立地条件和树龄对刺槐和小叶杨叶水力性状及抗旱性的影响

以形成黄土高原“小老树”的2种典型树种刺槐和小叶杨为对象,研究了立地条件(沟谷台地和沟间坡地)和树龄对两种树木叶水力学性质和抗旱性的影响,探讨“小老树”形成的水力生理机制.结果表明: 水分较好的沟谷台地上生长的两种树木的叶最大水力导度(K_max)明显大于水分较差的沟间坡地,叶水力脆弱性(P₅₀)也较高;随树龄增加,两种树木的K_max明显下降,但P₅₀差异不大.台地上生长的两种树木的叶表皮导度和PV曲线参数(膨压损失点时的相对含水量RWC_tlp、膨压损失点时的水势ψ_tlp、饱和含水量时的渗透势ψ_sat)均大于坡地;随树龄增加,两种树木的叶表皮导度显著下降,PV曲线参数出现不同程度的下降.两种树木K_max与ψ_tlp呈显著正相关,P₅₀与PV曲线参数之间存在一定的相关性,表明K_max与抗旱性之间存在一种权衡关系,P₅₀是反映两种树木的抗旱性特征之一.     

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.     

陕北沙地高龄小叶杨光合速率下降的水力限制

黄土高原地区“小老树”现象多出现在成年树,幼树相对较少.为探讨树龄影响“小老树”形成的机制,以该地区“小老树”发生面积最大的树种小叶杨为例,研究了3个不同树龄(低龄:13～15 a;中龄:31～34 a;高龄:49～54 a)树木的生长、光合、水力学特性.结果表明: 随树龄增加,小叶杨枯稍长度显著增加,叶片光合速率、气孔导度和蒸腾速率显著下降,整株水力导度也显著下降,不同测定时间的光合速率、气孔导度与整株水力导度呈显著正相关,表明树龄增加引起的光合速率下降可能与整株水力导度下降有关.与中、低龄相比,高龄小叶杨枝干和叶片抵抗空穴化的能力(P₅₀)更强,但通过脆弱性曲线估算的不同树龄正午时的枝干栓塞程度和叶片水力导度无显著差异,表明高龄小叶杨土-根系统水流阻力的增加可能是导致其整株水力导度降低的重要原因.     

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。     

科尔沁沙地黄柳气孔导度和水分状况随季节和地形的变化特征①

黄柳叶片气孔导度在上午８时后即呈下降趋势,下表皮气孔导度大于上表皮。干旱季节上下表皮气孔日间开启程度不同,下表皮气孔受环境因子的影响较大;湿润季节则相同。不同季节气孔导度值不同,下表皮气孔对季节变化反应灵敏。不同季节黄柳体内水分状况参数值不同,在５、６月份抗旱性最强;沙丘不同部位各参数值亦不相同,在沙丘中部和丘脚抗旱性最强,而在丘顶水分状况最好。湿润的７月黄柳一日内各时刻的膨压值均大于０,说明经期     

