Hydraulic lift: consequences of water efflux from the roots of plants

Hydraulic lift is the passive movement of water from roots into soil layers with lower water potential, while other parts of the root system in moister soil layers, usually at depth, are absorbing water. Here, we review the brief history of laboratory and field evidence supporting this phenomenon and discuss some of the consequences of this below-ground behavior for the ecology of plants. Hydraulic lift has been shown in a relatively small number of species (27 species of herbs, grasses, shrubs, and trees), but there is no fundamental reason why it should not be more common as long as active root systems are spanning a gradient in soil water potential (Ψ_s) and that the resistance to water loss from roots is low. While the majority of documented cases of hydraulic lift in the field are for semiarid and arid land species inhabiting desert and steppe environments, recent studies indicate that hydraulic lift is not restricted to these species or regions. Large quantities of water, amounting to an appreciable fraction of daily transpiration, are lifted at night. This temporary partial rehydration of upper soil layers provides a source of water, along with soil moisture deeper in the profile, for transpiration the following day and, under conditions of high atmospheric demand, can substantially facilitate water movement through the soil-plant-atmosphere system. Release of water into the upper soil layers has been shown to afford the opportunity for neighboring plants to utilize this source of water. Also, because soils tend to dry from the surface downward and nutrients are usually most plentiful in the upper soil layers, lifted water may provide moisture that facilitates favorable biogeochemical conditions for enhancing mineral nutrient availability, microbial processes, and the acquisition of nutrients by roots. Hydraulic lift may also prolong or enhance fine-root activity by keeping them hydrated. Such indirect benefits of hydraulic lift may have been the primary selective force in the evolution of this process. Alternatively, hydraulic lift may simply be the consequence of roots not possessing true rectifying properties (i.e., roots are leaky to water). Finally, the direction of water movement may also be downward or horizontal if the prevailing Ψ_s gradient so dictates, i.e., inverse, or lateral, hydraulic lift. Such downward movement through the root system may allow growth of roots in otherwise dry soil at depth, permitting the establishment of many phreatophytic species. Received: 2 June 1997 / Accepted: 24 September 1997     

植物根水倒流的证据及意义

对根水倒流的研究进展进行了综述,分析和介绍了其存在的类型、普遍性、发生量、发生位置、生理学基础等,讨论了其在植物生理学、生态学、营养学和作物载培学等学科中的重要意义,并是其存在的问题和进一步研究的方向。     

The redistribution of soil water by tree root systems

Plant roots transfer water between soil layers of different water potential thereby significantly affecting the distribution and availability of water in the soil profile. We used a modification of the heat pulse method to measure sap flow in roots of Grevillea robusta and Eucalyptus camaldulensis and demonstrated a redistribution of soil water from deeper in the profile to dry surface horizons by the root system. This phenomenon, termed “hydraulic lift” has been reported previously. However, we also demonstrated that after the surface soils were rewetted at the break of season, water was transported by roots from the surface to deeper soil horizons – the reverse of the “hydraulic lift” behaviour described for other woody species. We suggest that “hydraulic redistribution” of water in tree roots is significant in maintaining root viability, facilitating root growth in dry soils and modifying resource availability. Received: 26 January 1998 / Accepted: 15 April 1998     

The interaction between water and nitrogen translocation in a rhizomatous sedge (Carex flacca)

