树木体内水分长距离运输的负压传递机制

蒸腾能使木质部产生不超过-0.1MPa的有限负压,这个负压在木质部内自上而下依次向下传递。水分在导管或管胞内所受的吸附力、毛管力、真空力等综合作用力要大于或等于所受的重力,这个综合作用力可以将水分托住而不受重力作用而下移,使水分在负压作用下以分段移动的方式从根部上升到植物顶端。空穴和栓塞是两个不同的概念,其形成过程、恢复机制都有其本质的不同。空穴的发生与负压和水分的分段移动有密切的关系。     

内聚力学说不可能成立的依据

利用压力室技术测到的水势是缺乏修正的数值,比实际水势要高出许多,用该方法测定水势,只能反映出水势的大小变化,而不能用其代表木质部内真实的负压。蒸腾使木质部产生的负压最大不会超过-0．1Mpa,木质部内巨大负压的存在缺乏理论依据和直接的实验证据。空穴的必然发生使水柱无法保持连续,故连续水柱不存在。蒸腾状态下木质部水分的移动方式是自上而下分段依次移动的,这一现象与内聚力原理相抵触。蒸腾产生的负压能保持到蒸腾停止后继续使根系吸水。     

内聚力-张力学说中关于负压的几点疑问

内聚力-张力学说(C-T学说)是目前解释高大树木体内水分上升机理的唯一理论,但对其的质疑从来没有停过。本文就该理论中关于负压的使用提出几点疑问。在C-T学说中,张应力(单位面积上受到的张力)具有和大气压强、静水压强、水势相同的单位-帕斯卡,且都可为负值,即负压。由于四者单位相同,张应力的负压被等同于负的大气压强、负的静水压强和水势。然而,四者具有不同的物理意义,不能相互比较和代替。负的大气压强和负的静水压强在物理学上不存在;水分在水势的作用下可以自由扩散,水分子间内聚力消失,而被蒸腾拉力往上拽时水分子间需要具有强大的内聚力才能承受张应力,因此,植物体内水分的水势和张应力不可能共存。张应力、水势、大气压强和静水压强四种负压只有一种是存在的,C-T学说需要对该理论中涉及到的负压进行重新合理的诠释。     

美国红梣雄株和雌株茎导管分子的形态解剖比较

运用离析法和显微照相技术,比较美国红梣雄株和雌株茎次生木质部导管分子的特征.结果表明:美国红梣茎次生木质部导管分子中存在着许多不同的样式.雄株茎导管分子具有长导管性、宽导管性、梯纹-网纹导管和单穿孔等特征,这些特征为雄株在生长季节旺盛的水分需求提供强有力的结构保证.雌雄株导管分子差异的发现为该树种的性别鉴定奠定了理论与实践基础.     

活立木生理干燥过程中水分传输和散失机制探讨

木材中的水分会严重影响木材加工和使用性能,必须通过干燥使木材含水率控制在适宜范围内。常规蒸汽干燥耗能大、干燥缺陷多,热泵除湿干燥、太阳能干燥等新型节能干燥技术工业化应用尚不理想,因此,本文从木材水分来源和树木水分生理特性出发,探讨基于蒸腾作用降低木材水分的活立木生理干燥理论和技术,并从水分与植物生理的角度阐述活立木生理干燥的理论基础。通过分析树叶水分蒸发研究进展,总结叶内水分可能的3种蒸发位点,即暴露在内部气体空间的所有叶肉细胞和表皮细胞、气孔下腔室周围大部分区域的叶肉细胞和表皮细胞以及气孔下腔室周围其他区域的叶肉细胞和表皮细胞。通过分析植物体内水分传输机制研究进展和现状,总结植物叶内水分传输的3种可能途径,即通过胞间连丝的共质体传输途径、通过水孔蛋白的跨细胞传输途径以及通过未栓化细胞壁的质外体传输途径。阐明被广泛用于解释木质部水分长距离运输的内聚力-张力学说,分析其目前存在的争论及一些新提出的学说,如补偿压学说、多驱动力学说或水门学说,并分析木质部水分运输过程中时常发生的空穴和栓塞现象及其可能的恢复机制。在此基础上,提出今后研究活立木生理干燥过程中水分传输和散失机制的几个重点和方向:一是探讨生理干燥过程中处于水分胁迫状态下树叶叶孔蒸腾和角质层蒸腾之间的关系;二是探讨生理干燥过程中处于严重水分胁迫状态下树木叶内水分传输途径和蒸发位点;三是探讨纹孔等微观构造在木质部水分长距离传输中的作用以及在空穴和栓塞产生和恢复过程中的作用;四是探讨生理干燥过程中木质部内空穴和栓塞的产生和恢复机制及其对水分长距离传输的作用和影响。     

