油松、侧柏苗木水力结构特征的对比研究

油松侧柏是分同属两个不同耐旱类型的树种,为了进一步了解其在水分运输和耐旱机理方面的特点,应用水力结构的基本理论,在正常水分条件下,用改良的冲洗法测定４年生油松侧柏苗木各部位的水力结构参数。实验和模拟的结果表明,油松和侧柏苗木都存在限速区;在非限速区,二者各部分木质部的导水串、比导串、叶比导串和胡伯尔值都随功能木质部直径的增加而增大,但表现出不同的变化趋势,这些变化能较好地说明油松和侧柏苗木耐旱的生态策略。在功能木质部直径相同的情况下,油松苗木水力结构的各参数均较侧柏大。     

土壤干旱胁迫对樟树（Cinnamomum camphora (L.) Presl）苗木水力结构特征的影响

植物水力结构特征通常用导水率(Kh)、比导率(Ks)、叶比导率(LSC)、胡伯尔值(Hv)等参数来表征.设置了3种土壤干旱胁迫强度和对照共4种处理,每一处理5个重复,采用"冲洗法"对喀斯特地区造林树种两年生樟树(Cinnamomum camphora (L.) Presl)苗木的水力结构相关参数进行测定.结果表明:樟树苗木在正常水分和不同强度土壤干旱胁迫条件下,其茎段导水率、比导率、叶比导率、胡伯尔值均与茎段直径呈正相关关系,并可以用不同类型函数进行拟合.随着干旱胁迫强度的加剧,不同处理的导水率、比导率、叶比导率降低,胡伯尔值在不同处理间的差异不显著.这说明较粗的茎段,其单位横截面积的导管投入与使用效率明显高于较细茎段,且供给末端叶片水分所需要的压力梯度小于较细茎段.结果为进一步探讨土壤干旱对樟树苗木水力结构作用机理及在石漠化山地造林的适应性提供理论依据.     

干旱胁迫对油松和侧柏水分运输安全性和有效性的影响

通过采用改良的冲洗法测定5年生苗木油松(Pinus tabulaeformis)和侧柏(Platycladus orientalis)在不同干旱胁迫时期的水力结构参数,结果表明随着干旱胁迫增加,不同分枝级和茎段所在区域的导水率损失(PLC)增加,比导率(Ks)减少。油松和侧柏在0级,1级和2级分枝级的发生栓塞的水势阈值分别为-0.55、-0.49、-0.43和-0.90、-0.78、-0.74MPa。随着相对分枝级的增加,油松和侧柏的水势阈值增大,栓塞脆弱性变大。非限速区PLC大而Ks小,限速区PLC小而Ks大。油松和侧柏相对分枝级和茎段所在区域的栓塞脆弱性大小为2级1级0级,限速区非限速区,且油松大于侧柏。油松和侧柏在不同干旱胁迫,不同相对分枝级,不同茎段所在区域采取不同的方式来适应由水势降低而引起的栓塞变化。其采取的生态策略包括:保持较高的水分安全性;减轻安全性而对有效性的折衷;同时降低有效性和安全性但不终止任何生产力或树高的组织生长所需水的限制。     

CO_2浓度升高对4种乔木幼树水力结构的影响

以油松(Pinus tabuliformis)、侧柏(Platycladus orientalis)、元宝枫(Acer truncatum)和刺槐(Robinia pseudoacacia)幼树为研究对象,采用改良冲洗法测定不同分枝级和茎段所在区域的水力结构参数,研究CO_2浓度升高对4个树种水力结构及水分运输安全性和有效性的影响。结果表明:CO_2浓度升高,4个树种在3个分枝级(0、1和2级分枝)的比导率(Ks)和叶比导率(LSC)增加,而侧柏和刺槐在3个分枝的导水率损失(PLC)均下降,油松2级和元宝枫0和1级分枝的PLC下降,其中2级分枝的参数变化最为显著,侧柏、油松和刺槐2级分枝的Ks升高了12.8%、19.6%和51.24%,PLC下降了11.80%、9.6%和51.01%,而元宝枫2级分枝的Ks和PLC均升高。CO_2浓度升高对油松和侧柏非限速区的导水率(Kh)的提高大于限速区,元宝枫和刺槐则相反,而PLC不受CO_2浓度升高的影响。高CO_2浓度对侧柏和元宝枫提高非限速区Ks和LSC大于限速区,油松和刺槐则相反。4个树种在不同分枝级采取不同的方式来适应CO_2浓度的升高,采取的生态策略包括:保持较高的水分运输有效性,同时提高有效性和安全性,和减轻安全性而对有效性的折衷;而在不同茎段所在区域采取保持较高的水分运输效率来适应大气CO_2浓度的升高。     