In order to examine whether the translocation of water and nitrogen in clonal plants is interdependent, interramet translocation of these two resources was investigated in the greenhouse. Two-ramet systems of Carex flacca were imposed to different spatial patterns of water and nitrogen supply. The experimental design allowed to examine the effects of water heterogeneity on nitrogen sharing, and, vice versa, the effects of nitrogen heterogeneity on water sharing. Interramet translocation of both water and nitrogen was quantified by stable isotope labelling. If one of the ramets was deprived of water, nitrogen or both resources (parallel resource heterogeneity), resource translocation towards this ramet was markedly enhanced compared to a control treatment in which both ramets received ample water and ample nitrogen. Under these conditions, the amount of water or nitrogen translocated was not significantly affected by the pattern of heterogeneity of the other resource imposed on the two-ramet system. If one of the interconnected ramets was rooted in dry but nitrogen-rich soil and the other ramet was placed in nitrogen-deficient but well-watered soil (reciprocal resource heterogeneity), a significant amount of water was translocated towards the ramet in dry soil, while the low-N ramet hardly received any nitrogen. These results show that little nitrogen is translocated between ramets in a direction opposite to the transpiration stream within the rhizome. However, nitrogen may be translocated independently from water if both are transported in a similar direction within the clonal system. The effects of translocation on ramet performance (in terms of transpiration, nitrogen accumulation, and biomass) were assessed by comparing interconnected ramets with isolated (severed) ramets that were treated identically. Integration enhanced the performance of ramets deficient of one or both of the resources. In case of water translocation, the transpiration and growth of the water exporting (donor) ramets was similar to the transpiration and growth of their isolated counterparts. When nitrogen was heterogeneously supplied, however, nitrogen accumulation and growth of the donor ramet was reduced to the same extent as the performance of the nitrogen-deficient ramet was increased. Water translocation thus enhanced the performance of the whole plant, while nitrogen only reduced the differences in ramet performance within the plant. In the case of the reciprocal heterogeneity treatment, the benefits of translocation were strongly unidirectional towards the ramet in dry soil. The data for this treatment suggested that total nitrogen accumulation was enhanced by the acquisition of nitrogen from the dry pot as a result of “hydraulic lift” and water exudation in the dry soil. We conclude that nitrogen translocation in clonal plants, and the associated benefits in terms of resource utilization and growth, may strongly depend on the pattern of interramet water transport. The implications are discussed for studies of physiological integration in clonal plants and the patterns of interramet resource sharing in the field. Received: 2 November 1997 / Accepted: 9 April 1998     

Native root xylem embolism and stomatal closure in stands of Douglas-fir and ponderosa pine: mitigation by hydraulic redistribution

Hydraulic redistribution (HR), the passive movement of water via roots from moist to drier portions of the soil, occurs in many ecosystems, influencing both plant and ecosystem-water use. We examined the effects of HR on root hydraulic functioning during drought in young and old-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] and ponderosa pine (Pinus ponderosa Dougl. Ex Laws) trees growing in four sites. During the 2002 growing season, in situ xylem embolism, water deficit and xylem vulnerability to embolism were measured on medium roots (2–4-mm diameter) collected at 20–30 cm depth. Soil water content and water potentials were monitored concurrently to determine the extent of HR. Additionally, the water potential and stomatal conductance (g_s) of upper canopy leaves were measured throughout the growing season. In the site with young Douglas-fir trees, root embolism increased from 20 to 55 percent loss of conductivity (PLC) as the dry season progressed. In young ponderosa pine, root embolism increased from 45 to 75 PLC. In contrast, roots of old-growth Douglas-fir and ponderosa pine trees never experienced more than 30 and 40 PLC, respectively. HR kept soil water potential at 20–30 cm depth above –0.5 MPa in the old-growth Douglas-fir site and –1.8 MPa in the old-growth ponderosa pine site, which significantly reduced loss of shallow root function. In the young ponderosa pine stand, where little HR occurred, the water potential in the upper soil layers fell to about –2.8 MPa, which severely impaired root functioning and limited recovery when the fall rains returned. In both species, daily maximum g_s decreased linearly with increasing root PLC, suggesting that root xylem embolism acted in concert with stomata to limit water loss, thereby maintaining minimum leaf water potential above critical values. HR appears to be an important mechanism for maintaining shallow root function during drought and preventing total stomatal closure.     

Radial transport of water across cortical sleeves of excised roots ofZea mays L.

