Water and solute flow through mung bean roots under applied pressure

The technique of applying hydrostatic pressure on the root medium to study water and solute flows through excised plant roots and to study various characteristics of roots in relation to flow has been used by many workers but flows in excised roots have not been compared with those in intact transpiring plants. In the present study this comparison has been made using mung bean roots. Results show that excised roots under pressure lack the ion selectivity which is observed in intact plant roots and conduct salt many times higher than salt flows through intact plant roots. The role of stem resistance in the rates of water and salt flow through roots has been discussed. The suitability of this technique for solute flow studies through mung bean roots is questioned.     

Evidence for active water transport in a corn root preparation

Summary Water flow was measured in aZea mays root preparation consisting of a segment from which the central part had been excised. It was shown that water flow had two components, one osmotic and one non-osmotic. The non-osmotic flow was inhibited by cyanide. No correlation was found between water flow and solute flow. These findings suggest that active water transport occurred in the root preparation. The mechanism of such water movement is discussed.     

Radial Turgor and Osmotic Pressure Profiles in Intact and Excised Roots of Aster tripolium: Pressure Probe Measurements and Nuclear Magnetic Resonance-Imaging Analysis

High-resolution nuclear magnetic resonance images (using very short spin-echo times of 3.8 milliseconds) of cross-sections of excised roots of the halophyte Aster tripolium showed radial cell strands separated by air-filled spaces. Radial insertion of the pressure probe (along the cell strands) into roots of intact plants revealed a marked increase of the turgor pressure from the outermost to the sixth cortical layer (from about 0.1-0.6 megapascals). Corresponding measurements of intracellular osmotic pressure in individual cortical cells (by means of a nanoliter osmometer) showed an osmotic pressure gradient of equal magnitude to the turgor pressure. Neither gradient changed significantly when the plants were grown in, or exposed for 1 hour to, media of high salinity. Differences were recorded in the ability of salts and nonelectrolytes to penetrate the apoplast in the root. The reflection coefficients of the cortical cells were approximately 1 for all the solutes tested. Excision of the root from the stem resulted in a collapse of the turgor and osmotic pressure gradients. After about 15 to 30 minutes, the turgor pressure throughout the cortex attained an intermediate (quasistationary) level of about 0.3 megapascals. This value agreed well with the osmotic value deduced from plasmolysis experiments on excised root segments. These and other data provided conclusions about the driving forces for water and solute transport in the roots and about the function of the air-filled radial spaces in water transport. They also showed that excised roots may be artifactual systems.     

Effects of synthetic plant growth retardants and abscisic acid on root functions of Brassica rapa plants exposed to low root-zone temperature

Possible interactions of two synthetic plant-growth retardants during the short-term response of Brassica rapa L. ssp. oleifera (DC.) Metzger plants to low root-zone temperature were investigated by pretreating with mefluidide or paclobutrazol. Water and solute transfers were studied by measuring xylem sap volume flow (under root pressure exudation) and ion flow from the roots. Relations with nitrate uptake rate were also considered. Root pretreatment with paclobutrazol strongly restricted the cold-inducible processes which normally restore water and solute flow from the root xylem. Paclobutrazol decreased the rates of nitrate uptake and exudation flow from the root xylem (principally by reducing root hydraulic conductivity) with dramatic consequences for ion flow, especially that of nitrate.
The effects of root ABA pretreatment on plant response to root cooling were then studied separately or in association with a pretreatment with paclobutrazol. Despite a slight decrease in nitrate uptake rate, ABA pretreatment of the roots enabled the plant to develop rapid mechanisms for adaptation to cold constraint at the root level. Moreover, this action of exogenous ABA greatly reduced the effect of a simultaneous paclobutrazol pretreatment and partly restored water and solute flows.
Thus, the improvement of plant resistance to cold conditions brought about by treatments with mefluidide and paclobutrazol (previously shown in long-term experiments) cannot simply be explained by their short-term effects.     

