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.     

The blind men and the elephant: the impact of context and scale in evaluating conflicts between plant hydraulic safety and efficiency

Given the fundamental importance of xylem safety and efficiency for plant survival and fitness, it is not surprising that these are among the most commonly studied features of hydraulic architecture. However, much remains to be learned about the nature and universality of conflicts between hydraulic safety and efficiency. Although selection for suites of hydraulic traits that confer adequate plant fitness under given conditions is likely to occur at the organismal level, most studies of hydraulic architecture have been confined to scales smaller than the whole plant, such as small-diameter branches and roots. Here we discuss the impact of the spatial and temporal contexts in which hydraulic traits are studied on the interpretation of their role in maintaining plant hydraulic function. We argue that further advances in understanding the ecological implications of different suites of plant hydraulic traits will be enhanced by adopting an integrated approach that considers variation in hydraulic traits throughout the entire plant, dynamic behavior of water transport, xylem tension and water transport efficiency in intact plants, alternate mechanisms that modulate hydraulic safety and efficiency, and alternate measures of hydraulic safety and safety margins.     

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。     

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。     

Anisotropic hydraulic permeability in compressed articular cartilage

The extent to which articular cartilage hydraulic permeability is anisotropic is largely unknown, despite its importance for understanding mechanisms of joint lubrication, load bearing, transport phenomena, and mechanotransduction. We developed and applied new techniques for the direct measurement of hydraulic permeability within statically compressed adult bovine cartilage explant disks, dissected such that disk axes were perpendicular to the articular surface. Applied pressure gradients were kept small to minimize flow-induced matrix compaction, and fluid outflows were measured by observation of a meniscus in a glass capillary under a microscope. Explant disk geometry under radially unconfined axial compression was measured by direct microscopic observation. Pressure, flow, and geometry data were input to a finite element model where hydraulic permeabilities in the disk axial and radial directions were determined. At less than 10% static compression, near free-swelling conditions, hydraulic permeability was nearly isotropic, with values corresponding to those of previous studies. With increasing static compression, hydraulic permeability decreased, but the radially directed permeability decreased more dramatically than the axially directed permeability such that strong anisotropy (a 10-fold difference between axial and radial directions) in the hydraulic permeability tensor was evident for static compression of 20-40%. Results correspond well with predictions of a previous microstructurally-based model for effects of tissue mechanical deformations on glycosaminoglycan architecture and cartilage hydraulic permeability. Findings inform understanding of structure-function relationships in cartilage matrix, and suggest several biomechanical roles for compression-induced anisotropic hydraulic permeability in articular cartilage.     

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。     

Tree Height-Related Hydraulic Strategy to Cope with Freeze-Thaw Stress in Six Common Urban Tree Species in North China

Urban trees are sensitive to extreme weather events under climate change. Freeze-thaw induced hydraulic failure could induce urban tree dieback and nullify the services they provide. Plant height is a simple but significant trait for plant ecological strategies. Understanding how urban trees with different heights adapt to freeze-thaw stress is increasingly important under climate change. We investigated the relationship between tree height and stem hydraulic functional traits of six common urban tree species in North China to explore tree height-related hydraulic strategies to cope with freeze-thaw stress. Results showed that tall trees had wider vessels, higher hydraulic conductivity, more winter embolism, but lower vessel and wood densities. Positive relationships were found between tree height and vessel diameter, hydraulic conductivity, and freeze-thaw induced embolism, and negative relationships were found between tree height and vessel and wood densities, which implied that short trees employ more conservative ecological strategies than tall trees. Tall and short tree species were well separated by multiple stem hydraulic functional traits; this is consistent with the fact that tall and short trees occupy different niches and indicates that different hydraulic strategies for freeze-thaw stress exist between them. Tall trees might face more pressure to survive under extreme cold weather caused by climate change in the future. Therefore, more attention should be paid to tall urban tree management in North China to cope with extreme cold weather.     