木质部内负压传递方式的研究

利用杨树、法桐1年生枝条作试验材料,通过施加不同的负压进行传递实验,结果证明木质部内的负压是靠压力平衡方式传递的。水分的移动方式是从上到下分段依次进行的,负压的大小与移动速度呈显著的正相关关系。     

用根压法研究竹子的耐旱、耐寒性

水分在植物体内长距离传输是植物水分研究中一个很重要的研究方向(Tyree,1997;Steudle,2001;Zimmermann et al.,2002;2004)。木质部管道分子(导管或管胞)是植物体内水分长距离运输     

柚木次生木质部导管分子观察研究

运用细胞图象分析系统及显微照相的方法对柚木次生木质部导管分子进行了观察研究.在柚木的次生木质部导管分子中存在着许多不同的样式,分别对其进行了描述,并从导管分子个体发育与系统发育的角度进行了讨论.     

柳苗中的变异电波传递

盐击柳树幼苗根系或烧伤其茎引发的变异电波（ＶＰ）可顺利地从环剥后的木质部通过，并再横向传递至韧皮部（树皮）。但若将柳茎中的木质部切除一段，ＶＰ的传递即被阻断。在柳茎木质部中，ＶＰ传递随水分供应的调节而定，水分供应充分时，ＶＰ传递速度正常；水分供应亏缺时，ＶＰ传递速度减慢或停止。这表明，刺激产生的伤素是借助木质部导管中的蒸腾流传递的，ＶＰ是木质部中伤素传导的反映。     

木材干燥过程中声发射信号分析

对木材在干燥过程中产生的声发射信号进行采集和特征分析,结果表明:木材干燥过程中主要采集到两种不同类型的声发射信号:一种信号来自木材内部自由水的蒸发,与木质部导管内水分空穴化过程的声发射信号一致;另一种信号与木材干燥过快时自身形变开裂相关.作为一种木材干燥质量的控制方法,测量和分析声发射信号,可为木材干燥温度和湿度的控制提供依据,实现防止木材干裂、提高木材干燥质量的目的.     

Xylem cavitation in roots and stems of Douglas-fir and white fir

Roots of hardwoods have been shown to be more vulnerable to xylem cavitation than stems. This study examined whether this pattern is also observed in a conifer species. Vulnerability to cavitation was determined from the pressure required to inject air into the vascular system of hydrated roots and stems, and reduce hydraulic conductance of the xylem. According to the air-seeding hypothesis for the cavitation mechanism, these air pressures predict the negative xylem pressure causing cavitation in dehydrating stems. This was evaluated for stems of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and white fir (Abies concolor (Gord. & Glend.) Lindl.). The air-injection method was applied to roots and stems of different sizes and positions in Douglas-fir trees. Roots, especially smaller roots with a xylem diameter < 5 mm, were more vulnerable to cavitation than stems. Mean cavitation pressure for smaller roots was -2.09 +/- 0.42 versus -3.80 +/- 0.19 MPa for larger roots (> 8 mm diameter). Within the shoot system, smaller stems (< 5 mm diameter) were most vulnerable to cavitation, having a mean cavitation pressure of -4.23 +/- 0.565 versus -5.27 +/- 0.513 MPa for large stems (> 8 mm diameter). There was no correlation between tracheid diameter and mean cavitation pressure within root or stem systems, despite larger tracheid diameters in roots (23.3 +/- 3.9 micro m) than in stems (9.2 +/- 1.6 micro m). Smaller safety margins from cavitation in roots may be beneficial in limiting water use during mild drought, and in protecting the stem from low xylem pressures during extreme drought.     