Gradients of turgor, osmotic pressure, and water potential in the cortex of the hypocotyl of growing ricinus seedlings : effects of the supply of water from the xylem and of solutes from the Phloem

To evaluate the possible role of solute transport during extension growth, water and solute relations of cortex cells of the growing hypocotyl of 5-day-old castor bean seedlings (Ricinus communis L.) were determined using the cell pressure probe. Because the osmotic pressure of individual cells (πⁱ) was also determined, the water potential (ψ) could be evaluated as well at the cell level. In the rapidly growing part of the hypocotyl of well-watered plants, turgor increased from 0.37 megapascal in the outer to 1.04 megapascal in the inner cortex. Thus, there were steep gradients of turgor of up to 0.7 megapascal (7 bar) over a distance of only 470 micrometer. In the more basal and rather mature region, gradients were less pronounced. Because cell turgor ≈ πⁱ and ψ ≈ 0 across the cortex, there were also no gradients of ψ across the tissue. Gradients of cell turgor and πⁱ increased when the endosperm was removed from the cotyledons, allowing for a better water supply. They were reduced by increasing the osmotic pressure of the root medium or by cutting off the cotyledons or the entire hook. If the root was excised to interrupt the main source for water, effects became more pronounced. Gradients completely disappeared and turgor fell to 0.3 megapascal in all layers within 1.5 hours. When excised hypocotyls were infiltrated with 0.5 millimolar CaCl₂ solution under pressure via the cut surface, gradients in turgor could be restored or even increased. When turgor was measured in individual cortical cells while pressurizing the xylem, rapid responses were recorded and changes of turgor exceeded that of applied pressure. Gradients could also be reestablished in excised hypocotyls by abrading the cuticle, allowing for a water supply from the wet environment. The steep gradients of turgor and osmotic pressure suggest a considerable supply of osmotic solutes from the phloem to the growing tissue. On the basis of a new theoretical approach, the data are discussed in terms of a coupling between water and solute flows and of a compartmentation of water and solutes, both of which affect water status and extension growth.     

Responses of a CAM Plant to Drought and Rainfall: Capacitance and Osmotic Pressure Influences on Water Movement

Daily and seasonal patterns in water flow and water potentialwere investigated for the Crassulacean acid metabolism succulentAgave deserti during an extended summer drought and for a periodfollowing rainfall. Field measurements of transpiration andof osmotic pressure changes over selected 24 h periods wereused as input variables for a computer model of water flow thatwas based on an electrical circuit analog of the whole plant.Parameters such as root resistance and tissue capacitance werealso varied to reflect the effects of changing plant or soilwater status. The model predicted internal water flow and waterpotential during the drought cycle and was used to assess therole of tissue osmotic properties in water uptake from the soiland in internal water redistribution. For plants under wet soil conditions, 55% of the night-timetranspiration was derived from water storage, this storage beingrecharged during the day. As drought progressed, transpirationand the nocturnal increase in osmotic pressure declined, althoughthe osmotic pressure itself increased. The difference in osmoticpressure between the water storage tissue and the chlorenchymacaused a net flow of water into the chlorenchyma after 3 weeksof drought, thereby increasing chlorenchyma turgor pressure.Simulations also indicated that a large increase in root resistancemust occur to prevent substantial water loss from the plantto the dry soil. After rainfall, recharge of plant water storagewas complete within one week, although full recovery in theamplitude of daily osmotic pressure variations took longer. Key words: Agave deserti, transpiration, water potential, water storage     

A model for radial water transport across plant roots

Summary A model based on the canal theory (Katou andFurumoto 1986 a, b) is proposed for the absorption of solute and water at the root periphery. The present canal model in the periphery and the model which was previously proposed for the exudation in the stele (Katou et al. 1987), are organized into a model for radial transport across excised plant roots, in the light of anatomical and physiological knowledge of maize roots. The canal equations for both canals are numerically solved to give quite a good explanation for the observed exudation of maize roots. It is found that the regulation of solute transport has a primary importance in the regulation of water transport across excised roots. The internal cell pressure of the symplast adjusts the water absorption at the root periphery to the water secretion into the vessels. There seems no need for this explanation of the radial water transport across roots to assume cell membranes with low reflection coefficient or variable water permeability. It would seem that the apoplast wall layers play a crucial role in metabolic control of water transport in roots as well as in hypocotyls.Abbreviations J _s ^ex* the theoretically estimated rate of solute exudation per unit surface area of model maize roots - J that of volume exudation per unit surface area of model maize roots - the reflection coefficient of the cell membrane against solutes     