Distribution of xylem hydraulic resistance in fruiting truss of tomato influenced by water stress

In this study xylem hydraulic resistances of peduncles (truss stalk), pedicels (fruit stalk) and the future abscission zone (AZ) halfway along the pedicel of tomato (Lycopersicon esculentum L.) plants were directly measured at different stages of fruit development, in plants grown under two levels of water availability in the root environment. The xylem hydraulic connection between shoot and fruits has previously been investigated, but contradictory conclusions were drawn about the presence of a flow resistance barrier in the pedicel. These conclusions were all based on indirect functional measurements and anatomical observations of water-conducting tissue in the pedicel. In the present study, by far the largest resistances were measured in the AZ where most individual vessels ended. Plants grown at low water availability in the root environment had xylem with higher hydraulic resistances in the peduncle and pedicel segments on both sides of the AZ, while the largest increase in hydraulic resistance was measured in the AZ. During fruit development hydraulic resistances in peduncle and pedicel segments decreased on both sides of the AZ, but tended to increase in the AZ. The overall xylem hydraulic resistance between the shoot and fruit tended to increase with fruit development because of the dominating role of the hydraulic resistance in the AZ. It is discussed whether the xylem hydraulic resistance in the AZ of tomato pedicels in response to water stress and during fruit development contributes to the hydraulic isolation of fruits from diurnal cycles of water stress in the shoot.     

Stem and leaf hydraulic properties are finely coordinated in three tropical rain forest tree species

Coordination of stem and leaf hydraulic traits allows terrestrial plants to maintain safe water status under limited water supply. Tropical rain forests, one of the world's most productive biomes, are vulnerable to drought and potentially threatened by increased aridity due to global climate change. However, the relationship of stem and leaf traits within the plant hydraulic continuum remains understudied, particularly in tropical species. We studied within‐plant hydraulic coordination between stems and leaves in three tropical lowland rain forest tree species by analyses of hydraulic vulnerability [hydraulic methods and ultrasonic emission (UE) analysis], pressure‐volume relations and in situ pre‐dawn and midday water potentials (Ψ). We found finely coordinated stem and leaf hydraulic features, with a strategy of sacrificing leaves in favour of stems. Fifty percent of hydraulic conductivity (P₅₀) was lost at ?2.1 to ?3.1 MPa in stems and at ?1.7 to ?2.2 MPa in leaves. UE analysis corresponded to hydraulic measurements. Safety margins (leaf P₅₀ – stem P₅₀) were very narrow at ?0.4 to ?1.4 MPa. Pressure‐volume analysis and in situ Ψ indicated safe water status in stems but risk of hydraulic failure in leaves. Our study shows that stem and leaf hydraulics were finely tuned to avoid embolism formation in the xylem.     

Dynamic changes in root hydraulic properties in response to nitrate availability

Changes in root hydraulic resistance in response to alterations in nitrate supply were explored in detail as a potential mechanism that allows plants to respond rapidly to changes in their environment. Sunflower (Helianthus annuus cv. Holiday) plants grown hydroponically with limited nitrate availability (200 micromol l(-1)) served as our model system. Experimental plants were 6-9-weeks-old with total dry mass of 2-4 g. Root pressurization of intact plants and detached root systems was used to elucidate the temporal dynamics of root hydraulic properties in sunflower plants following changes in external nitrate availability. The response was rapid, with a 20% decrease in hydraulic resistance occurring within the first hour after the addition of 5 mM nitrate and the magnitude of the effect was dependent on nitrate concentration. The change in root hydraulic resistance was largely reversible, although the temporal dynamics of the response to nitrate addition versus nitrate withdrawal was not symmetric (a gradual decrease in resistance versus its fast increase), raising the possibility that the underlying mechanisms may also differ. Evidence is presented that the observed changes in root hydraulic properties require the assimilation of nitrate by root cells. The hydraulic resistance of roots, previously stimulated by the addition of nitrate, increased more than in control plants in low nitrate under anoxia and that suggests a key role of aquaporin activity in this response. It is proposed that a rapid decrease in root hydraulic resistance in the presence of increased nitrate availability is an important trait that could enhance a plant's ability to compete for nitrate in the soil.     