伊犁河谷退化野果林中新疆野苹果茎的水力结构

为揭示伊犁河谷退化野果林中新疆野苹果植株个体的死亡机理,对比研究3种长势(Ⅰ级,20%枯枝率;Ⅱ级,40%~60%枯枝率;Ⅲ级,>80%枯枝率)新疆野苹果茎的水力结构特征的差异以及水分运输有效性和安全性及其影响因素。结果表明: 随野果林退化程度的增加,3种长势的新疆野苹果茎的边材比导水率和叶比导水率呈下降趋势,但差异不显著;傍晚时的栓塞程度、栓塞脆弱性呈显著增加的趋势,其中Ⅰ、Ⅱ和Ⅲ级长势个体的导水率损失50%时木质部水势分别为-1.87、-1.35和-0.53 MPa;黎明前和正午叶水势、水力安全边际均为Ⅰ级>Ⅱ级>Ⅲ级;与水力学相关的木质部解剖结构以及枝叶性状在3种长势个体之间存在显著差异。相关性分析表明,新疆野苹果茎木质部水分运输的有效性和安全性之间存在弱的权衡关系。新疆野苹果茎的水力结构变化伴随着新疆野果林的衰退过程。野果林的退化会加剧新疆野苹果木质部导管的栓塞化程度,降低其抵抗空穴化栓塞的能力,新疆野苹果面临水力失衡的威胁更大。     

沙柳和柠条茎水力学特性对模拟降雨改变的响应

利用模拟降雨控制试验(对照、降雨增加45%和减少50%),研究了黄土高原水蚀风蚀交错带典型灌木沙柳和柠条茎水力学特性对模拟降雨改变的响应,以揭示两种灌木对未来降雨改变的适应性.结果表明: 沙柳茎比导水率(K_s)、比叶导水率(K_l)和Huber值对增水有显著响应,而对干旱无显著响应;柠条黎明前和正午叶水势、水分传输效率(K_s及K_l)对干旱有显著响应,但对增水无响应.两种灌木不同处理间抵抗栓塞能力无显著差异,沙柳不同处理间正午原位栓塞程度亦无显著差异,而柠条干旱处理正午原位栓塞程度显著增大.沙柳增水处理导管直径和导管面积占边材面积的比例显著增加,干旱导致沙柳导管密度显著增大,水力直径变小;柠条增水处理的木质部结构无明显改变,干旱导致其导管密度和木材密度显著增加.说明增水提升了沙柳的水力功能,而长期干旱显著降低了柠条水力功能,预测在未来气候旱化条件下,柠条的水力适应性可能不如沙柳.     

