Pressure-Induced Water and Solute Flow Through Plant Roots

Water and salt flows through detopped sunflower, tomato andred kidney bean roots under applied pressure were studied usinga pressure chamber. Values of J_v for these root systems weremeasured applying variable pressure on the root medium, andL_p calculated. The K, Na and Cl fluxes under applied pressure were comparedwith those in intact plants at the same water flow rates. Tento 100 times higher Na and Cl fluxes were observed through detoppedroots under pressure as compared to those in the unpressurized,intact plants. It is suggested that the roots under pressureare not completely analogous to intact plant roots, and thatpressure-induced flow may not be a reliable method of determiningcharacteristics of ion flow in roots in relation to water flow. Key words: Volume flow, Hydraulic conductivity, K selectivity     

Resistance to Water Flow in Xylem Vessels

Experimental data on flow resistances in xylem vessels withdifferent lumen wall surface sculptures are presented. The techniqueinvolved using determinable forces at menisci to pull waterthrough isolated undamaged metaxylem and protoxylem vesselswhich were empty but had water-saturated walls. In the horizontalorientation, the surface tension forces moved the water at velocitiesthat the resisting viscous forces at the vessel walls wouldallow since inertial forces were found negligible. A high speedcamera was used to determine the meniscus translation rates.Vessel diameters as well as average dimensions of the microscopicinternal surface irregularities were measured with respect toaxial position from the inlet. From these, flow resistanceswere determined in terms of dimensionless friction factor, f,as functions of Reynolds number, Re. It was found that, at certain helical ring thicknesses and spacing,resistance to flow was lowest. Deviations from these parametervalues cause dramatic increases in resistance to flow. Resultsare applicable to normal flow in plants, i.e. without meniscipresent.     

Water Movement Through Plant Roots

Mathematical analysis of the hydraulics of water movement throughplant roots, in terms of radial and axial resistances, has ledto equations which provide new insights into the effects ofthe component resistan ces on water uptake by and movement throughindividual roots and root systems. The ratio of axial to radialresistance determines the optimum length of a root and its totalresistance to water movement. The equations permit direct calculationsof the plant water potentials necessary, at the base of theplant, to sustain given flow rates through root systems withgiven characteristics. Lateral spacing and the resistance ofindividual laterals are the dominant factors determining totalflux per unit area into a root. When soil water potential increases with depth (surface layersdrier) root resistance tends to decrease with increasing flowrate; the reverse occurs when the surface is wetter than thelower layers. Calculated patterns of water movement into andthrough roots, in relation to soil water potential and flowrate through the root, indicate efflux from root to soil undercertain conditions. This is considered to reflect reality, althoughthe fluxes are probably transient or intermittent. The equations presented should be combined with equations describingwater movement through soil to define the behaviour of the wholeroot-soil system adequately.     

Shoot Resistance to Water Flow in Cotton

Studies using excised cotton (Gossypium hirsutum L.) plants,attached to a free water source and undergoing transpirationcycles, were conducted at intervals over a 2 year period inorder to quantify shoot resistance components of cotton canopies.Leaf water potential was found to be a linear function of transpirationrate at rates above 0.1 mm h^–1, so shoot resistance wasevaluated as the slope of this function. The value of 4.8 104h (0.48 MPa h mm^–1) total shoot resistance was consistentfor 1.10 m tall, well irrigated, fruit-bearing cotton plants.Further tests, with pre-wrapped and exposed leaves, revealedthat total shoot resistance was comprised of an axial component(40%) and a leaf component (60%). The total shoot resistanceof 0.48 MPa h mm^–1 is likely to be relevant for modellingcotton water relations when LWP is evaluated on exposed, topof the canopy leaves, such as in the ‘big leaf’type models. Key words: Leaf water potential, axial resistance, leaf resistance     

Resistance to Water Flow in the Seminal Roots of Wheat

Water uptake by the seminal root system of wheat is describedby a model which incorporates radial and axial resistance toflow in the root and the resistance to flow in the soil. Inan experiment where wheat plants were dependent on subsoil waterextracted from below 45 cm by one, three, or five seminal roots,calculations indicated that the axial resistance was the dominantresistance within the soil-root system and that flow rate changedpredictably with changing axial resistance. The model also indicatedthat the observed shape of the metaxylem vessel in the seminalaxes has important effects on the pattern of water uptake.     

