中亚热带喀斯特常绿落叶阔叶混交林典型树种的木质部解剖与功能特征分析

树木木质部主要由导管、纤维和薄壁组织组成, 分别具有运输、支撑和贮存的生理功能。由于木质部空间限制, 一种组织比例的增加会导致其他组织比例的降低, 因而可能表现出权衡关系。分析木质部组织比例和权衡关系有助于了解植物的生理生态适应性。该研究选择中亚热带喀斯特常绿落叶阔叶混交林21种典型树种(10种落叶树种, 11种常绿树种), 测定枝条木质部各组织比例, 计算水力相关指标并分析性状之间的相关性。结果表明: (1)与全球木质部解剖数据对比分析, 喀斯特树种木质部趋向具有较高比例的薄壁组织; (2)喀斯特树种导管组织比例与薄壁和纤维组织比例之间没有显著的相关性, 但是薄壁和纤维组织比例之间有显著的权衡关系; (3)常绿和落叶树种的木质部水力运输安全性(导管壁加固系数)和效率性(理论导水率)均具有显著的权衡关系, 但是这两个类群线性回归的截距存在显著差异, 即在相同的理论导水率条件下, 落叶树种比常绿树种具有较高的导管壁加固系数(安全性), 可能与常绿树种具有更多的轴向薄壁组织有关。喀斯特树种木质部解剖特征表明薄壁组织的贮存功能对喀斯特树种(尤其是常绿树种)的干旱适应具有重要作用。     

极端干旱环境下的胡杨木质部水力特征

胡杨作为我国西北干旱区重要的乔木树种,研究其木质部水力特征对了解此树种适应极端干旱环境的生物学背景具有较重要的意义。本研究以塔里木河下游的胡杨成株和2年生胡杨幼苗为研究材料,对其木质部最大导水能力（ks(max)）和自然栓塞程度（PLC）等木质部水力特征及其水力特征有关的木质部导管(或管饱)数量特征进行研究。结果表明,成株胡杨多年生枝条和侧根（2≤d<5 mm）木质部自然栓塞程度均较高,PLC均值高于50%,其中多年生枝条栓塞程度具有一定的日变化规律,清晨的PLC均值（58%）小于正午的（67%）;河道边上成株胡杨侧根的均ks(max)和PLC均值都小于距河道200 m处的。随着土壤干旱程度的加剧,幼苗胡杨侧根的自然栓塞程度随之增加,而叶片气孔导度随之降低,土壤含水率与侧根自然栓塞程度,叶片气孔导度之间分别存在显著负相关关系（R =-0.9、R =-0.811）。在统一直径范围内(2≤d<5 mm),成株胡杨侧根均导管直径（dmean）和水力直径均大于（d95%、dh）胡杨幼苗,而导管密度胡杨幼苗高于成株胡杨;胡杨侧根木质部最大导水能力与均导管直径、水力直径之间具有显著正相关关系（R>0.9）.     

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.     

古尔班通古特沙漠梭梭木射线径向运输特征研究

以古尔班通古特沙漠地区优势植物种梭梭（Haloxylon ammodendron）为试验材料,通过观察测定不同树龄、不同季节梭梭木质部解剖结构,计算梭梭木质部非结构性碳水化合物（NSC）及其组分的径向运输速率,分析NSC运输特征与木质部射线和导管解剖结构之间的相关性。结果表明：（1）随着树龄的增加,梭梭导管和射线解剖特征均表现出增加趋势;5月梭梭导管长度和导管直径高于其他季节,10月梭梭射线高度、射线宽度和射线细胞壁厚度均高于其他季节。（2）梭梭木质部NSC、可溶性糖和淀粉的径向运输速率随树龄增长呈显著递增趋势,但13年生与18年生树龄间的NSC、可溶性糖和淀粉的径向运输速率差异不显著。（3）5月和10月的梭梭木质部NSC径向运输速率显著高于7月（P<0.05）;可溶性糖径向运输速率表现为5月最高、7月最低;淀粉径向运输速率从5月到10月呈显著递减趋势。（4）相关分析结果显示,可溶性糖径向运输速率与导管长度（P＜0.05）、导管直径和射线高度呈显著正相关关系（P＜0.01）;淀粉径向运输速率与导管长度、导管直径间呈极显著正相关关系（P＜0.01）;NSC径向运输速率与导管直径、射线高度间呈极显著正相关关系。研究认为,梭梭木质部径向运输功能与木射线和导管解剖结构有关,且呈明显季节动态变化。     