两种苹果砧木根系水力结构及其PV曲线水分参数对干旱胁迫的响应

研究干旱胁迫对平邑甜茶(Malus hupehensis)和楸子(Malus prunifolia)根系水力结构及其PV曲线水分参数的影响.设定正常与干旱2种水分处理,对2种苹果砧木进行氯化汞-巯基乙醇处理和压力室-容积(PV)曲线测定试验,并利用高压流速仪(HPFM),测定平邑甜茶和楸子根系水力结构.结果表明:随着水分胁迫的加重,平邑甜茶和楸子的根系导水率、根系叶比导水率、根系茎比导水率出现减少趋势.在适宜水分和重度干旱条件下,平邑甜茶根系叶比导水率分别为楸子根系叶比导水率的95％和92％,平邑甜茶根系茎比导水率分别为楸子根系茎比导水率的52％和62％,楸子与平邑甜茶相比在根系茎比导水率和根系叶比导水率上出现增加趋势.干旱胁迫可能会导致水通道蛋白的活性受到抑制,致使其根系导水率出现降低,继而导致了地上部分气体交换受到影响.严重干旱时,楸子与平邑甜茶相比可能具有更大的水孔蛋白表达量来抵御干旱胁迫.在2种水分条件下,楸子的初始质壁分离时的渗透势(ψstlp)、饱和含水时的渗透势(Ψssal)、初始质壁分离时的相对水含量(RWCtlp)、初始质壁分离时的相对渗透水含量(ROWCtlp)、组织细胞总体弹性模量(ε')值与平邑甜茶相比较均处于较低水平,束缚水含量(Va)值处在较高水平.对PV曲线水分参数进行隶属函数综合评价得出的△值为楸子大于平邑甜茶,平邑甜茶和楸子之间b值差异不明显.在适宜水分和重度干旱条件下楸子所体现出的输水策略优于平邑甜茶.PV曲线水分参数同苹果砧木根系的水力结构一样能够随着植物所处的环境做出相应的调整.对于PV曲线水分参数研究发现,楸子在膨压保持方面与平邑甜茶相比,其抗旱性优于平邑甜茶.     

科尔沁沙地黄柳气孔导度和水分状况随季节和地形的变化特征

黄柳叶片气孔导度在上午8时后即呈下降趋势,下表皮气孔导度大于上表皮。干旱季节上下表皮气孔日间开启程度不同,下表皮气孔受环境因子的影响较大;湿润季节则相同。不同季节气孔导度值不同,下表皮气孔对季节变化反应灵敏。不同季节黄柳体内水分状况参数值不同,在5、6月份抗旱性最强;沙丘不同部位各参数值亦不相同,在沙丘中部和丘脚抗旱性最强,而在丘顶水分状况最好。湿润的７月黄柳一日内各时刻的膨压值均大于0,说明虽然此期间水势随环境条件和气孔调节有所升降,但未影响植株的正常生长。     

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。     

应用PV技术对7种针阔叶幼树抗旱性的研究

为进一步探讨PV(Pressure-Volume)技术在树木抗旱性研究中的应用前景,比较成、幼龄树木的抗旱性差别,在对樟子松等7种成龄树木抗旱性生理指标研究的基础上,对其幼树的抗旱性进行了研究。结果表明,对成、幼龄树木一年生小枝用PV曲线所测得的诸水分状况参数随年生长发育进程的的变化规律是一致的;用嫩枝生长初期和枝条完全木质化时期两个阶段的水分参数比较树木的抗旱性、具有较大的可靠性和实用性;幼树在嫩枝生长初期的抗旱性较成树弱,当新枝完全木质化之后,二者的抗旱性基本相近。     

Leaf hydraulic traits of larch and ash trees in response to long-term nitrogen addition in northeastern China

兴安落叶松和水曲柳叶片水力性状对长期氮添加的响应 大气氮沉降影响树木水力结构，进而影响树木的生长和生存。然而，目前关于叶片水力性状对氮沉降的响应尚不明确，而且该响应可能与物种或植物功能型有关。本研究以中国东北地区水曲柳 (Fraxinus mandshurica，阔叶被子植物)和兴安落叶松(Larix gmelinii，针叶裸子植物)人工林为研究对象，利用长达16年的施氮(10 g N m⁻² yr⁻¹)试验研究了氮添加对两种树种叶片水力性状的影响。用自然干燥法测定了叶片压力-容积曲线，用复水动力学法量化了叶片最大水力导度(K_{leaf_max})和抗栓塞阻力(P_50leaf)。研究结果表明，与水曲柳相比，兴安落叶松具有较高的K_{leaf_max}和较强的干旱容忍性(即较低的质壁分离点的相对含水量(RWC_tlp)和弹性模量(ε)，较负的P_50leaf)。此外，氮添加增加了水曲柳的叶膨压损失点水势(π_tlp)、叶饱和含水量时的渗透势(π₀)和叶水容(C_{leaf_mass})，但对兴安落叶松的这些性状影响不显著，表明水曲柳对氮添加更敏感。氮添加增加了水曲柳和兴安落叶松的K_{leaf_max}和P_50leaf。水曲柳的π_tlp和π₀均与叶密度(LD)正相关，而C_{leaf_mass}与LD负相关。兴安落叶松的K_{leaf_max}与LD正相关，P_50leaf与LD负相关。两个树种的K_{leaf_max}与P_50leaf均呈负相关关系。我们的研究表明，长期氮添加降低了这两个重要造林树种的叶片干旱容忍性，这一发现加深了我们对氮沉降背景下树木水力表现的理解。     