Influence of soil texture on hydraulic properties and water relations of a dominant warm-desert phreatophyte

We investigated hydraulic constraints on water uptake by velvet mesquite (Prosopis velutina Woot.) at a site with sandy-loam soil and at a site with loamy-clay soil in southeastern Arizona, USA. We predicted that trees on sandy-loam soil have less negative xylem and soil water potentials during drought and a lower resistance to xylem cavitation, and reach E(crit) (the maximum steady-state transpiration rate without hydraulic failure) at higher soil water potentials than trees on loamy-clay soil. However, minimum predawn leaf xylem water potentials measured during the height of summer drought were significantly lower at the sandy-loam site (-3.5 +/- 0.1 MPa; all errors are 95% confidence limits) than at the loamy-clay site (-2.9 +/- 0.1 MPa). Minimum midday xylem water potentials also were lower at the sandy-loam site (-4.5 +/- 0.1 MPa) than at the loamy-clay site (-4.0 +/- 0.1 MPa). Despite the differences in leaf water potentials, there were no significant differences in either root or stem xylem embolism, mean cavitation pressure or Psi(95) (xylem water potential causing 95% cavitation) between trees at the two sites. A soil-plant hydraulic model parameterized with the field data predicted that E(crit) approaches zero at a substantially higher bulk soil water potential (Psi(s)) on sandy-loam soil than on loamy-clay soil, because of limiting rhizosphere conductance. The model predicted that transpiration at the sandy-loam site is limited by E(crit) and is tightly coupled to Psi(s) over much of the growing season, suggesting that seasonal transpiration fluxes at the sandy-loam site are strongly linked to intra-annual precipitation pulses. Conversely, the model predicted that trees on loamy-clay soil operate below E(crit) throughout the growing season, suggesting that fluxes on fine-textured soils are closely coupled to inter-annual changes in precipitation. Information on the combined importance of xylem and rhizosphere constraints to leaf water supply across soil texture gradients provides insight into processes controlling plant water balance and larger scale hydrologic processes.     

A case-study of water transport in co-occurring ring- versus diffuse-porous trees: contrasts in water-status, conducting capacity, cavitation and vessel refilling

Recent work has suggested that the large earlywood vessels of ring-porous trees can be extraordinarily vulnerable to cavitation making it necessary that these trees maintain a consistent and favorable water status. We compared cavitation resistance, vessel refilling, transport capacity and water status in a study of ring-porous Quercus gambelii Nutt. (oak) and diffuse-porous Acer grandidentatum Nutt. (maple). These species co-dominate summer-dry foothills in the western Rocky Mountains of the USA. Native embolism measurements, dye perfusions and balance pressure exudation patterns indicated that the large earlywood vessels of 2-3-year-old oak stems cavitated extensively on a daily basis as predicted from laboratory vulnerability curves, resulting in a more than 80% reduction in hydraulic conductivity. Maple branches showed virtually no cavitation. Oak vessels refilled on a daily basis, despite negative xylem pressure in the transpiration stream, indicating active pressurization of embo-lized vessels. Conductivity and whole-tree water use in oak were between about one-half and two-thirds that in maple on a stem-area basis; but were similar or greater on a leaf-area basis. Oak maintained steady and modest negative xylem pressure potentials during the growing season despite little rainfall, indicating isohydric water status and reliance on deep soil water. Maple was markedly anisohydric and developed more negative pressure potentials during drought, suggesting use of shallower soil water. Although ring porosity may have evolved as a mechanism for coping with winter freezing, this study suggests that it also has major consequences for xylem function during the growing season.     