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.     

Interaction between Osmotic- and Pressure-induced Water Flow in Plant Roots

When the pressure gradient across a root alters, there is often an apparent change in the permeability of the root to water. Fiscus (Plant Physiol. 1975. 55: 917-922) has suggested that this can be explained by a simple two-compartment model which takes into account rates of solute uptake into the xylem. A method of testing actual data against the Fiscus model is proposed; this shows that in some cases the apparent changes in permeability cannot be explained by the model. The model is not adequate to predict the amounts of solute reaching the xylem by passive drag: a three-compartment model would be more realistic.     

Xylem Hydraulic Characteristics, Water Relations and Wood Anatomy of the Resurrection PlantMyrothamnus flabellifoliusWelw.

Myrothamnus flabellifoliusWelw. is a desiccation-tolerant (‘resurrection’)plant with a woody stem. Xylem vessels are narrow (14 µmmean diameter) and perforation plates are reticulate. This leadsto specific and leaf specific hydraulic conductivities thatare amongst the lowest recorded for angiosperms (k_s0.87 kg m^-1MPa^-1s^-1;k_l3.28x10^-5kg m^-1MPa^-1s^-1, stem diameter 3 mm). Hydraulic conductivitiesdecrease with increasing pressure gradient. Transpiration ratesin well watered plants were moderate to low, generating xylemwater potentials of -1 to -2 MPa. Acoustic emissions indicatedextensive cavitation events that were initiated at xylem waterpotentials of -2 to -3 MPa. The desiccation-tolerant natureof the tissue permits this species to survive this interruptionof the water supply. On rewatering the roots pressures thatwere developed were low (2.4 kPa). However capillary forceswere demonstrated to be adequate to account for the refillingof xylem vessels and re-establishment of hydraulic continuityeven when water was under a tension of -8 kPa. During dehydrationand rehydration cycles stems showed considerable shrinking andswelling. Unusual knob-like structures of unknown chemical compositionwere observed on the outer surface of xylem vessels. These maybe related to the ability of the stem to withstand the mechanicalstresses associated with this shrinkage and swelling.Copyright1998 Annals of Botany Company cavitation, desiccation, hydraulic conductivity, refilling, resurrection plant, root pressure, xylem anatomy,Myrothamnus flabellifolius     

Membrane Potentials in the Xylem in Roots of Intact Plants

The membrane potential differences (PDs) of root cells of intact,illuminated Trifolium repens L. and Lolium perenne L. have beenmeasured. In T. repens the PDs were the same for all cell typesexcept for the xylem vessels, which were more positive, andfor some cells immediately adjacent to the xylem vessels whichwere 10 mV more negative. The mean PD for all cells was emdash164.6 ± 0.6 mV and the mean for cells adjacent to thexylem vessels with elevated PDs was 178.4 ± 2.4 mV. Whenthe electrode tip was in a xylem vessel a low but stable PD(mean = emdash 89.9 mV) was recorded. The results for L. perennewere similar except that there were no cells with elevated PDsadjacent to the xylem vessels. An inhibitor of ion transport from the root to the shoot, p-fluorophenylalanine(p-FPA), caused a depolarization of 10 mV in the cell PDs butin the xylem vessels the depolarization was 50 mV. The possibility that the elevated PDs of cells adjacent to thexylem vessels are related to the transport of ions into thevessels is discussed.     

The Interaction between Osmotic- and Pressure-induced Water Flow in Plant Roots

This paper presents a general model for coupled solute and water flow through plant roots based on the thermodynamics of irreversible processes. The model explains in a straight-forward manner such experimentally observed phenomena as changes in root resistance, increased solute flux, and apparent negative resistance, which have been reported for root systems under the influence of a hydrostatic pressure gradient. These apparent anomalies are explained on the basis of the interaction between the osmotic and hydrostatic driving forces and the well known “sweeping away” or dilution effect. We show that with a constant hydraulic conductivity the only features necessary to explain these phenomena are some type of membrane or membranelike structure and a mechanism for actively accumulating solutes.     

