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。     

木本植物木质部内水柱不连续的证据

利用压力注水的方法使枝干木质部内形成连续水柱,然后与未处理的枝干进行压力测试对比,结果显示只有连续水柱才能在底部产生0.01Mpa/m的压力,而水柱不连续的情况下底部测不到因重力产生的压力。树干液流时滞和木质部空穴现象对木质部水柱不连续均可提供有力的证据。     

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。     

杨树苗木完整植株根压的昼夜节律及其影响因素

【目的】对84K杨树(Populus alba×P.glandulosa)苗木完整植株的根压进行连续测定,研究根压昼夜节律和蒸腾作用、木质部汁液渗透质的关系,为揭示在自然条件下完整植物的根压昼夜节律及发生规律提供参考。【方法】在自然条件下,运用侧面开孔法对杨树苗木完整植株的根压昼夜周期进行测定,并测定叶片水势、根系和茎干木质部汁液的渗透势、无机离子及可溶性糖含量的日变化。【结果】完整杨树植株的根压可以通过侧面开孔法进行监测,测定结果可靠性强。在自然条件下完整植株的根压和去顶根系的根压具有相似的昼夜节律,即昼高夜低,但是完整植株的最大根压值相对较小,且受蒸腾作用的影响。上午蒸腾拉力导致的水势下降并没有抑制根压的上升,中午水势降到一定程度后,根压才迅速下降。1天内任何时间,杨树苗木茎干不同部位木质部汁液存在自上而下渗透势梯度,越到上部渗透势越低,中午尤为突出。此外,茎干木质部汁液渗透势比根系木质部汁液低,根系木质部汁液渗透势在凌晨最低。根系木质部汁液的渗透物质主要是由矿质元素组成,其次是可溶性糖。【结论】完整植株的根压和去顶根系的根压一样,具有明显的昼夜节律,但是在蒸腾作用的影响下最大根压值较小。根压具有较强的自主性,在一定程度上能够抗衡蒸腾拉力。茎干木质部汁液渗透势比根系木质部汁液低,根系木质部汁液的渗透势在凌晨最低,吸水动力最强。     

渗透压在植物体内水分运输中的作用

对渗透压和毛细现象的定量研究表明：在植物体内水分运输过程中，毛细上升只能说明小型植物水分向上移动的范围，而对高大植物（几十米高的大树）来说，渗透压起重要作用。植物生理学中认为，植物体内水分上升的动力来自“根压”和“蒸腾拉力”。“根压”的产生是由于植物细胞内存在着渗透压，“蒸腾”则是由于植物顶部水势高，而顶部大气水势低，致使水分向上渗透而产生拉力。因此，在植物体内水分运输过程中，可认为渗透压起重要作用。     

内聚力-张力学说的新证据(英文)

虽然内聚力 -张力学说已经建立了一个多世纪 ,但要完全接受它仍有一些问题 ,因为它的一个推论是很难证实的 ,即木质部导管中存在大的负压 .特别是近年来木质部压力探针和Z型管试验的结果表明 ,引起空穴的木质部压力仅在 - 0 5MPa以上 .有人根据这一结果认为必须对木质部水分运输的内聚力 -张力学说加以修正 .但是 ,目前又有两个研究小组应用相似改良的Z型管方法测定了产生空穴的木质部压力 .他们的结果显示 ,不同树种木质部导管保持水分传导的压力范围在 - 1 2MPa到- 3 5MPa以下 .这表明树木木质部导管中存在较大负压 ,从而有力支持了内聚力 -张力学说     

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。     

关于植物吸水动力的新见解

叶肉细胞产生的渗透吸力和根细胞产生的渗透压力是水分吸收运输的主要动力。根压是吸水结果而不是吸水动力。蒸腾只是一个失水过程而不能直接产生吸水动力,所谓蒸腾拉力是叶肉细胞通过渗透吸水在导管内产生的负压。水分的吸收运输完全是建立在渗透平衡基础上进行的。     

宁夏盐池沙地水分动态研究初探

采用沙地土壤定位研究方法,系统地研究了毛乌素沙地土壤的水分特性及SPAC(土壤-植物-大气连续体)中水势的变化规律.结果表明(1)经验方程θ=AS-B对该地区的土壤水分特征曲线有良好的模拟性,该区土壤的水分特性为持水量低,供水力小,耐旱性差;(2)水分从树叶扩散到大气中的阻力是土壤-植物-大气连续体中水分传输的主要阻力,土壤水势的变化受降雨、土壤蒸发和林木蒸腾的共同影响,林地的土壤水分表现出明显的季节性变化,树木的叶水势日变化呈明显的单峰曲线,树木受到水分胁迫时水势最低值出现的时间前移,大气水势、土壤水势与叶水势之间均具有一定的相关性.     

黄腐酸钠对侧柏造林生理调节作用的初步研究

侧柏造林苗木叶片蘸黄腐酸钠能够使气孔开张度减小,蒸腾量降低,水份消耗速率减慢,叶片水势和叶绿素含量提高,衰老速度延缓。用0.1%浓度的黄腐酸钠蘸侧柏苗叶,可使造林成活率提高11.3%。     

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.     

Adaptations to soil drying in woody seedlings of African locust bean, (Parkia biglobosa (Jacq.) Benth.)