CO2浓度倍增与干旱胁迫对油松（Pinus tabulaeformis）相对分枝级水力结构的影响

在密闭式生长箱内经过13个月高CO2浓度培养的5年生油松(Pinus tabulaeformis)为实验对象,采用改良冲洗法研究了CO2浓度倍增(720μmolmol-1)与干旱胁迫交互作用对油松相对分枝级水力结构参数的影响.通过测定油松不同分枝级的水力结构参数分别在720μmolmol-1 CO2和380μmolmol-1 CO2(大气现有CO2浓度)浓度下随着干旱胁迫的变化,得出不同分枝级的导水率(Kh)、比导率(Ks)和胡伯尔值(Hv)在2个CO2浓度下均随着干旱胁迫的增加而逐渐下降,叶比导率(Lsc)在720μmolmol-1CO2浓度下随着干旱胁迫的增加非线性变化(0级>2级>1级)不同于380μmolmol-1 CO2(0级>1级>2级).同期干旱胁迫条件下,720μmolmol-1CO2浓度下的Kh、Ks、Lsc 和Hv均大于380μmolmol-1 CO2且差异显著.根据整株苗木的水势将苗木的水分状况分为4个梯度,在正常水分(-0.45～-0.65MPa)、轻度干旱(-1.15～-0.75MPa)和中度干旱(-1.95～-1.35MPa)胁迫时,3个分枝级均在720μmolmol-1CO2条件下的Kh和Ks较380μmolmol-1 CO2增加,说明交互作用能提高导水能力,同时加快水分运输效率.在重度干旱(＜-2.80MPa)胁迫时Kh比380μmolmol-1 CO2增加,而Ks比380μmolmol-1 CO2减小,即交互作用提高了水分运输的安全性,却减少了有效性.     

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。     

干旱胁迫下毛白杨和元宝槭的水力学调控

毛白杨(Populus tomentosa)和元宝槭(Acer truncatum)是华北平原人工林的主要树种, 研究两者水力结构和干旱-复水过程中茎非结构性碳水化合物(NSC)含量动态, 可揭示其水力学调控策略, 为全球气候变化背景下华北人工林水分平衡的科学管理提供理论依据。该研究以相同生境下分布的毛白杨和元宝槭幼树为研究材料, 测量两者的茎抗栓塞能力与水力安全阈、水力面积、叶膨压损失点等水力结构参数; 开展干旱-复水实验, 测定茎NSC含量动态以及干旱胁迫解除后复水阶段的木质部栓塞修复能力。结果表明: 毛白杨导水率损失50%对应的水势(-1.289 MPa)高于元宝槭(-2.894 MPa), 且膨压损失点时的渗透势低, 水力安全阈小, 木材密度小, 气孔调节偏向于变水行为, 表现为易栓塞的低水势忍耐脱水耐旱特性, 水分调节对策趋于冒险; 元宝槭则倾向于不易栓塞的高水势延迟脱水耐旱特性, 水分调节对策趋于保守。在干旱-复水实验中, 毛白杨可溶性糖、淀粉和茎NSC含量先减后增, 元宝槭则先增后减; 并且毛白杨表现出比元宝槭更高的栓塞修复能力, 这与植物体内茎NSC含量变化差异具有一定联系。毛白杨较高的栓塞修复能力也为其易栓塞的低水势忍耐脱水耐旱特性及冒险的水分调节对策提供水力安全保障。两树种在水力学调控上表现出的较大差异可能与其生活史特性相关。     

浙江天童常绿阔叶林不同演替阶段木本植物的水力结构特征

水力结构是植物应对环境形成的与水分运输相关的形态策略.探索不同演替阶段和群落不同高度层植物的水力结构特征, 有助于理解植物的水分运输和利用策略.该研究以浙江天童常绿阔叶林演替前中后期群落的上层木(占据林冠层的树种)和下层木(灌木层物种)为对象, 测定了演替共有种(至少存在于两个演替阶段的物种)和更替种(仅存在于某一演替阶段的物种)的枝边材比导率,叶比导率和胡伯尔值, 以及边材疏导面积,末端枝总叶面积和枝条水势, 分析植物水力结构在群落上层木和下层木间以及在演替阶段间的差异, 及其与枝叶性状的相关关系.结果显示: (1)上层木植物边材比导率和叶比导率显著高于下层木植物(p < 0.05); (2)上层木和下层木的边材比导率与叶比导率在演替阶段间均无显著差异(p > 0.05); 上层木的胡伯尔值在演替阶段间无显著差异, 下层木的胡伯尔值随演替显著下降(p < 0.05); (3)上层木共有种仅边材比导率随演替进行显著降低(p < 0.05), 更替种的3个水力结构参数在演替阶段间无显著差异; 下层木共有种水力结构参数在演替阶段间无明显差异, 更替种仅胡伯尔值随演替减小(p < 0.05); (4)植物边材比导率与枝疏导面积和末端枝所支撑的总叶面积显著正相关(p < 0.01), 胡伯尔值与枝条水势及末端枝总叶面积显著负相关(p < 0.01).以上结果表明: 天童常绿阔叶林演替各阶段上层木比下层木具有更大的输水能力和效率; 随着演替进行, 上层木与下层木的共有种和更替种边材比导率的相反变化表明上层木水力结构的变化可能由微生境变化引起, 而下层木水力结构特征的变化可能由物种更替造成.     