Osmotic Adjustment in Cotton (Gossypium hirsutum L.) Leaves and Roots in Response to Water Stress

The relative magnitude of adjustment in osmotic potential (ψ_s) of water-stressed cotton (Gossypium hirsutum L.) leaves and roots was studied using plants raised in pots of sand and grown in a growth chamber. One and three water-stress preconditioning cycles were imposed by withholding water, and the subsequent adjustment in solute potential upon relief of the stress and complete rehydration was monitored with thermocouple psychrometers. Both leaves and roots exhibited a substantial adjustment in ψ_s in response to water stress with the former exhibiting the larger absolute adjustment. The osmotic adjustment of leaves was 0.41 megapascal compared to 0.19 megapascal in the roots. The roots, however, exhibited much larger percentage osmotic adjustments of 46 and 63% in the one and three stress cycles, respectively, compared to 22 and 40% in the leaves in similar stress cycles. The osmotically adjusted condition of leaves and roots decreased after relief of the single cycle stress to about half the initial value within 3 days, and to the well-watered control level within 6 days. In contrast, increasing the number of water-stress preconditioning cycles resulted in significant percentage osmotic adjustment still being present after 6 days in roots but not in the leaves. The decrease in ψ_s of leaves persisted longer in field-grown cotton plants compared to plants of the same age grown in the growth chamber. The advantage of decreased ψ_s in leaves and roots of water-stressed cotton plants was associated with the maintenance of turgor during periods of decreasing water potentials.     

盐胁迫下大麦根系木质部压力的自调节现象

用植物木质部压力探针测定的结果表明,水培大麦幼苗根的木质部压力在环境条件恒定不变时始终保持波动,并且在受到轻度的盐胁迫和当盐胁迫解除时表现出高度的自调节现象.这种波动和自调节现象将对植物水势的测定和根的径向反射系数的测定产生很大的影响,并可能与植物的抗盐性有关.小麦根在同样条件下未表现出上述现象.     

Electrochemical Relations in the Transfer of Ions to the Xylem Sap of Maize Roots

The difference in electrical potential between the xylem exudateof isolated roots of 1-week-old maize seedlings and the culturesolution surrounding the roots has been determined togetherwith the concentrations of potassium, calcium, chloride, andsulphate in the xylem sap. The difference in electro-chemicalpotential (µ) for these ions has been calculated fromthe measurements. The effects on µ of varying the saltstatus of the roots, the composition of the culture solutionand of 2-4-dinitrophenol have been examined. µ for potassiumand chloride was always positive, implying that movement ofthese ions to the xylem sap was under metabolic constraint.However, pretreatment of the maize seedlings with potassiumchloride and increasing levels of dinitrophenol in the culturesolution over the exudation period caused little or no significantchange in µ for potassium although the rate of movementof ions to the xylem was substantially reduced. For calcium,µ was negative with roots in dilute culture solutionsin the absence of dinitrophenol, implying that calcium couldenter the xylem by passive diffusion. Addition of dinitrophenolchanged the sign of µ and thus brought about an apparentlyactive transport of calcium.Only in the absence of competinganions was the rate of entry of chloride significantly correlatedwith µ, and no such relationship was found for potassiumor calcium individually. These results encourage doubt as towhether measurements of µ provide a valid basis for decidingto what extent the movement of individual ions depends on specific‘active’ or ‘passive’ transport processes.Thedifference in electrical potential appears to be a characteristicof the living root and to depend on the total concentrationof ions in the external solution and on their rate of transferinto the xylem sap.     