Water transport through tall trees: A vertically explicit,analytical model of xylem hydraulic conductance in stems

Trees grow by vertically extending their stems, so accurate stem hydraulic models are fundamental to understanding the hydraulic challenges faced by tall trees. Using a literature survey, we showed that many tree species exhibit continuous vertical variation in hydraulic traits. To examine the effects of this variation on hydraulic function, we developed a spatially explicit, analytical water transport model for stems. Our model allows Huber ratio, stem‐saturated conductivity, pressure at 50% loss of conductivity, leaf area, and transpiration rate to vary continuously along the hydraulic path. Predictions from our model differ from a matric flux potential model parameterized with uniform traits. Analyses show that cavitation is a whole‐stem emergent property resulting from non‐linear pressure‐conductivity feedbacks that, with gravity, cause impaired water transport to accumulate along the path. Because of the compounding effects of vertical trait variation on hydraulic function, growing proportionally more sapwood and building tapered xylem with height, as well as reducing xylem vulnerability only at branch tips while maintaining transport capacity at the stem base, can compensate for these effects. We therefore conclude that the adaptive significance of vertical variation in stem hydraulic traits is to allow trees to grow tall and tolerate operating near their hydraulic limits.     

Substantial hydraulic signals are triggered by leaf-biting insects in tomato

In various plants, defence responses can be induced throughoutthe shoot by localized damage or insect attack. Activation ofsuch systemic defence responses must involve a rapid long-distancesignal of wounding. There is firm evidence that, in the caseof localized heat wounds, systemic signalling occurs by hydraulicdispersal of chemical elicitors. However, more natural wounds(such as those imposed by leaf-biting insects) may trigger onlysmall hydraulic events, and it is not clear whether hydraulicdispersal could account for wound signalling in these cases. It is shown here that partial defoliation offers a method foramplifying wound-induced hydraulic events in tomato. Using thisamplification, it is demonstrated that brief feeding by individualleaf-eating insects triggers substantial hydraulic events. Themass flows associated with these events are shown to be sufficientto drive hydraulic dispersal of elicitors through the tomatoplant. It is concluded that hydraulic dispersal could be ofmajor importance for wound signalling in plants in the naturalenvironment. Key words: Lycopersicon esculentum, Spodoptera lit-toralis, wound signalling, systemic defence responses, hydraulic signals     

The relationship between wound-induced proteinase inhibitors and hydraulic signals in tomato seedlings

Localized wounding is known to induce systemic proteinase inhibitors (PI) in seedlings of tomato (Lycopersicon esculentum L.). Inhibitors of elastase (EC 3.4.21.36) were shown here to be among those systemically induced by wounding, and a simple rapid assay for PI based on elastase was developed. Using this assay, the nature of the systemic signalling system (‘PIIF’) was investigated. Hydraulic signals were shown to be induced in tomato by localized wounds. These signals travelled throughout the plant well within the lag time before appearance of systemic wound-induced PI. A number of correlations were drawn between the occurrence of the hydraulic signals and induction of systemic PI, suggesting that hydraulic signals might be the PIIF, or a component of it. It was shown that systemic hydraulic signals could be triggered, without significant wounding, by excision of a single leaflet through the submerged petiole. These hydraulic signals were similar in both kinetics and magnitude to those induced by localized wounding. However, they did not induce systemic PI. In addition, it was shown that systemic events almost as rapid as wound-induced hydraulic signals could be induced without wounding, under certain environmental conditions. This indicates that rapid hydraulic signals do not provide a specific signal of wounding. These findings demonstrate that hydraulic signals per se are not the PIIF.     