The stationary radial volume flows across maize (Zea mays L.) root segments without steles (sleeves) were measured under isobaric conditions. The driving force of the volume flow is an osmotic difference between the internal and external compartment of the root preparations. It is generated by differences in the concentrations of sucrose, raffinose or polyethylene glycol. The flows are linear functions of the corresponding osmotic differences ( ) up to osmotic values which cause plasmolysis. The straight lines obtained pass through the origin. No asymmetry of the osmotic barrier could be detected within the range of driving forces applied ( =±0.5 MPa), corresponding to volume-flow densities of j_{v, s}=±7·10^–8 m·s^–1. Using the literature values for the reflection coefficients of sucrose and polyethylene glycol in intact roots (E. Steudle et al. (1987) Plant Physiol.84, 1220–1234), values for the sleeve hydraulic conductivity of about 1·10^–7 m·s^–1 MPa^–1 were calculated. They are of the same order of magnitude as those reported in the literature for the hydraulic conductivity of intact root segments when hydrostatic pressure is applied.Abbreviations and symbols a _s outer surface of sleeve segment - c concentration of osmotically active solute - j _{v, s} radial volume flow density across sleeve segment - Lp_s hydraulic conductivity of sleeves - Lp_r hydraulic conductivity of intact roots - _N thickness of Nernst diffusion layer - reflection coefficient of root for solute - osmotic value of bulk phase - osmotic coefficient     

土壤水势对水曲柳幼苗水分生态的影响

采用根区渗灌控水技术,将土壤水势长期控制在0～-20kPa(W1)、-20～-40kPa(W2)、-40～-60kPa(W3)、-60～-80kPa(W4)、-80～-160kPa(W5)范围内,系统地研究了不同土壤水势条件下水曲柳幼苗的蒸腾过程、吸水过程、根叶水势13动态过程及SPAC体系的水流阻力,结果表明,在亚饱和土壤水分状态下(W1),细根水势最高,水分由土壤进入细根的阻力最小,根系吸水速率最高,从而支持了13间强烈的蒸腾作用．在田间持水量土壤水分状态下(W2),细根吸水阻力成倍增加,吸水速率和蒸腾速率显著下降,但尚未改变蒸腾作用13动态过程的单峰模式．当土壤水分在田间持水量状态以下(W3-W5)时,随着土壤水势递降,细根吸水阻力急剧增加至几倍乃至几十倍,根系吸水速率过低,吸水与蒸腾矛盾加剧,叶水势降至很低,气孔关闭,蒸腾作用受到严重抑制,呈现明显的午休低谷．在实验范围内(0～-160kPa),土壤水分对水曲柳幼苗是非等效的,当土壤水分在田间持水量状态以下(<-40kPa)时,水曲柳全光苗发生显著的水分胁迫。     

土壤水势对水曲柳幼苗水分生态的影响

采用根区渗灌控水技术,将土壤水势长期控制在0～-20kPa(W₁)、-20～-40kPa(W₂)、-40～-60kPa(W₃)、-60～-80kPa(W₄)、-80～-160kPa(W₅)范围内,系统地研究了不同土壤水势条件下水曲柳幼苗的蒸腾过程、吸水过程、根叶水势日动态过程及SPAC体系的水流阻力.结果表明,在亚饱和土壤水分状态下(W₁),细根水势最高,水分由土壤进入细根的阻力最小,根系吸水速率最高,从而支持了日间强烈的蒸腾作用.在田间持水量土壤水分状态下(W₂),细根吸水阻力成倍增加,吸水速率和蒸腾速率显著下降,但尚未改变蒸腾作用日动态过程的单峰模式.当土壤水分在田间持水量状态以下(W₃～W₅)时,随着土壤水势递降,细根吸水阻力急剧增加至几倍乃至几十倍,根系吸水速率过低,吸水与蒸腾矛盾加剧,叶水势降至很低,气孔关闭,蒸腾作用受到严重抑制,呈现明显的午休低谷.在实验范围内(0～-160kPa),土壤水分对水曲柳幼苗是非等效的,当土壤水分在田间持水量状态以下(<-40kPa)时,水曲柳全光苗发生显著的水分胁迫.     