A biophysical analysis of stem and root diameter variations in woody plants

A comprehensive model of stem and root diameter variation was developed. The stem (or root) was represented using two coaxial cylinders corresponding with the mature xylem and the extensible tissues. The extensible tissues were assumed to behave as a single cell separated from the mature xylem by a virtual membrane. The mature xylem and the extensible tissues are able to dilate with temperature and grow. Moreover, the extensible tissues are able to shrink and swell according to water flow intensity. The model is mainly based on the calculation of water volume flows in the "single cell" that are described using the principles of irreversible thermodynamics. The elastic response to storage volume and plastic extension accompanying growth are described. The model simulates diameter variation due to temperature, solute accumulation, and xylem, water potential. The model was applied to the peach (Prunus persica) stem and to the plum (Prunus domestica x Prunus spinosa) root. The simulation outputs corresponded well with the diameter variation observed. The model predicts that variations of turgor pressure and osmotic potential are smaller than the variations of xylem water potential. It also demonstrates correlations between the xylem water potential, the turgor pressure, the elastic modulus, and the osmotic potential. The relationship between the diameter and the xylem water potential exhibits a substantial hysteresis, as observed in field data. A sensitivity analysis using the model parameters showed that growth and shrinkage were highly sensitive to the initial values of the turgor pressure and to the reflection coefficient of solutes. Shrinkage and growth were sensitive to elastic modulus and wall-yielding threshold pressure, respectively. The model was not sensitive to changes in temperature.     

Water uptake by plant roots: an integration of views

A COMPOSITE TRANSPORT MODEL is presented which explains the variability in the ability of roots to take up water and responses of water uptake to different factors. The model is based on detailed measurements of 'root hydraulics' both at the level of excised roots (root hydraulic conductivity, Lp_r) and root cells (membrane level; cell Lp) using pressure probes and other techniques. The composite transport model integrates apoplastic and cellular components of radial water flow across the root cylinder. It explains why the hydraulic conductivity of roots changes in response to the nature (osmotic vs. hydraulic) and intensity of water flow. The model provides an explanation of the adaptation of plants to conditions of drought and other stresses by allowing for a `coarse regulation of water uptake' according to the demands from the shoot which is favorable to the plant. Coarse regulation is physical in nature, but strongly depends on root anatomy, e.g. on the existence of apoplastic barriers in the exo- and endodermis. Composite transport is based on the composite structure of roots. A `fine regulation' results from the activity of water channels (aquaporins) in root cell membranes which is assumed to be under metabolic and other control.     

Comparison between gradient-dependent hydraulic conductivities of roots using the root pressure probe: the role of pressure propagations and implications for the relative roles of parallel radial pathways

Hydrostatic pressure relaxations with the root pressure probe are commonly used for measuring the hydraulic conductivity (Lp(r)) of roots. We compared the Lp(r) of roots from species with different root hydraulic properties (Lupinus angustifolius L. 'Merrit', Lupinus luteus L. 'Wodjil', Triticum aestivum L. 'Kulin' and Zea mays L. 'Pacific DK 477') using pressure relaxations, a pressure clamp and osmotic gradients to induce water flow across the root. Only the pressure clamp measures water flow under steady-state conditions. Lp(r) determined by pressure relaxations was two- to threefold greater than Lp(r) from pressure clamps and was independent of the direction of water flow. Lp(r) (pressure clamp) was two- to fourfold higher than for Lp(r) (osmotic) for all species except Triticum aestivum where Lp(r) (pressure clamp) and Lp(r) (osmotic) were not significantly different. A novel technique was developed to measure the propagation of pressure through roots to investigate the cause of the differences in Lp(r). Root segments were connected between two pressure probes so that when root pressure (P(r)) was manipulated by one probe, the other probe recorded changes in P(r). Pressure relaxations did not induce the expected kinetics in pressure in the probe at the other end of the root when axial hydraulic conductance, and probe and root capacitances were accounted for. An electric circuit model of the root was constructed that included an additional capacitance in the root loaded by a series of resistances. This accounted for the double exponential kinetics for intact roots in pressure relaxation experiments as well as the reduced response observed with the double probe experiments. Although there were potential errors with all the techniques, we considered that the measurement of Lp(r) using the pressure clamp was the most unambiguous for small pressure changes, and provided that sufficient time was allowed for pressure propagation through the root. The differences in Lp(r) from different methods of measurement have implications for the models describing water transport through roots and the potential role of aquaporins.     