Xylem conductivity and vulnerability to cavitation of ponderosa pine growing in contrasting climates

We examined the effects of increased transpiration demand on xylem hydraulic conductivity and vulnerability to cavitation of mature ponderosa pine (Pinus ponderosa Laws.) by comparing trees growing in contrasting climates. Previous studies determined that trees growing in warm and dry sites (desert) had half the leaf/sapwood area ratio (A(L)/A(S)) and more than twice the transpiration rate of trees growing in cool and moist sites (montane). We predicted that high transpiration rates would be associated with increased specific hydraulic conductivity (K(S)) and increased resistance to xylem cavitation. Desert trees had 19% higher K(S) than montane trees, primarily because of larger tracheid lumen diameters. Predawn water potential and water potential differences between the soil and the shoot were similar for desert and montane trees, suggesting that differences in tracheid anatomy, and therefore K(S), were caused primarily by temperature and evaporative demand, rather than soil drought. Vulnerability to xylem cavitation did not differ between desert and montane populations. A 50% loss in hydraulic conductivity occurred at water potentials between -2.61 and -2.65 MPa, and vulnerability to xylem cavitation did not vary with stem size. Minimum xylem tensions of desert and montane trees did not drop below -2.05 MPa. Foliage turgor loss point did not differ between climate groups and corresponded to mean minimum xylem tensions in the field. In addition to low A(L)/A(S), high K(S) in desert trees may provide a way to increase tree hydraulic conductivity in response to high evaporative demand and prevent xylem tensions from reaching values that cause catastrophic cavitation. In ponderosa pine, the flexible responses of A(L)/A(S) and K(S) to climate may preclude the existence of significant intraspecific variation in the vulnerability of xylem to cavitation.     

Regulation of water loss in populations of Populus trichocarpa: the role of stomatal control in preventing xylem cavitation

Variations in resistance to drought-induced xylem cavitation, xylem air-entry points, stomatal behavior, and hydraulic conductivity were measured in four populations of Populus trichocarpa Torr. & A. Gray collected along an east-west humidity and temperature gradient in Washington State, USA. Xylem air-entry points were less negative in trees from moist environments (-0.71 and -1.32 MPa in the Hoh and Nisqually populations, respectively) than in trees from dry environments (-1.55 and -1.67 MPa in the Palouse and Yakima populations, respectively). Xylem cavitation in response to experimental drought was consistent with air-injection measures of xylem air-entry points for a given population. Populations vulnerable to cavitation also exhibited higher stem specific hydraulic conductivities and limited stomatal control compared with resistant populations. Populations exhibiting vulnerability to cavitation and limited stomatal control desiccated more rapidly during drought compared with resistant populations. This study provides evidence of interpopulation variation in resistance to drought-induced xylem cavitation, stomatal behavior, and hydraulic conductivity within Populus trichocarpa.     

Vulnerability of several conifers to air embolism

Hydraulic properties of xylem in seven species of conifer were studied during late winter and early spring 1991. Vulnerability to cavitation and air embolism was investigated using hydraulic conductivity and acoustic techniques. Embolisms were induced in branches excised from mature trees by air-drying them in the laboratory. Both techniques gave comparable results indicating that they both assess the same phenomenon. Within a tree, vulnerability was related to the permeability of the xylem, the largest stems tended to cavitate before the smallest ones when water deficits developed in a branch. Interspecific comparisons showed large differences in the xylem water potential needed to induce significant embolism, values ranged from -2.5 MPa in Pinus sylvestris to -4 MPa in Cedrus atlantica, but these differences did not correlate with differences in the xylem permeability of the species. The vulnerability of a species to air embolism was found to be consistent with its ecophysiological behavior in the presence of water stress, drought-tolerant species being less vulnerable than drought-avoiding species.     