Species-specific patterns of hydraulic lift in co-occurring adult trees and grasses in a sandhill community

Plants can significantly affect ecosystem water balance by hydraulic redistribution (HR) from dry to wet soil layers via roots (also called hydraulic lift, HL, when the redistribution is from deep to shallow soil). However, the information on how co-occurring species in natural habitats differ in HL ability is insufficient. In a field study, we compared HL ability of four tree species (including three congeneric oak species) and two C4 bunch grass species that co-occur in subxeric habitats of fall-line sandhills in southeastern USA. Soil water potentials (_s) were recorded hourly for 3 years both in large chambers that isolated roots for each species and outside the chambers. Outside of root chambers, soil drying occurred periodically in the top 25 cm and corresponded with lack of precipitation during the summer growing season. Soil moisture was continuously available at a 1 m depth. HL activity was observed in three of the tree species, with greater frequency for Pinus palustris than for Quercus laevis and Q. incana. The fourth tree species Q. margaretta did not exhibit HL activity even though it experienced a similar _s gradient. For the C4 bunch grasses, Aristida stricta exhibited a small amount of HL activity, but Schizachyrium scoparium did not. The capacity for HL activity may be linked to the species ecological distribution. The four species that exhibited HL activity in this subxeric habitat are also dominant in adjacent xeric sandhill habitats, whereas the species that did not exhibit HL are scarcely found in the xeric areas. This is consistent with other studies that found greater fine root survival in dry soil for the four xeric species exhibiting HL activity. The differential ability of these species to redistribute water from the deep soil to the rapidly drying shallow soil likely has a strong effect on the water balance of sandhill plant communities, and is likely linked to their differential distribution across edaphic gradients.An erratum to this article can be found at     

玉米生育期土壤—植物—大气连续体水流阻力与水势的分布

根据玉米生育期的田间试验资料分析了土壤-植物-大气连续体中水势和水流阻力的分布,结果表明土壤与植物叶片之间的水势差在玉米抽雄期前达0.8—1.0MPa,到抽雄期以后达1.0—1.5MPa,叶片与大气之间的水势差则在抽雄期前后分别达80—120MPa和60—80MPa;连续体内的水流阻力主要在叶片与大气之间。建立了连续体中玉米叶片水势的动态模拟公式,模拟叶水势具有较高的精度。最后,揭示了叶片蒸腾速率与叶-气系统水势差和水流阻力的关系,当叶片与大气之间的水势差达90—100MPa之后,蒸腾速率随叶-气间水势差增加而减小。     

沙丘多枝柽柳灌丛根层土壤含水量变化特征与根系水力提升证据

分析干旱区深根型荒漠植物的根层土壤水分是揭示荒漠植物与土壤水分关系机理的重要方面。在黑河中游一片风沙侵蚀区域的多枝柽柳(Tamarix ramosissima)人工林地中, 对表层0.3 m到3 m深的土壤不同深度的含水量进行了连续的动态观测。结果显示, 多枝柽柳根系层土壤含水量可以分为明显不同的3层: 浅层(0.2-1.7 m深)相对湿润层、中间(1.7-2.7 m深)相对干层和深层(2.7 m以下)有效含水层。在多枝柽柳生长盛期, 浅层相对湿润层土壤含水量呈现明显的昼夜变化特征, 同时, 在晚上植物根系与浅层土壤之间存在正水势梯度, 这说明存在根系水力提升现象。水力提升是干旱气候下根层浅层土壤含水量保持相对湿润的主要原因, 并因此维系浅层根系的发育, 也为多枝柽柳具备的防风固沙功能提供了可能的解释。据初步估算, 多枝柽柳根系水力提升占每天耗水量的5%-8%, 耗水的主要水分来源仍然是充足的土壤深层有效含水层。     