The response of European beech (Fagus sylvatica L.) seedlings from two Italian populations to drought and recovery

The response of gas exchange, leaf chlorophyll content, relative fluorescence to decreasing water potential and recovery was followed in European beech seedlings from two Italian populations, differing in their native precipitation amounts. A population from Sicily (southern Italy) was selected as representative of a xeric population while a population from central Italy, Abetone, represented a mesic one. Dry-matter partitioning, leaf area, hydraulic sufficiency and xylem embolism were evaluated in both well-watered control plants as well as in plants subjected to drought. With the onset of water stress, values of water potential, leaf relative water content, net photosynthesis, leaf conductance and leaf chlorophyll concentration decreased concurrently while relative fluorescence remained unchanged. The population from Sicily showed a delay in effects of the imposed drought. Within 5 days of rewatering, leaf conductance was not fully recovered while all of the other parameters recovered to control levels, in both populations. Total, shoot, stem and root dry weight tended to be higher in seedlings from Abetone, even though both populations had similar photosynthetic rates. The population from Sicily exhibited about 3% greater (even if not significant) allocation to roots than the population from Abetone. Seedlings from Abetone had higher, but not significant, leaf specific conductivity and per cent loss in hydraulic conductivity than seedlings from Sicily. Drought resulted in a reduction of hydraulic conductivity and hydraulic sufficiency in seedlings from both populations. Photosynthesis of water stressed plants from both populations appeared to be reduced primarily by carbon dioxide diffusion through stomata and perhaps secondarily by changes in chlorophyll concentration rather than by efficiency of photosystem II. The effect of hydraulic factors on gas exchange during drought and recovery was not clearly evident.     

Rapid hydraulic recovery in Eucalyptus pauciflora after drought: linkages between stem hydraulics and leaf gas exchange

In woody plants, photosynthetic capacity is closely linked to rates at which the plant hydraulic system can supply water to the leaf surface. Drought‐induced embolism can cause sharp declines in xylem hydraulic conductivity that coincide with stomatal closure and reduced photosynthesis. Recovery of photosynthetic capacity after drought is dependent on restored xylem function, although few data exist to elucidate this coordination. We examined the dynamics of leaf gas exchange and xylem function in Eucalyptus pauciflora seedlings exposed to a cycle of severe water stress and recovery after re‐watering. Stomatal closure and leaf turgor loss occurred at water potentials that delayed the extensive spread of embolism through the stem xylem. Stem hydraulic conductance recovered to control levels within 6 h after re‐watering despite a severe drought treatment, suggesting an active mechanism embolism repair. However, stomatal conductance did not recover after 10 d of re‐watering, effecting tighter control of transpiration post drought. The dynamics of recovery suggest that a combination of hydraulic and non‐hydraulic factors influenced stomatal behaviour post drought.     

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.     

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）.     

Relationship of water transport to anatomical features in the mangrove Laguncularia racemosa grown under contrasting salinities

The purpose of this study was to investigate the xylem anatomy and hydraulic characteristics of the mangrove Laguncularia racemosa grown under contrasting salinities. The study addressed the hypothesis that, at high salinity, water transport capacity may decrease in association with higher water use efficiency. Plants were grown in media to which 0, 15 and 30 NaCl was added. Vessel density and diameter were determined in transverse sections of stem and midrib leaves in terminal shoots, and hydraulic parameters were measured. In stems, the vessel density increased with salinity, while the anatomical diameter (d(a)) and hydraulic diameter (d(h)) declined; in leaves, these parameters remained unchanged with salinity. Huber value and hydraulic and specific conductivities decreased with salinity. Leaf blade resistance increased with salinity and represented the largest fraction of twig resistance. Xylem anatomy and leaf tissue of L. racemosa appeared to be modulated by salinity, which led to a coordinated decline in hydraulic properties as salinity increased. Therefore, these structural changes would reflect functional water use characteristics of leaves under salinity.     