不同林龄刺槐解剖结构及生理特性

全球变暖导致干旱强度增大、频率增加，探究黄土高原不同林龄刺槐林对干旱的响应机制对于理解该区人工林生产力稳定性具有重要意义。本研究以黄土高原半干旱区刺槐幼龄林、中龄林和成熟林为对象，探究不同林龄木质部解剖特征、水力特征和非结构性碳水化合物含量及其之间的相关关系。结果表明：幼龄林、中龄林的导管直径、纹孔总面积、纹孔面积、附着物面积和附着比例均显著大于成熟林，而纹孔密度显著小于成熟林。导管直径、纹孔面积和附着物面积与导水率之间存在显著相关性。幼龄、中龄和成熟林枝条导水率分别为2.30、2.12和0.76 kg·m^-1·s^-1·MPa^-1,栓塞程度分别为54.5%、53.8%和45.1%。导水率与可溶性糖和淀粉含量之间存在显著相关性。幼龄、中龄和成熟林枝条木质部可溶性糖含量分别为4.9%、4.2%和3.8%。成熟林刺槐木质部生长力下降，形成具有较多较小纹孔的小导管，在维持水力安全的同时降低了水分运输效率，组织内维持生长代谢的非结构性碳水化合物含量显著降低。本研究解释了不同林龄刺槐人工林对干旱的响应机制，为黄土高原不同林龄刺槐人工...     

Drought effects on the plasticity in vessel traits of two endemic Magnolia species in the tropical montane cloud forests of eastern Mexico

本文旨在探讨气候变化如何影响两种特有墨西哥木兰的导管性状可塑性和其每年的年轮宽度。尽管如此，很少有研究评估干旱对墨西哥东部热带山地云林导管性状的影响。通过生长年轮的数字图像，我们对所研究的墨西哥木兰树种的径向生长率、树龄和导管性状可塑性进行了评估，比较了两种墨西哥木兰树种在干旱年和非干旱年的导管密度、水力直径和传导面积百分比。本研究首次对墨西哥两个热带山地云林中的两个濒危木兰树种(Magnolia vovidesii 和M. schiedeana)的多孔木质部导管性状对长期气候变化的可塑性进行了分析。结果发现，当比较干旱和非干旱年份时，温度和降水量与年轮宽度的差异密切相关。扩散型多孔木质导管解剖结构具有很高的可塑性，这与所研究的两种木兰树种的温度和/或水的有效性有关。我们的结论是，与非干旱年份相比，对干旱年份的解剖学适应导致了导管性状的大量减少；这些可塑的适应性在植物长期环境胁迫下的水分运输以及生存的安全性方面起着重要的作用。     

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”。     

摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响

以3年生刺槐（Robinia pseudoacacia Linn.）为研究对象,通过对其进行连续3次摘叶造成严重碳限制,检测摘叶后刺槐的生物量分配、叶片形态和不同部位的非结构性碳（NSC）浓度,同时检测其根压和根系导水率、枝条水势和导水率损失值（PLC）及茎的抗栓塞能力,研究摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响。结果显示,摘叶显著降低了刺槐不同部位的生物量,其中细根生物量降低程度最大;摘叶还造成了刺槐不同部位NSC浓度显著降低,茎韧皮部、茎木质部、根韧皮部和根木质部的NSC浓度分别为对照的29.6%、20.2%、10.2%和8.7%,且根部NSC的降低程度显著高于茎;碳限制显著降低了刺槐苗木的根压和根系导水率,增加了枝条凌晨和正午的PLC,降低了其抗栓塞能力。研究结果表明摘叶造成的碳限制改变了刺槐的碳素分配模式,限制了碳素向根的分配,抑制细根的发生,进而限制根的水分吸收能力,加重枝条栓塞程度,同时还会导致枝条抗栓塞能力下降,从而降低植物水分输导的安全性。     

亚低温与干旱胁迫对番茄植株水分传输和形态解剖结构的影响

为探讨亚低温和干旱对植株水分传输的影响机制,以番茄幼苗为试材,利用人工气候室设置常温(昼25 ℃/夜18 ℃)和亚低温(昼15 ℃/夜8 ℃)环境,采用盆栽进行正常灌水(75%～85%田间持水量)和干旱处理(55%～65%田间持水量),分析了温度和土壤水分对番茄植株水分传输、气孔和木质部导管形态解剖结构的影响。结果表明: 与常温正常灌水处理相比,干旱处理使番茄叶水势、蒸腾速率、气孔导度、水力导度、茎流速率、气孔长度和叶、茎、根导管直径显著减小,而使叶、茎、根导管细胞壁厚度和抗栓塞能力增强;亚低温处理下番茄叶水势、蒸腾速率、气孔导度、水力导度和叶、茎、根导管直径显著降低,但气孔变大,叶、根导管细胞壁厚度和叶、茎、根抗栓塞能力显著升高。亚低温条件下土壤水分状况对番茄叶水势、蒸腾速率、气孔导度、水力导度、气孔形态、叶、根导管结构均无显著影响。总之,干旱处理下番茄通过协同调控叶、茎、根结构使植株水分关系重新达到稳态;亚低温处理下番茄植株水分关系的调控主要通过改变叶和根导管结构实现,且受土壤水分状况的影响较小。     