Aquaporin-mediated changes in hydraulic conductivity of deep tree roots accessed via caves

Although deep roots can contribute substantially to whole-tree water use, little is known about deep root functioning because of limited access for in situ measurements. We used a cave system on the Edwards Plateau of central Texas to investigate the physiology of water transport in roots at 18-20 m depth for two common tree species, Quercus fusiformis and Bumelia lanuginosa. Using sap flow and water potential measurements on deep roots, we found that calculated root hydraulic conductivity (RHC) fluctuated diurnally for both species and decreased under shading for B. lanuginosa. To assess whether these dynamic changes in RHC were regulated during initial water absorption by fine roots, we used an ultra-low flowmeter and hydroxyl radical inhibition to measure in situ fine root hydraulic conductivity (FRHC) and aquaporin contribution to FRHC (AQPC), respectively. During the summer, FRHC and AQPC were found to cycle diurnally in both species, with peaks corresponding to the period of highest transpirational demand at midday. During whole-tree shade treatments, B. lanuginosa FRHC ceased diurnal cycling and decreased by 75 and 35% at midday and midnight, respectively, while AQPC decreased by 41 and 30% during both time periods. A controlled growth-chamber study using hydroponically grown saplings confirmed daily cycling and shade-induced reductions in FRHC and AQPC. Winter measurements showed that the evergreen Q. fusiformis maintained high FRHC and AQPC throughout the year, while the deciduous B. lanuginosa ceased diurnal cycling and exhibited its lowest annual values for both parameters in winter. Adjustments in FRHC and AQPC to changing canopy water demands may help the trees maintain the use of reliable water resources from depth and contribute to the success of these species in this semi-arid environment.     

Xylem structure and water transport in a twiner,a scrambler,and a shrub of Lonicera (Caprifoliaceae)

Summary Wood structure and function was investigated in different growth forms of temperate honeysuckles (Lonicera spp.). All three species had many narrow vessels and relatively few wide ones, with the measured K _h (flow rate/pressure gradient) approximately 24–55% of the theoretical K _h predicted by Poiseuille's law. Only the twiner, Lonicera japonica, had some vessels greater than 50 m in diameter. The twiner also had the narrowest stem xylem diameters, suggesting the greater maximum vessel diameter hydraulically compensated for narrow stems. Conversely, the free-standing shrub, L. maackii, had the greatest annual increments of xylem but the least percent conductive xylem implying that a great portion of the wood was involved with mechanical support. The scrambler, L, sempervirens had low maximum vessel diameter, high Huber values (= xylem area/leaf area), and low specific conductivities (= measured K _h/xylem area), much like the shrub. The greatest vessel frequency occurred in the scrambler (901 vessels · mm^-2), the highest thus far recorded in vines. The lowest Huber value and highest specific conductivity occurred in the twiner, suggesting little self-support but relatively efficient water conduction. LSC (= measured K _h/leaf area) and maximum vessel diameter of Lonicera vines were near the low end of the range for vines in general.     

玉米生育期土壤-植物-大气连续体水流阻力与水势的分布

根据玉米生育期的田间试验资料分析了土壤-植物-大气连续体中水势和水流阻力的分布,结果表明土壤与植物叶片之间的水势差在玉米抽雄期前达0.8—1.0MPa,到抽雄期以后达1.0—1.5MPa,叶片与大气之间的水势差则在抽雄期前后分别达80—120MPa和60—80MPa;连续体内的水流阻力主要在叶片与大气之间.建立了连续体中玉米叶片水势的动态模拟公式,模拟叶水势具有较高的精度.最后,揭示了叶片蒸腾速率与叶-气系统水势差和水流阻力的关系,当叶片与大气之间的水势差达90—100MPa之后,蒸腾速率随叶-气间水势差增加而减小.     