Combined application of computer tomography and light microscopy for analysis of conductive xylem area in coarse roots of European beech and Norway spruce

Axial water transport in trees is mainly determined by the gradient of negative water pressure and the structure of conductive xylem elements (i.e. conduits) connecting the fine roots with the foliage. There is still an essential lack of knowledge concerning the relationship between wood structure and hydraulic properties, especially of coarse roots. To this end, the study aimed (1) to work out a novel approach, based on the combination of computer tomography (CT) and light microscopy (LM), for determining the cumulative cross-sectional lumen area of conduits involved in the water transport of coarse roots in European beech (Fagus sylvatica) and Norway spruce (Picea abies) and (2) to demonstrate its adequacy in quantifying the functional relationship between sapwood anatomy and ascending water mass flow in the xylem. The cross-sectional sapwood area of coarse roots was assessed through CT. The cumulative cross-sectional lumen area of conduits in the sapwood (i.e. the lumen area of conductive conduits) was measured by LM in combination with interactive image analysis. The new approach was developed with coarse roots of both the tree species growing in a 60-year-old mixed forest in Bavaria, Germany. The combination of the two methods unveiled spruce to possess a distinct sapwood/heartwood boundary in small-diameter roots, whereas such roots of beech reflected a gradual transition zone; only large-diameter roots displayed a distinct boundary in beech. Additionally, the cumulative lumen area of conductive conduits was found to be approximately 12% of the total coarse root cross-sectional area in both the tree species. The new approach of measuring the conductive lumen area of coarse-root conduits yielded levels of specific sap flow (i.e. axial conductivity) that substantially differed from those derived from commonly applied methods, which were based on sap flow per unit of total cross-sectional root area or xylem cross-sectional area of individual roots. The combination of CT and LM will facilitate functional comparisons of woody roots differing in diameter and of tree species of different anatomical xylem structure.     

Mechanism of cavitation development in the pine wilt disease

Volatile terpenes increase in xylem tissue after infection of Pinus thunbergii with the pine wood nematode (Bursaphelenchus xylophilus). The role of these terpenes in traeheid cavitation, which blocks xylem-sap ascent and leads to water deficit in pine trees, was assessed. Volatile terpene concentration increased long before initiation of tracheid cavitation. After the volatile terpenes reached the highest concentration, severe cavitation developed. Direct injection of α-pinene into healthy pine trunks formed artificial cavitation in xylem. These observations support the hypothesis that excessively produced volatiles, which are hydrophobic and have lower surface tension than water can promote tracheid cavitation in pine wilt disease.     

Differential summer water use by Pinus edulis and Juniperus osteosperma reflects contrasting hydraulic characteristics

Previous studies of pinyon-juniper woodlands show that Pinus edulis Engelm. makes better use of soil water from summer precipitation pulses than does co-occurring Juniperus osteosperma (Torr.) Little. To investigate the basis of this difference, we examined seasonal variation in cavitation and hydraulic conductance. Pinus edulis remained isohydric over the growing season. Minimum water potentials never fell below -2.3 MPa, and the extent of xylem cavitation remained near constant during the dry season. In contrast, J. osteosperma was anisohydric, reaching water potentials as low as -6.9 MPa, and experiencing progressively greater xylem cavitation as the dry season progressed despite having more cavitation-resistant xylem than P. edulis. We conducted an irrigation experiment to observe the responses of the study species to a summer pulse of water. Although sap flow increased in both species in response to the 25-mm irrigation pulse, only J. osteosperma responded to the 10-mm pulse. This was inconsistent with the response of P. edulis to light rain events and may have been due to a difference in the distribution of irrigation water and rain water between the under- and between-canopy areas. Whole-plant conductance increased following the 25-mm irrigation in P. edulis but remained constant in J. osteosperma. We hypothesized that this difference was caused, in part, by differential refilling of embolized xylem. Area specific hydraulic conductivity was 66% higher in roots of irrigated P. edulis trees relative to roots of control trees 3 days after the 25-mm irrigation (t = 2.14, P = 0.02, df = 16). There was no change in hydraulic conductivity of the roots of J. osteosperma or in the stems of either species. Our results indicate that the response to an irrigation pulse in P. edulis depended on cavitation avoidance in stems and the reversal of cavitation in roots, resulting in increased whole-plant conductance and water uptake. In contrast, J. osteosperma failed to exploit light summer rain events but was able to extract deep soil water at low water potentials.