Water Flow Through Vessel Perforation Plates--A Fluid Mechanical Approach

The effect of scalariform perforation plates on the flow ofwater through plant vessels remains poorly understood. In thisstudy, a new computational tool based on finite element methodsolutions to the Navier-Stokes equation was applied to modellingfluid flow through these structures in plant vessels. Modelsdeveloped for Liriodendron tulipifera vessel elements were solvedfor cells with and without the perforation plate to study effectsof the plate on the pressure drop along the cell. Results indicatethat the pressure gradient was 5-fold greater through the platethan for regions before and after the plate. However, the perforationplate in this species accounts for only about 8% of the resistanceto flow through typical vessels because the plate influencesflow for only a short distance along the cell relative to itslength. Details of the flow characteristics through pores ofthe perforation plate are also described. Key words: Conductance, finite element method, perforation plate, vessel, water flow     

Electron Microscope Study of Cuticular Abrasion on Cotton Leaves in Relation to Water Potential Measurements

Wullschleger, S. D. and Oosterhuis, D. M. 1987. Electron microscopestudy of cuticular abrasion on cotton leaves in relation towater potential measurements.—J. exp. Bot 38: 660–667. Accurate determination of plant water potential using thermocouplepsychrometers requires vapour equilibrium between the tissuesample and the sensing psychrometer junction. Failure to achievethis equilibrium due to cuticular resistances to vapour movementmay introduce significant errors into psychrometrically measured. The effect of cuticular abrasion on W equilibration timesfor cotton (Gossypium hirsutum L.) was studied with three typesof thermocouple psychrometers, and the extent of surface andcuticular damage was determined with electron microscopy. Water vapour equilibration between the leaf and psychrometerchamber was achieved in approximately 4 h for unabraded controls,whereas abrasion of the leaf with carborundum powder consistentlyreduced equilibration times to below 2?5 h for all three typesof psychrometers. Microscopic observation of abraded leaf tissueindicated that substantial damage to surface structures occurredduring the cuticular abrasion process. Scanning electron micrographsrevealed localized cellular damage to anastomosing leaf veinsand physical disruption of both the stomatal complex and glandulartrichomes. Transverse sections viewed with a transmission electronmicroscope indicated substantial direct damage to the cuticlewith large sections of cell wall devoid of a cuticular layer.Although the exposed cell walls were intact, the lateral cellwalls were physically compressed and distorted during abrasion.In addition, the cytoplasmic and vacuolar membranes of the epidermalcells were also frequently ruptured. Evaluation of the damagefollowing abrasion indicated that the release of turgor by theaffected cells may contribute to increased sample variabilityand possibly to errors in measurements. Key words: Leaf water potential, cuticular resistance, thermocouple psychrometer     

植物根系吸水过程中根系水流阻力的变化特征

以植物根系吸水的人工模拟试验所测得的数据为依据,运用水流的电模拟原理,定理分析了不同土壤水分水平处理下植物根系吸水过程中根系水流阻力各主要分量的大小、变化规律及其相对重要性．结果表明,在同一水分水平处理中,植物根内木质部传导阻力（Ｒｃ）随生长时间的推移而减小,随土层深度的加深而增大,土根接触阻力（Ｒｓｒ）、植物根系吸收阻力（Ｒｒ）随生长时间表现出先下降后上升阶段的动态变化特征;在不同水分水平处理中,Ｒｃ、Ｒｓｒ、Ｒｒ均随土壤湿度减小而大幅度增大;在植物根系水流阻力各分量中,Ｒｒ占根系水流阻力的比例为５５％～９６％,Ｒｓｒ约占根系水流阻力的４％～４５％,而Ｒｃ仅占根系水流阻力的７×１０－６,故Ｒｒ是决定植物根系吸水速率的重要因素     

Studies on Uptake and Loss of Water by Barley Roots in Relation to Changes in Root Resistance

The uptake and loss of water by roots of young barley plantshave been measured using tracer techniques in sand culture undercontrasting conditions of transpiration and water potentialdifference. The results are compared with direct potometricmeasurements of water uptake. The apparent resistance to flowof water changed in response to differences in water potentialbetween the leaves and the root medium and in transpirationrate. We examine the significance of this in relation to netloss of water and possible mobilization of nutrients from drysoil. With the aid of a mathematical model, some considerationis given to the role of the endodermis as a barrier to diffusivemovement of water in the older parts of the root system.