Effect of apoplastic solutes on water potential in elongating sugarcane leaves

Solute concentration in the apoplast of growing sugarcane (Saccharum spp. hybrid) leaves was measured using one direct and several indirect methods. The osmotic potential of apoplast solution collected directly by centrifugation of noninfiltrated tissue segments ranged from −0.25 megapascal in mature tissue to −0.35 megapascal in tissue just outside the elongation zone. The presence of these solutes in the apoplast manifested itself as a tissue water potential equal to the apoplast osmotic potential. Since the tissue was not elongating, the measurements were not influenced by growth-induced water uptake and no significant tension was detected with the pressure chamber. Further evidence for a significant apoplast solute concentration was obtained from pressure exudation experiments and comparison of methods for estimating tissue apoplast water fraction. For elongating leaf tissue the centrifugation method could not be used to obtain direct measurements of apoplast solute concentration. However, several other observations suggested that the apoplast water potential of −0.35 to −0.45 megapascal in elongating tissue had a significant osmotic component and small, but significant tension component. Results of experiments in which exudate was collected from pressurized tissue segments of different ages suggested that a tissue age-dependent dynamic equilibrium existed between intra- and extracellular solutes.     

Hydraulic and osmotic properties of oak roots

Hydraulic and osmotic properties of root systems of 2.5–8-months-oldoak seedlings (Quercus robur and Q. petraea) were measured usingthe root pressure probe. Root pressures of excised roots rangedbetween 0.05 and 0.15 MPa which was similar to values obtainedfor herbaceous species. Root hydraulic conductivity (Lp_r; perunit of root surface area) was much larger in the presence ofhydrostatic than in the presence of osmotic pressure differencesdriving water flow across the roots. Differences were as largeas a factor of 20 to 470. Roots of the young seedlings of Q.robur grew more rapidly than those of Q. petraea and had a hydraulicconductivity which was substantially higher. Nitrogen nutritionaffected root growth of Q. robur more than that of Q. petraea,but did not affect root Lp_r of either species. For Q. robur,Lp_r decreased with root age (size) which is interpreted by aneffect of suberization during the development of fine roots.Root hydraulic conductance remained constant for both species.For Q. robur, this was due to the fact that the overall decreasein Lp_r was compensated for by an increase in root surface area.Root reflection coefficients (_sr) were low and ranged between_sr=0.1 and 0.5 for solutes for which cell membranes exhibitreflection coefficients of virtually unity (salts, sugars etc.).Solute permeability was small and was usually not measurablewith the technique. When root systems were attached to the rootpressure probe for longer periods of time (up to 10d), solutepermeability increased due to ageing effects which, however,did not cause a general leakiness of the roots as Lp_r decreased.Hence, values were only used from measurements taken duringthe first day. Transport properties of oak roots are comparedwith those recently obtained for spruce (Rdinger et al., 1994).They are discussed in terms of a composite transport model ofthe root which explains low root _sr at low solute permeabilityand reasonable rootLp_r The model predicts differences betweenosmotic and hydraulic water flow and differences in the transportproperties of roots of herbs and trees as found. Key words: Composite transport, hydraulic conductivity, nitrogen nutrition, Quercus, reflection coefficient, root transport, water relations     

Permeation of Uncharged Organic Molecules and Water Through Tomato Roots

Permeation of ethylene glycol, D-mannitol, L glucose, and raffinosethrough excised tomato roots was investigated. Solute transportfrom external solution to the xylem sap was determined at variousrates of exudate flow achieved in turn by application of differentpressures to the solution surrounding the roots. At an applied pressure of two bar, steady-state solute concentrationsexpressed as a percentage of external concentrations were, 52per cent for ethylene glycol, 3–4 per cent for mannitoland glucose, and I per cent for raffinose. Such relationshipsbetween the chemical structure of the solute and its concentrationin the xylem sap are similar to those demonstrated by otherworkers for solute permeation into single cells. Thus in thepresent experiments most of these solutes presumably flowedthrough at least one membrane before reaching the xylem. The data also indicate that the flow of water via wholly extracellularpathways was slight, at most, I per cent of the total flow reachingthe xylem via this route.     

An interpretation of some whole plant water transport phenomena

A treatment of water flow into and through plants to the evaporating surface of the leaves is presented. The model is driven by evaporation from the cell wall matrix of the leaves. The adsorptive and pressure components of the cell wall matric potential are analyzed and the continuity between the pressure component and the liquid tension in the xylem established. Continuity of these potential components allows linking of a root transport function, driven by the tension in the xylem, to the leaf water potential. The root component of the overall model allows for the solvent-solute interactions characteristic of a membrane-bound system and discussion of the interactions of environmental variables such as root temperature and soil water potentials. A partition function is developed from data in the literature which describes how water absorbed by the plant might be divided between transpiration and leaf growth over a range of leaf water potentials.