6个耐旱树种木质部结构与栓塞脆弱性的关系

木质部栓塞脆弱性对干旱响应的研究已成为全球气候变化背景下的热点和重要内容。该文以6个耐旱树种刺槐(Robinia pseudoacacia)、沙棘(Hippophae rhamnoides)、榆树(Ulmus pumila)、元宝枫(Acer truncatum)、旱柳(Salix matsudana)、榛(Corylus heterophylla)为研究对象, 采用Cochard Cavitron离心机技术建立木质部栓塞脆弱曲线, 计算木质部栓塞脆弱性, 利用染色法、硅胶注射法等测定木质部导管直径、导管内径跨度、导管连接度、导管密度、导管长度和木质部密度, 探究木质部结构与栓塞脆弱性的关系, 区分6个耐旱树种木质部结构在抗栓塞性上的差异, 以期建立6个耐旱树种在木质部结构方面的抗栓塞性指标。结果表明: 6个耐旱树种木质部栓塞脆弱性大小为刺槐>榆树>沙棘>旱柳>元宝枫>榛, 其中, 刺槐、沙棘和榆树的栓塞脆弱曲线为“r”形, 而元宝枫、旱柳和榛的栓塞脆弱曲线为“s”形, 脆弱曲线为“r”形的树种与脆弱曲线为“s”形的树种栓塞脆弱性差异极显著(p < 0.01)。线性分析表明: 木质部结构影响各树种的栓塞脆弱性, 其中, 木质部密度影响最大(t = 0.702), 导管直径次之(t = 0.532), 导管长度影响最小(t = 0.010)。     

Anatomical adaptations of Astragalus gombiformis Pomel. under drought stress

The present study was designed to study the effect of drought on root, stem and leaf anatomy of Astragalus gombiformis Pomel. Several root, stem and leaf anatomical parameters (cross section diameter, cortex, root cortical cells, pith, leaf lamina and mesophyll thickness) were reduced under moderate to severe water deficit (20–30 days of withheld irrigation). The stele/cross section root ratio increased under moderate water deficit. The root’s and stems vascular systems showed reduced xylem vessel diameter and increased wall thickness under water deficit. In addition, the root xylem vessel density was increased in these drought conditions while it was unchanged in the stems. The stomata density was increased under prolonged drought conditions whereas the stomata size was untouched. The leaf vascular system showed reduced xylem and phloem tissue thickness in the main vein under moderate to severe water deficit. However, in the lamina the vascular tissue and the distance between vascular bundle were unaffected. Our findings suggest a complex network of anatomical adaptations such as a reduced vessel size with increased wall thickness, lesser cortical and mesophyll parenchyma formation and increased stomata density. These proprieties are required for the maintenance of water potential and energy storage under water stress which can improve the resistance of A. gombiformis to survive in arid areas.     

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.     

Impact of extreme drought on tree-ring width and vessel anatomical features of Chukrasia tabularis

Multiple sources of evidence suggest an increasing frequency of extreme climatic events during the past century. In Bangladesh, a country strongly influenced by the South Asian monsoon climate, the years 1999 and 2006 were the most severe droughts among the ten drought events identified over the last four decades. We investigated the impact of these two drought events on radial growth and xylem anatomical features of the brevi-deciduous tree species Chukrasia tabularis in a moist tropical forest in Bangladesh. Tree radial growth declined by 54% during the 1999 and 48.7% during the 2006 droughts, respectively. Among the wood anatomical features, the number of vessels (NV) showed the highest sensitivity to drought, with a 45% decrease in the 1999 drought year, followed by total vessel area (TVA) and mean vessel area (MVA). On the other hand, Vessel density (VD) increased by 13% during the 1999 drought but the increase in VD was very low in the drought year 2006. The decreasing vessel area and increasing vessel density indicate xylem hydraulic adaptation of C. tabularis to minimize drought induced cavitation risk and to avoid hydraulic failure. The significant correlations between the Standardized Precipitation Evapotranspiration Index (SPEI) and time series of tree-ring width and vessel variables imply that decline in radial growth and changes in vessel features in C. tabularis are likely to be caused by drought induced water stress. Our analyses suggest that radial growth and wood anatomical features of C. tabularis are highly sensitive to extreme drought events in South Asian moist tropical forests and can be used to reconstruct past droughts and to model tree response to drought stress under future climate conditions.     