Hydraulic architecture of plants of Helianthus annuus L. cv. Margot: evidence for plant segmentation in herbs

The hydraulic architecture of sunflower (Helianthus annuus L. cv. Margot) was studied in terms of the partitioning of the hydraulic conductance (Kleaf) of leaves inserted at progressively more apical nodes both in growing plants (GP) and in plants at full anthesis (mature plants, MP). Leaf conductance to water vapour (gL), leaf water potential (PsiL), leaf water potential at zero turgor (Psi tlp), and leaf osmotic potential at full turgor (pi0) were also measured. Sunflower plants showed gL and Kleaf values significantly increasing in the acropetal direction, while PsiL of basal leaves was significantly more negative than that of distal leaves; Psi tlp markedly decreased in the acropetal direction in MP so that leaves of MP retained increasingly more turgor the more apical they were. This hydraulic pattern, already present in very young plants (GP), strongly favours apical leaves. These data suggest that the progressive leaf dieback starting from the stem base, as observed when the inflorescence of sunflower reached maturity, might be due to time-dependent loss of hydraulic conductance. In fact, Kleaf loss was correlated with PsiL drop and stomatal closure. Leaf dehydration was aggravated by solute exportation from the basal towards the apical leaves, as revealed by the acropetal decrease of pi0. Kleaf was shown to be linearly and positively related to the prevailing ambient irradiance during plant growth, thus suggesting that leaf hydraulics is very sensitive to environmental conditions. It was concluded that the pronounced apical dominance of some sunflower cultivars is determined, among other factors, by plant hydraulic architecture.     

Sap flow as an indicator of transpiration and the water status of young apricot trees

The relationship between water loss via transpiration and stem sap flow in young apricot trees was studied under different environmental conditions and different levels of soil water status. The experiment was carried out in a greenhouse over a 2-week period (November 2–14, 1997) using three-year-old apricot trees (Prunus armeniaca cv. Búlida) growing in pots. Diurnal courses of leaf water potential, leaf conductance and leaf turgor potential also were recorded throughout the experiment. Data from four days of different enviromental conditions and soil water availability have been selected for analysis. On each of the selected days the leaf water potential and the mean transpiration rates were well correlated. The slope of the linear regression of this correlation, taken to indicate the total hydraulic resistance of the tree, confirmed an increasing hydraulic resistance under drought conditions. When the trees were not drought stressed the diurnal courses of sap flow and transpiration were very similar. However, when the trees were droughted, measured of sap flow slightly underestimated actual transpiration. Our heat-pulse measurements suggest the amount of readily available water stored in the stem and leaf tissues of young apricot trees is sufficient to sustain the peak transpiration rates for about 1 hour. This revised version was published online in June 2006 with corrections to the Cover Date.     