Relationships between the overall system conductance and the conductance coefficients of the various plant parts (roots, xylem, leaf matrix) are established and the influence of each of these discussed.

The whole plant flow model coupled to the partition function is used to simulate several possible relationships between leaf water potential and transpiration rate. The effects of changing some of the partition function coefficients, as well as the root medium water potential on these simulations is illustrated.

In addition to the general usefulness of the model and its ability to describe a wide range of situations, we conclude that the relationships used, dealing with bulk fluid flow, diffusion, and solute transport, are adequate to describe the system and that analogically based theoretical systems, such as the Ohm's law analogy, probably ought to be abandoned for this purpose.

     

Xylem Flow and its Driving Forces in a Tropical Liana: Concomitant Flow-Sensitive NMR Imaging and Pressure Probe Measurements

Abstract: Flow-sensitive NMR imaging and pressure probe techniques were used for measuring xylem water flow and its driving forces (i.e., xylem pressure as well as cell turgor and osmotic pressure gradients) in a tropical liana, Epipremnum aureum. Selection of tall specimens allowed continuous and simultaneous measurements of all parameters at various distances from the root under diurnally changing environmental conditions. Well hydrated plants exhibited exactly linearly correlated dynamic changes in xylem tension and flow velocity. Concomitant multiple-probe insertions along the plant shoot revealed xylem and turgor pressure gradients with changing magnitudes due to environmental changes and plant orientation (upright, apex-down, or horizontal). The data suggest that in upright and - to a lesser extent - in horizontal plants the transpirational water loss by the cells towards the apex during the day is not fully compensated by water uptake through the night. Thus, longitudinal cellular osmotic pressure gradients exist. Due to the tight hydraulic coupling of the xylem and the tissue cells these gradients represent (besides the transpiration-induced tension in the xylem) an additional tension component for anti-gravitational water movement from the roots through the vessels to the apex.     

Permeability of Iris germanica's multiseriate exodermis to water, NaCl, and ethanol

The exodermis of Iris germanica roots is multiseriate. Its outermost layer matures first with typical Casparian bands and suberin lamellae. But as subsequent layers mature, the Casparian band extends into the tangential and anticlinal walls of their cells. Compared with roots in which the endodermis represents the major transport barrier, the multiseriate exodermis (MEX) was expected to reduce markedly radial water and solute transport. To test this idea, precocious maturation of the exodermis was induced with a humid air gap inside a hydroponic chamber. Hydraulic conductivity (Lp(pc)) was measured on completely submerged roots (with an immature exodermis) and on air-gap-exposed root regions (with two mature exodermal layers) using a pressure chamber. Compared with regions of roots with no mature exodermal layers, the mature MEX reduced Lp(pc) from 8.5×10(-8) to 3.9×10(-8) m s(-1) MPa(-1). Puncturing the MEX increased Lp(pc) to 19×10(-8) m s(-1) MPa(-1), indicating that this layer constituted a substantial hydraulic resistance within the root (75% of the total). Alternatively, a root pressure probe was used to produce pressure transients from which hydraulic conductivity was determined, but this device measured mainly flow through the endodermis in these wide-diameter roots. The permeability of roots to NaCl and ethanol was also reduced in the presence of two mature MEX layers. The data are discussed in terms of the validity of current root models and in terms of a potential role for I. germanica MEX during conditions of drought and salt stress.     

Control of scion vigour by kiwifruit rootstocks is correlated with spring root pressure phenology

