自然保护区3个不同海拔落叶阔叶木本植物当年生小枝内的叶大小与数量间的关系进行研究。结果显示, 无论叶大小和小枝大小是用面积或干重表示, 在每个海拔, 叶大小与出叶强度均存在显著的等速负相关关系, 表明在落叶阔叶木本植物中发现的叶大小与出叶强度之间的权衡关系在不同生境物种中是普遍存在的, 植物在叶大小方面的种间变化, 可能不是自然选择的直接产物, 而是叶片数量变化权衡关系的一个副产物。不同海拔间的比较显示, 高海拔物种的叶面积或干重与出叶强度相关关系的y轴截距比中、低海拔物种小, 表明在出叶强度一定的情况下, 高海拔物种比低海拔物种具有更小的叶大小。与高海拔物种相比, 中海拔物种的共同斜率沿共同主轴有一个向上的位移, 表明中海拔物种比高海拔物种具有更大的叶大小, 但出叶强度更小。这些结果表明生境对叶大小-数量的权衡关系具有显著的影响, 中海拔生境具有更适宜植物生长的气候及养分条件, 而高海拔的低温等不利影响使得叶片变小。     

Leaf hydraulic safety margin and safety–efficiency trade-off across angiosperm woody species

Leaf hydraulic conductance and the vulnerability to water deficits have profound effects on plant distribution and mortality. In this study, we compiled a leaf hydraulic trait dataset with 311 species-at-site combinations from biomes worldwide. These traits included maximum leaf hydraulic conductance (K_leaf), water potential at 50% loss of K_leaf (P50_leaf), and minimum leaf water potential (Ψ_min). Leaf hydraulic safety margin (HSM_leaf) was calculated as the difference between Ψ_min and P50_leaf. Our results indicated that 70% of the studied species had a narrow HSM_leaf (less than 1 MPa), which was consistent with the global pattern of stem hydraulic safety margin. There was a positive relationship between HSM_leaf and aridity index (the ratio of mean annual precipitation to potential evapotranspiration), as species from humid sites tended to have larger HSM_leaf. We found a significant relationship between K_leaf and P50_leaf across global angiosperm woody species and within each of the different plant groups. This global analysis of leaf hydraulic traits improves our understanding of plant hydraulic response to environmental change.     

Functional relationships between hydraulic traits and the timing of diurnal depression of photosynthesis

The hydraulic coordination along the water transport pathway helps trees provide adequate water supply to the canopy, ensuring that water deficits are minimized and that stomata remain open for CO₂ uptake. We evaluated the stem and leaf hydraulic coordination and the linkages between hydraulic traits and the timing of diurnal depression of photosynthesis across seven evergreen tree species in the southern Andes. There was a positive correlation between stem hydraulic conductivity (k_s) and leaf hydraulic conductance (K_Leaf) across species. All species had similar maximum photosynthetic rates (A_max). The species with higher k_s and K_Leaf attained A_max in the morning, whereas the species with lower k_s and K_Leaf exhibited their A_max in the early afternoon concurrently with turgor loss. These latter species had very negative leaf water potentials, but far from the pressure at which the 88% of leaf hydraulic conductance is lost. Our results suggest that diurnal gas exchange dynamics may be determined by leaf hydraulic vulnerability such that a species more vulnerable to drought restrict water loss and carbon assimilation earlier than species less vulnerable. However, under stronger drought, species with earlier CO₂ uptake depression may increase the risk of hydraulic failure, as their safety margins are relatively narrow.     

东北温带森林20种乔木树种叶片干旱容忍性特征

全球范围内干旱频率和强度的增加严重影响树木生长,甚至导致森林大面积死亡。压力-容积(PV)曲线能够反映树木对干旱的容忍能力,但在局域尺度上尚未确定哪个PV曲线参数具有最优指示性。通过测定东北温带森林20种主要树种(包括16种被子植物和4种裸子植物)的PV曲线性状,包括质壁分离时的相对含水量(RWC_tlp)、失膨点叶水势(TLP)、饱和含水时的叶渗透势(π₀)、细胞弹性模量(ε)、叶水容(C_leaf)及叶结构性状(比叶面积和叶密度),研究局域尺度上叶片耐旱性的最佳指示性状,并分析叶片PV性状与结构性状间的相关性。结果表明: 被子植物的RWC_tlp 显著大于裸子植物,但其C_leaf 显著小于裸子植物,这表明用RWC_tlp和C_leaf可以指示东北温带森林不同功能型树种间耐旱性的大小。在被子植物中,TLP和π₀与叶密度呈显著负相关,且均与比叶面积呈显著正相关;而ε与比叶面积呈显著负相关。然而,裸子植物PV曲线性状与叶结构性状之间呈现与被子植物完全相反的趋势。裸子植物与被子植物树种之间PV曲线性状与叶结构性状关系的差异,可能归因于二者采取不同的干旱响应和适应策略。     

Leaf hydraulics and drought stress: response, recovery and survivorship in four woody temperate plant species

Efficient conduction of water inside leaves is essential for leaf function, yet the hydraulic-mediated impact of drought on gas exchange remains poorly understood. Here we examine the decline and subsequent recovery of leaf water potential ( Ψ _leaf), leaf hydraulic conductance ( K _leaf), and midday transpiration ( E ) in four temperate woody species exposed to controlled drought conditions ranging from mild to lethal. During drought the vulnerability of K _leaf to declining Ψ _leaf varied greatly among the species sampled. Following drought, plants were rewatered and the rate of E and K _leaf recovery was found to be strongly dependent on the severity of the drought imposed. Gas exchange recovery was strongly correlated with the relatively slow recovery of K _leaf for three of the four species, indicating conformity to a hydraulic-stomatal limitation model of plant recovery. However, there was also a shift in the sensitivity of stomata to Ψ _leaf suggesting that the plant hormone abscisic acid may be involved in limiting the rate of stomatal reopening. The level of drought tolerance varied among the four species and was correlated with leaf hydraulic vulnerability. These results suggest that species-specific variation in hydraulic properties plays a fundamental role in steering the dynamic response of plants during recovery.     

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。