Independence of stem and leaf hydraulic traits in six Euphorbiaceae tree species with contrasting leaf phenology

Hydraulic traits and hydraulic-related structural properties were examined in three deciduous (Hevea brasiliensis, Macaranga denticulate, and Bischofia javanica) and three evergreen (Drypetes indica, Aleurites moluccana, and Codiaeum variegatum) Euphorbiaceae tree species from a seasonally tropical forest in south-western China. Xylem water potential at 50% loss of stem hydraulic conductivity (P50_stem) was more negative in the evergreen tree, but leaf water potential at 50% loss of leaf hydraulic conductivity (P50_leaf) did not function as P50_stem did. Furthermore, P50_stem was more negative than P50_leaf in the evergreen tree; contrarily, this pattern was not observed in the deciduous tree. Leaf hydraulic conductivity overlapped considerably, but stem hydraulic conductivity diverged between the evergreen and deciduous tree. Correspondingly, structural properties of leaves overlapped substantially; however, structural properties of stem diverged markedly. Consequently, leaf and stem hydraulic traits were closely correlated with leaf and stem structural properties, respectively. Additionally, stem hydraulic efficiency was significantly correlated with stem hydraulic resistance to embolism; nevertheless, such a hydraulic pattern was not found in leaf hydraulics. Thus, these results suggest: (1) that the evergreen and deciduous tree mainly diverge in stem hydraulics, but not in leaf hydraulics, (2) that regardless of leaf or stem, their hydraulic traits result primarily from structural properties, and not from leaf phenology, (3) that leaves are more vulnerable to drought-induced embolism than stem in the evergreen tree, but not always in the deciduous tree and (4) that there exists a trade-off between hydraulic efficiency and safety for stem hydraulics, but not for leaf hydraulics.     

Decreased hydraulic conductance in plants at elevated carbon dioxide

Previous work indicated that long-term exposure to elevated carbon dioxide levels can reduce hydraulic conductance in some species, but the basis of the response was not determined. In this study, hydraulic conductance was measured at concentrations of both 350 and 700 cm³ m^–3 carbon dioxide for plants grown at both concentrations, to determine the reversibility of the response. In Zea mays and Amaranthus hypochondriacus , exposure to the higher carbon dioxide concentration for several hours reduced whole-plant transpiration rate by 22–40%, without any consistent change in leaf water potential, indicating reversible reductions in hydraulic conductance at elevated carbon dioxide levels. Hydraulic conductance in these species grown at both carbon dioxide concentrations responded similarly to measurement concentration of carbon dioxide, indicating that the response was reversible. In Glycine max , which in earlier work had shown a long-term decrease in hydraulic conductance at elevated carbon dioxide levels, and in Abutilon theophrasti , no short-term changes in hydraulic conductance with measurement concentration of carbon dioxide were found, despite lower transpiration rates at elevated carbon dioxide. In G. max and Medicago sativa , growth at high dew-point temperature reduced transpiration rate and decreased hydraulic conductance. The results indicate that both reversible and irreversible decreases in hydraulic conductance can occur at elevated carbon dioxide concentrations, and that both could be responses to reduced transpiration rate, rather than to carbon dioxide concentration itself.     

从水力结构的角度解释生长温度对烟草叶片形状和大小的影响（英文）

叶片形状和大小在不同的生长温度下变化非常大,但少有从水力结构的角度解释其变化原因的研究。本研究测定了生长于两个不同温度下(24℃/18℃昼/夜;32℃/26℃昼/夜)的烟草叶片的解剖结构,导水率,叶片长宽比和叶面积。生长在24℃/18℃下的烟草叶片与生长在32℃/18℃的叶片相比,更狭窄,并有更小的叶柄导管直径,更低的叶脉密度和导水率。然而,在不同的生长温度下,烟草叶面积并没有显著差异。叶片导水率与叶脉密度呈正相关,但与叶片长宽比呈负相关。结果表明在不同的生长温度下叶片解剖结构和叶片导水率可能对于改变叶宽比起着重要作用。     