Factors controlling plasticity of leaf morphology in Robinia pseudoacacia: III. biophysical constraints on leaf expansion under long-term water stress

In this article, we measured the relative growth rate (RGR) of leaves of Robinia pseudoacacia seedlings under well-watered and water-stressed conditions (mid-day Ψ(w) = leaf water potential estimated with a pressure bomb of -0.48 and -0.98 MPa, respectively). Pressure-volume (PV) curves were done on growing leaves at 25, 50 and 95% of the mature size (growth stage) in order to compute solute potential (Ψ) and turgor pressure (Ψ(P) ) as a function of Ψ(w) . The PV curves and diurnal measurements of Ψ(w) and RGR allowed us to evaluate the parameters (cell wall extensibility m and growth turgor threshold Y) of the Lockhart equation, RGR = m(Ψ(P)-Y), at each growth stage. Our data showed that m and Y did change with leaf age, but the changes were slow enough to evaluate m and Y on any given day. We believe this is the first study to provide evidence that the Lockhart equation adequately quantifies leaf growth of trees over a range of time domains. The value of m linearly declined and Y linearly increased with growth stage. Also, mild drought stress caused a decline in m and increase in Y relative to controls. Although water stress caused an osmotic adjustment which, in turn, increased Ψ(P) in stressed plants relative to controls, the RGR and final leaf sizes were reduced in water-stressed plants because of the impact of water stress on decreased m and increased Y.     

九种不同材性的温带树种叶水力性状及其权衡关系

不同材性树种的解剖、叶脉分布等结构性状差异会影响树木的水分运输效率和水分利用策略, 进而限制树木的生存、生长和分布。然而, 材性对叶导水率、水力脆弱性及其潜在的权衡关系的影响尚不清楚。该研究选择东北温带森林中不同材性的9种树种(散孔材: 山杨(Populus davidiana)、紫椴(Tilia amurensis)、白桦(Betula platyphylla); 环孔材: 蒙古栎(Quercus mongolica)、水曲柳(Fraxinus mandshurica)、胡桃楸(Juglans mandshurica); 无孔材: 红皮云杉(Picea koraiensis)、樟子松(Pinus sylvestris var. mongolica)、红松(Pinus koraiensis), 测量其基于叶面积和叶质量的叶导水率(K_area和K_mass)、水力脆弱性(P₅₀)、膨压丧失点水势(TLP)及叶结构性状, 以比较不同材性树种叶水力性状的差异, 并探索叶水力效率与安全的权衡关系。结果表明: 3种材性树种的K_area、K_mass和P₅₀均差异显著(p < 0.05)。无孔材树种的K_area和K_mass最低, 而散孔材和环孔材树种差异不显著; 环孔材树种P₅₀最高, 而散孔材和无孔材树种差异不显著。K_area和K_mass均与P₅₀显著负相关(p < 0.05), 但散孔材、环孔材和无孔材树种的相关关系分别呈线性、幂函数和指数函数关系。这表明叶水力效率与安全之间存在一定的权衡关系, 但该关系受树木材性的影响。K_mass与TLP显著负相关(p < 0.01), 其中散孔材和环孔材树种呈线性负相关, 无孔材树种呈负指数函数关系; P₅₀随TLP的增加而增加, 这表明树木在面临水分胁迫时, 其质外体和共质体抗旱阻力共同协调保护叶片活细胞, 防止其水分状况到达临界阈值。K_mass与叶干物质含量、叶密度、比叶重均显著负相关, 而P₅₀与之显著正相关(p < 0.01, P₅₀与比叶重的关系除外), 表明树木叶水力特性的变化受相同叶结构特性驱动, 树木增加对水力失调的容忍需要在叶水力系统构建上增加碳投资。     

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.     

黄土区树种抗旱性指数的研究

根据ＰＶ曲线主要水分参数测定数据,进行了黄土区６个乔,灌木树种抗旱性季节变化趋势和树种间抗旱性指数排序分析,结果表明,树种抗旱性指数在生长季（５－１０月）呈具有阶段降低的持续增大趋势,仅在雨季中后期（即树木年第２４生长高峰期）略有减小,在年生长季土壤严重干旱阶段,树种的抗旱性指数大小排序为：海红〉河北杨〉柠条〉小叶杨〉北京杨〉刺槐。