Leaf size serves as a proxy for xylem vulnerability to cavitation in plantation trees

Hybrid poplars are an important renewable forest resource known for their high productivity. At the same time, they are highly vulnerable to water stress. Identifying traits that can serve as indicators for growth performance remains an important task, particularly under field conditions. Understanding which trait combinations translate to improved productivity is key in order to satisfy the demand for poplar wood in an uncertain future climate. In this study, we compared hydraulic and leaf traits among five hybrid poplar clones at 10 plantations in central Alberta. We also assessed the variation of these traits between 2‐ to 3‐year‐old branches from the lower to mid‐crown and current‐year long shoots from the mid to upper crown. Our results showed that (1) hybrid poplars differed in key hydraulic parameters between branch type, (2) variation of hydraulic traits among clones was relatively large for some clones and less for others, and (3) strong relationships between measured hydraulic traits, such as vessel diameter, cavitation resistance, xylem‐specific and leaf‐specific conductivity and leaf area, were observed. Our results suggest that leaf size could serve as an additional screening tool when selecting for drought‐tolerant genotypes in forest management and tree improvement programmes.     

Could rapid diameter changes be facilitated by a variable hydraulic conductance?

Adequate radial water transport between elastic bark tissue and xylem is crucial in trees, because it smoothens abrupt changes in xylem water potential, greatly reducing the likelihood of suffering dangerous levels of embolism. The radial hydraulic conductance involved is generally thought to be constant. Evidence collected about variable root and leaf hydraulic conductance led us to speculate that radial hydraulic conductance in stem/branches might also be variable and possibly modulated by putative aquaporins. We therefore correlated diameter changes in walnut (Juglans regia L.) with changes in water potential, altered by perfusion of twig samples with D-mannitol solutions having different osmotic potentials. Temperature and cycloheximide (CHX; a protein synthesis inhibitor) treatments were performed. The temperature response and diameter change inhibition found in CHX-treated twigs underpinned our hypothesis that radial hydraulic conductance is variable and likely mediated by a putative aquaporin abundance and/or activity. Our data demonstrate that radial water transport in stem/branches can take two routes in parallel: an apoplastic and a cell-to-cell route. The contribution of either route depends on the hydraulic demand and is closely linked to a boost of putative aquaporins, causing radial conductance to be variable. This variability should be considered when interpreting and modelling diameter changes.     

Cavitation,stomatal conductance,and leaf dieback in seedlings of two co-occurring Mediterranean shrubs during an intense drought

Seedling shrubs in the Mediterranean semi-arid climate are subjected to intense droughts during summer. Thus, seedlings often surpass their limits of tolerance to water stress, resulting in the loss of hydraulic conductivity due to xylem cavitation. The response in terms of stomatal conductance, vulnerability to cavitation, leaf dieback, and survival were analysed in two co-occurring seedlings of mastic tree (Pistacia lentiscus L.) and kermes oak (Quercus coccifera L.) during an intense drought period. Both species reacted to drought with steep decreases in stomatal conductance before the critical water potential brought about the onset of cavitation events. Q. coccifera showed wider safety margins for avoiding runaway embolism than P. lentiscus and these differences could be related to the particular drought strategy displayed by each species: water saver or water spender. The limits for survival, resprout capacity and leaf dieback were also analysed in terms of loss of conductivity. By contrast with previous studies, the species showing higher seedling survival in the presence of drought also showed higher susceptibility to cavitation and operated with a lower safety margin for cavitation. Both species showed a leaf specific conductivity (LSC) threshold below which leaf biomass had to be regulated to avoid runaway embolism. However, each species displayed a different type of response: P. lentiscus conserved total leaf area up to 100% loss of LSC, whereas Q. coccifera continuously adjusted leaf biomass throughout the drought period in order to maintain the LSC very close to the maximum values recorded without loss of conductivity. Both species maintained the capacity for survival until the loss of conductivity was very nearly 100%.