干旱区水力提升的生态作用

水力提升是某些植物通过深层根系从较湿的深层土壤中吸取水分再通过浅层根系在较干的浅层土壤中释出的过程。水力提升所释出水量及其释出过程具有积极的生态学意义,它不但为伴生植物提供新的水源、改善其邻近植物的蒸腾作用、缓解水分亏缺、提高水分利用效率．而且促进植物的养分吸收、促进上层干土中分解、硝化和矿化等生化过程．尤其是发生在克隆植物植株间的水分共享这种特殊的水力提升,还影响群落的形成。然而,对于一般的水力提升而言,水力提升所释出水量到底有多少。其生态学作用有多大．有待深入研究。结合研究实践认为．要研究水力提升具有多大的生态学作用．必须准确地区分水;勺提升的水量和毛管水的上升水量。还建议了研究水力提升新的研究方法。     

Hydraulic redistribution in a stand of <Emphasis Type="Italic">Artemisia tridentata</Emphasis>: evaluation of benefits to transpiration assessed with a simulation model

The significance of soil water redistribution facilitated by roots (an extension of "hydraulic lift", here termed hydraulic redistribution) was assessed for a stand of Artemisia tridentata using measurements and a simulation model. The model incorporated water movement within the soil via unsaturated flow and hydraulic redistribution and soil water loss from transpiration. The model used Buckingham-Darcy's law for unsaturated flow while hydraulic redistribution was developed as a function of the distribution of active roots, root conductance for water, and relative soil-root (rhizosphere) conductance for water. Simulations were conducted to compare model predictions with time courses of soil water potential at several depths, and to evaluate the importance of root distribution, soil hydraulic conductance and root xylem conductance on transpiration rates and the dynamics of soil water. The model was able to effectively predict soil water potential during a summer drying cycle, and the rapid redistribution of water down to 1.5 m into the soil column after rainfall events. Results of simulations indicated that hydraulic redistribution could increase whole canopy transpiration over a 100-day drying cycle. While the increase was only 3.5% over the entire 100-day period, hydraulic redistribution increased transpiration up to 20.5% for some days. The presence of high soil water content within the lower rooting zone appears to be necessary for sizeable increases in transpiration due to hydraulic redistribution. Simulation results also indicated that root distributions with roots concentrated in shallow soil layers experienced the greatest increase in transpiration due to hydraulic redistribution. This redistribution had much less effect on transpiration with more uniform root distributions, higher soil hydraulic conductivity and lower root conductivity. Simulation results indicated that redistribution of water by roots can be an important component in soil water dynamics, and the model presented here provides a useful approach to incorporating hydraulic redistribution into larger models of soil processes.     

Changes in hydraulic conductance of citrus trees following a reduction in wetted soil volume

Abstract The effcct of the transition from fully to partially wetted soil voluine on transpiration rate and hydraulic conductance of mature citrus trees was examined in a 23-year-old, coninicrcial, sprinklerirrigated, Shanio u t i orange orchard. I rriga t i on frequency was determined by the rate of water loss from the soil, a s measured by neutron probes. The hydraulic conductance of tlic tree was coniputed from the rclationship between sap flow i n the trunk and leaf water potential. The diurnal valucs of leaf water potential and sap flow shifted towards lower levels as tlie water stored in the root zone was depleted. In the fully wetted soil volume the tree hydraulic conductance remained constant throughout the irrigation period, from June to Novcniber. However, partial wetting of the soil volume (40%) caused a reduction in the hydraulic conductance of the tree. Tlie decreased hydraulic conductance is attributed to tlie permanent interruption of water transport in part of tlie root system. Tlie rcsults of tlie experiment suggest that despite tlie increase of irrigation frequency, partial wetting intensifies water stress in tlie trees.