Root pressure was measured continuously over spring in eight clonal kiwifruit rootstocks selected from seven Actinidia species (A. chrysantha, A. deliciosa, A. eriantha, A. hemsleyana, A. kolomikta, A. macrosperma, A. polygama), using pressure transducers and miniature compression fittings. Rootstocks that promoted scion vigour developed root pressures up to 0.15 MPa before or during scion budburst, whereas those that reduced scion vigour developed root pressure up to 0.05 MPa only after scion shoot expansion. When several seasons were compared, the date of onset of root pressure and the magnitude of pressure achieved were consistent for each rootstock. Root pressure was first recorded between late July and early September in vigour-promoting rootstocks, while scion budburst and initial shoot growth were in late August and early September. Vigour-reducing rootstocks did not develop significant root pressure until October. The date of onset was similar for the grafted rootstock and ungrafted plant of the same clone, but was not clearly related to the timing of shoot growth by the ungrafted plant. In the grafted plants the leaf and xylem water potentials of the scion were more negative, midday turgor was 0.3-0.5 MPa lower, and wilting was sometimes observed in developing shoots growing on low-vigour rootstocks, indicating that water stress was contributing to reductions in growth. Leaf turgor was correlated with average root pressure but not pressure measured during the day, suggesting that root pressure was not supporting transpiration during peak flows and was, instead, indicative of higher root hydraulic conductance. The rapid temporal rise in root pressure observed each spring in the various rootstocks was not accompanied by changes in xylem sap solute potential, but when rootstock clones were compared those that developed higher root pressures had higher sap solute potentials. Xylem sap solute potential varied between rootstocks from -0.07 MPa to -0.15 MPa, while root pressures measured at the same time varied between 0.0 MPa and 0.09 MPa, suggesting that an osmotic mechanism could account for the observed root pressure. Differences in phenology between the rootstocks and scion appeared to account for the rootstock effects on shoot growth, and changes in root pressure provided a useful indication of seasonal changes in root hydraulic properties and solute transport behaviour.     

Metabolic inhibition of root water flow in red-osier dogwood (Cornus stolonifera) seedlings.

The short-term effects of sodium azide (NaN(3)) on water flow in red-osier dogwood (Cornus stolonifera Michx.) seedlings were examined in excised roots at a constant pressure of 0.3 MPa. NaN(3) significantly decreased root water flow rates (Q(v)). It also induced a significant reduction in root respiration and reduced stomatal conductance to a greater extent in intact seedlings than in excised shoots. Apoplastic flow of water increased with the NaN(3)-induced decreases in Q(v). Mercuric chloride (HgCl(2)) was also used to characterize the water flow responses and respiration of dogwood roots. Similarly to NaN(3), 0.1 and 0.3 mM HgCl(2) decreased root respiration rates and Q(v). The lower, 0.05 mM HgCl(2) treatment, reduced Q(v), but had no significant effect on root oxygen uptake. The reduction of Q(v) in HgCl(2)-treated plants was only partly reversed by 50 mM mercaptoethanol. The mercurial inhibition of Q(v) suggested the presence of Hg-sensitive water channels in dogwood roots. The results indicate that root-absorbed NaN(3) metabolically inhibited water channel activities in roots and in shoots and resulted in stomatal closure. It is suggested that the inhibition of respiration that occurs in plants stressed with environmental factors such as flooding, cold soils, and drought may be responsible for the closure of water channels in root cells and inhibition of root water flow.     

Water transport in barley roots

Radial transport of water in excised barley (Hordeum distichon, cv. Villa) roots was measured using a new method based on the pressure-probe technique. After attaching excised roots to the probe, root pressures of 0.9 to 2.9 bar were developed. They could be altered either by changing the root pressure artificially (with the aid of the probe) or by changing the osmotic pressure of the medium in order to induce water flows across the root. The hydraulic conductivity of the barley roots (per cm² of outer root surface) was obtained in different types of experiments (initial water flow, pressure relaxations, constant water flow) and was (0.3–4.3)·10^-7 cm s^-1 bar^-1. The hydraulic conductivity of the root was by an order of magnitude smaller than the hydraulic conductivity of the cell membranes of cortical and epidermal cells (0.8–2.2)·10^-6 cm s^-1 bar^-1. The half-times of water exchange of these cells was 1–21 s and two orders of magnitude smaller than that of entire excised roots (100–770 s). Their volumetric elastic modulus was 15–305 bar and increased with increasing turgor. Within the root cortex, turgor was independent of the position of the cell within a certain layer and turgor ranged between 3 and 5 bar. The large difference between the hydraulic conductivity of the root and that of the cell membranes indicates that there is substantial cell-to-cell (transcellular plus symplasmic) transport of water in the root. When it is assumed that 10–12 membrane layers (plasmalemma plus tonoplast) in the epidermis, cortex and endodermis form the hydraulic resistance to water flow, a value for the hydraulic conductivity of the root can be calculated which is similar to the measured value. This picture for water transport in the root contradicts current models which favour apoplasmic water transport in the cortex.