Stomatal conductance, transpiration and xylem pressure potential of African locust bean (Parkia biglobosa (Jacq.) Benth.) seedlings subjected from the sixth week after emergence to four weeks of continuous soil drought did not differ from those of well-watered, control plants until two-thirds of the available soil water had been used. In both well-watered and drought-treated plants, stomatal conductance was highest early in the day when vapor pressure deficits were low, but decreased sharply by midday when evaporative demand reached its highest value. There was no increase in stomatal conductance later in the day as vapor pressure deficit declined. The relationship between transpiration rate and xylem pressure potential showed non-linearity and hysteresis in both control and drought-treated plants, which seems to indicate that the plants had a substantial capacity to store water. The rate of leaf extension in African locust bean seedlings subjected to six consecutive 2-week cycles of soil drought declined relative to that of well-watered, control plants, whereas relative root extension increased. It appears that African locust bean seedlings minimized the impact of drought by: (1) restricting transpiration to the early part of the day when a high ratio of carbon gain to water loss can be achieved; (2) utilizing internally stored water during periods of rapid transpiration; (3) reducing the rate of leaf expansion and final leaf size in response to soil drought without reducing the rate of root extension, thereby reducing the ratio of transpiring leaf surface area to absorbing root surface area.     

Growth, nutrition and response to water stress of Pinus pinaster inoculated with ten dikaryotic strains of Pisolithus sp

Reconstituted dikaryons of Pisolithus sp. (Pers.) Coker & Couch from South Africa influenced growth parameters (shoot length, shoot/root ratio and leaf area), nutrition and physiological indicators (transpiration rate, stomatal conductance and xylem water potential) of maritime pine (Pinus pinaster Ait.) seedlings during drought and recovery from drought. Seedlings colonized with certain dikaryons were more sensitive to water stress and showed less mycorrhiza formation under water stress than seedlings colonized with other dikaryons. Control (uninoculated) seedlings were significantly smaller than those inoculated with dikaryons. Transpiration rate, stomatal conductance and xylem water potential varied among mycorrhizal treatments during the water stress and recovery periods. After rewatering, the controls and seedlings inoculated with dikaryon 34 x 20 had a weaker recovery in transpiration rate, stomatal conductance and xylem water potential than the other treatments and appeared to have experienced damage due to the water stress. Concentrations of various elements differed in the shoots of Pinus pinaster colonized by the various dikaryons. It is suggested that breeding of ectomycorrhizal fungi could constitute a new tool for improving reforestation success in arid and semi-arid zones.     

Influences of environmental factors on the radial profile of sap flux density in Fagus crenata growing at different elevations in the Naeba Mountains, Japan

Sap flux density was measured continuously during the 1999 and 2000 growing seasons by the heat dissipation method in natural Fagus crenata Blume (Japanese beech) forests growing between 550 and 1600 m on the northern slope of the Kagura Peak of the Naeba Mountains, Japan. Sap flux density decreased radially toward the inner xylem and the decrease was best expressed in relation to the number of annual rings from the cambium, or in relation to the relative depth between the cambium and the trunk center, rather than as a function of absolute depth. The relative influences of radiation, vapor pressure deficit and soil water on sap flux density during the growing season were similar for the outer and inner xylem, and at all sites. Measurements of soil water content and water potential at a depth of 0.25 m demonstrated that sap flux density responded similarly and sensitively to water potential changes in this soil layer, despite large differences in rooting depth at different elevations, localizing one important control point in the functioning of this forest ecosystem. Identification of the relative influences of radiation, vapor pressure deficit and drying of the upper soil layer on sap flux density provides a framework for in-depth analysis of the control of transpiration in Japanese beech forests. In addition, the finding that the same general controls are operating on sap flux density despite climate gradients and large differences in overall forest stand structure will enhance understanding of water use by forests along elevation gradients.     

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.     

The response of Pinus sylvestris to drought: stomatal control of transpiration and hydraulic conductance

We investigated the impact of drought on the physiology of 41-year-old Scots pine (Pinus sylvestris L.) in central Scotland. Measurements were made of the seasonal course of transpiration, canopy stomatal conductance, needle water potential, xylem water content, soil-to-needle hydraulic resistance, and growth. Comparison was made between drought-treated plots and those receiving average precipitation. In response to drought, transpiration rate declined once volumetric water content (VWC) over the top 20 cm of soil reached a threshold value of 12%. Thereafter, transpiration was a near linear function of soil water content. As the soil water deficit developed, the hydraulic resistance between soil and needles increased by a factor of three as predawn needle water potential declined from -0.54 to -0.71 MPa. A small but significant increase in xylem embolism was detected in 1-year-old shoots. Stomatal control of transpiration prevented needle water potential from declining below -1.5 MPa. Basal area, and shoot and needle growth were significantly reduced in the drought treatment. In the year following the drought, canopy stomatal conductance and soil-to-needle hydraulic resistance recovered. Current-year needle extension recovered, but a significant reduction in basal area increment was evident one year after the drought. The results suggest that, in response to soil water deficit, mature Scots pine closes its stomata sufficiently to prevent the development of substantial xylem embolism. Reduced growth in the year after a severe soil water deficit is most likely to be the result of reduced assimilation in the year of the drought, rather than to any residual embolism carried over from one year to the next.     

Die Messung des Wasserpotentials mit der SCHOLANDER-Methode und ihre Bedeutung für die Forstwissenschaft

Summary This deals with a method for determining the water potential of plants and soil and includes details of technique and application. The discussed procedure has found a wide following in the United States, but is nearly unknown in Europe. With different examples of forest research work it is shown that the procedure can be used in manifold ways, is very simple to handle and provides reliable dates. The principle is to place a detached leaf, a twig or a part of a living plant gastight into a steel container— just into the pressure bomb—in such a way that only the plan section is visible. Then the pressure inside the container is increased by means of compressed air or a nitrogen bottle until xylem water appears on the surface. The theory shows that the pressure read at this moment from the pressure gauge agrees with the water potential of the plant.

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Nach einer Gastvorlesung an der Forstlichen Forschungsanstalt München am 13, Mai 1970.     

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.