Apoplastic transport of abscisic acid through roots of maize: effect of the exodermis

The exodermal layers that are formed in maize roots during aeroponic culture were investigated with respect to the radial transport of cis-abscisic acid (ABA). The decrease in root hydraulic conductivity (Lp_r) of aeroponically grown roots was stimulated 1.5-fold by ABA (500 nM), reaching Lp_r values of roots lacking an exodermis. Similar to water, the radial flow of ABA through roots (J_ABA) and ABA uptake into root tissue were reduced by a factor of about three as a result of the existence of an exodermis. Thus, due to the cooperation between water and solute transport the development of the ABA signal in the xylem was not affected. This resulted in unchanged reflection coeffcients for roots grown hydroponically and aeroponically. Despite the well-accepted barrier properties of exodermal layers, it is concluded that the endodermis was the more effective filter for ABA. Owing to concentration polarisation effects, ABA may accumulate in front of the endodermal layer, a process which, for both roots possessing and lacking an exodermis, would tend to increase solvent drag and hence ABA movement into the xylem sap at increased water flow (J_Vr). This may account for the higher ABA concentrations found in the xylem at greater pressure difference. Received: 26 January 1999 / Accepted: 26 May 1999     

Abscisic acid and hydraulic conductivity of maize roots: a study using cell- and root-pressure probes

Using root- and cell-pressure probes, the effects of the stress hormone abscisic acid (ABA) on the water-transport properties of maize roots (Zea mays L.) were examined in order to work out dose and time responses for root hydraulic conductivity. Abscisic acid applied at concentrations of 100–1,000 nM increased the hydraulic conductivity of excised maize roots both at the organ (root Lp_r: factor of 3–4) and the root cell level (cell Lp: factor of 7–27). Effects on the root cortical cells were more pronounced than at the organ level. From the results it was concluded that ABA acts at the plasmalemma, presumably by an interaction with water channels. Abscisic acid therefore facilitated the cell-to-cell component of transport of water across the root cylinder. Effects on cell Lp were transient and highly specific for the undissociated (+)-cis-trans-ABA. The stress hormone ABA facilitates water uptake into roots as soils start drying, especially under non-transpiring conditions, when the apoplastic path of water transport is largely excluded. Received: 26 February 2000 / Accepted: 17 August 2000     

Mercurial inhibition of root hydraulic conductance in Vitis spp. rootstocks under water stress

This work aims to quantify the contribution to whole-root water transport of the fraction controlled by cellular metabolism in grape rootstocks upon water stress. We used mercuric chloride as inhibitor of cell metabolism on genotypes obtained from hybridization of Vitis berlandieri with either Vitis rupestris or Vitis riparia, and we found that the fraction of root water transport under metabolic control is higher in former, which are known to be more resistant to water stress. In addition, as these rootstocks showed lower vessel embolization during water stress, we suggested a possible role of cellular metabolism on the control of root embolism.     

The integration of whole-root and cellular hydraulic conductivities in cereal roots