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。     

Poplar vulnerability to xylem cavitation acclimates to drier soil conditions

Xylem vulnerability to cavitation differs between tree species according to their drought resistance, more xerophilous species being more resistant to xylem cavitation. Variability in xylem vulnerability to cavitation is also found within species, especially between in situ populations. The origin of this variability has not been clearly identified. Here we analyzed the response of xylem hydraulic traits of Populus tremula×Populus alba trees to three different soil water regimes. Stem xylem vulnerability was scored as the xylem water potential causing 12, 50 and 88% loss of conductivity (P₁₂, P₅₀ and P₈₈). Vulnerability to cavitation was found to acclimate to growing conditions under different levels of soil water content, with P₅₀ values of ?1.82, ?2.03 and ?2.45 MPa in well‐watered, moderately water‐stressed and severely water‐stressed poplars, respectively. The value of P₁₂, the xylem tension at which cavitation begins, was correlated with the lowest value of midday leaf water potential (ψm) experienced by each plant, the difference between the two parameters being approximately 0.5 MPa, consistent with the absence of any difference in embolism level between the different water treatments. These results support the hypothesis that vulnerability to cavitation is a critical trait for resistance to drought. The decrease in vulnerability to cavitation under growing conditions of soil drought was correlated with decreased vessel diameter, increased vessel wall thickness and a stronger bordered pit field (t/b)². The links between these parameters are discussed.     

Hydraulic, biomechanical, and anatomical interactions of xylem from five species of Acer (Aceraceae)

Possible trade-offs between hydraulic conductivity and mechanical properties of woody stems from five species were assessed. Acer negundo is a ruderal tree, A. saccharinum, and A. rubrum are fast-growing and shade-intolerant soft maples, whereas A. nigrum and A. saccharum are slow-growing and shade-tolerant hard maples. It was hypothesized that the ruderal and soft maples would have lower modulus of elasticity (MOE) and modulus of rupture (MOR), but higher maximum specific conductivity (K(s max)) than hard maples. Many anatomical and general morphological characteristics were measured in an attempt to correlate them to water transport and/or mechanical strength differences between species. No difference was found between species in vessel diameter, fiber wall thickness, initial hydraulic conductivity (K(h initial)), specific conductivity (K(s max)), native percent embolism, or Huber value. Similarly, no trade-off was found between K(s max) and MOE or MOR across the genus. However, fiber lumen diameter was inversely correlated to MOE and MOR. Surprisingly, percentage of ray parenchyma was positively related to MOE. The results suggest transport/mechanical trade-offs do not occur in Acer and differences in mechanical properties may be due to fiber lumen differences that do not influence the efficient transport of water.     

三种锦鸡儿属植物水力结构特征及其干旱适应策略

水分胁迫是干旱半干旱区限制植物生长的主要因素。以干旱半干旱区的3种锦鸡儿属植物为研究对象,从生态适应策略角度来分析3种锦鸡儿植物产生生态分离的原因。对三种锦鸡儿属植物茎干叶片的显微结构、生理功能(导水率、光合速率以及水分利用效率)进行测定,并统计了3种锦鸡儿植株的形态特征,如一、二级枝的直径、长度、末端叶面积。结果表明:三种锦鸡儿属植物都能形成较小的导管直径来适应旱生环境,但是在导水结构上又表现出一定的差异性。中间锦鸡儿的导管直径最小,次脉密度和最大净光合速率最大;柠条锦鸡儿的导管直径、叶片厚度和比叶重(LMA)最大。小叶锦鸡儿在导水率下降50%时的水势(P_(50))最大,水分胁迫时极易发生栓塞,但正是由于导管的栓塞降低了水分运输效率,使其在旱生环境中能够通过减少水分的供应来降低水分的丧失,从而保证自身生长的水分需求;而中间锦鸡儿则主要通过减小导管直径来适应旱生环境;柠条锦鸡儿的水分利用效率最高,抗栓塞能力最强,抗旱性最好,同时柠条锦鸡儿可以通过减少蒸腾面积来减少水分的丧失。植物的导管直径大小、叶片厚度、LMA、叶脉密度对植物导水速率、光合速率等生理功能都有一定的影响。