The hydraulic conductivities of excised whole root systems of wheat (Triticum aestivum L. cv. Atou) and of single excised roots of wheat and maize (Zea mays L. cv. Passat) were measured using an osmotically induced back-flow technique. Ninety minutes after excision the values for single excised roots ranged from 1.6·10^-8 to 5.5·10^-8 m·s^-1·MPa^-1 in wheat and from 0.9·10^-8 to 4.8·10^-8 m·s^-1·MPa^-1 in maize. The main source of variation was a decrease in the value as root length increased. The hydraulic conductivities of whole root systems, but not of single excised roots, were smaller 15 h after excision. This was not caused by occlusion of the xylem at the cut end of the coleoptile. The hydraulic conductivities of epidermal, cortical and endodermal cells were measured using a pressure probe. Epidermal and cortical cells of both wheat and maize roots gave mean values of 1.2·10^-7 m·s^-1·MPa^-1 but in endodermal cells (measured only in wheat) the mean value was 0.5·10^-7 m·s^-1·MPa^-1. The cellular hydraulic conductivities were used to calculate the root hydraulic conductivities expected if water flow across the root was via transcellular (vacuole-to-vacuole), apoplasmic or symplasmic pathways. The results indicate that, in freshly excised roots, the bulk of water flow is unlikely to be via the transcellular pathway. This is in contrast to our previous conclusion (H. Jones, A.D. Tomos, R.A. Leigh and R.G. Wyn Jones 1983, Planta 158, 230–236) which was based on results obtained with whole root systems of wheat measured 14–15 h after excision and which probably gave artefactually low values for root hydraulic conductivity. It is now concluded that, near the root tip, water flow could be through a symplasmic pathway in which the only substantial resistances to water flow are provided by the outer epidermal and the inner endodermal plasma membranes. Further from the tip, the measured hydraulic conductivities of the roots are consistent with flow either through the symplasmic or apoplasmic pathways.Symbols L _{p, cell} cell hydraulic conductivity - L _{p, root} root hydraulic conductivity - L _{p, root} calculated root hydraulic conductivity - root reflection coefficient     

Lateral hydraulic conductivity of early metaxylem vessels in Zea mays L. roots

The hydraulic conductivity of the lateral walls of early metaxylem vessels (Lp_x in m · s^–1 · MPa^–1) was measured in young, excised roots of maize using a root pressure probe. Values for this parameter were determined by comparing the root hydraulic conductivities before and after steam-ringing a short zone on each root. Killing of living tissue virtually canceled its hydraulic resistance. There were no suberin lamellae present in the endodermis of the roots used. The value of Lp_x ranged between 3 · 10^–7 and 35 · 10^–7 m · s^–1 · MPa^–1 and was larger than the hydraulic conductivity of the untreated root (Lp_r = 0.7 · 10^–7 to 4.0 · 10^–7 m · s^–1 · MPa^–1) by factor of 3 to 13. Assuming that all flow through the vessel walls was through the pit membranes, which occupied 14% of the total wall area, an upper limit of the hydraulic conductivity of this structure could be given(Lp_pm=21 · 10^–7 to 250 · 10^–7 m · s^–1 · MPa^–1). The specific hydraulic conductivity (Lp_cw) of the wall material of the pit membranes (again an upper limit) ranged from 0.3 · 10^–12 to 3.8 · 10^–12 m² · s^–1 · MPa^–1 and was lower than estimates given in the literature for plant cell walls. From the data, we conclude that the majority of the radial resistance to water movement in the root is contributed by living tissue. However, although the lateral walls of the vessels do not limit the rate of water flow in the intact system, they constitute 8–31% of the total resistance, a value which should not be ignored in a detailed analysis of water flow through roots.Abbreviatations and Symbols k_wr (T ₁ ^2/W ) rate constant (half-time) of water exchange across root (s^–1 or s, respectively) - Lp_cw specific hydraulic conductivity of wall material (m² · s^–1 · MPa^–1) - Lp_pm hydraulic conductivity of pit membranes (m · s ^–1 · MPa^–1) - Lp_r hydraulic conductivity of root (m · s^–1 · MPa^–1) - Lp_x lateralhydraulic conductivity of walls of root xylem (m · s ^–1 · MPa^–1) This research was supported by a grant from the Bilateral Exchange Program funded jointly by the Natural Sciences and Engineering Research Council of Canada and the Deutsche Forschungsgemeinschaft to C.A.P., and by a grant from the Deutsche Forschungsgemeinschaft, Sonderforschungsbereich 137, to E.S. The expert technical help of Mr. Burkhard Stumpf and the work of Ms. Martina Murrmann and Ms. Hilde Zimmermann in digitizing chart-recorder strips is gratefully acknowledged.