Water Transport across Maize Roots : Simultaneous Measurement of Flows at the Cell and Root Level by Double Pressure Probe Technique

A double pressure probe technique was used to measure simultaneously water flows and hydraulic parameters of individual cells and of excised roots of young seedlings of maize (Zea mays L.) in osmotic experiments. By following initial flows of water at the cell and root level and by estimating the profiles of driving forces (water potentials) across the root, the hydraulic conductivity of individual cell layers was evaluated. Since the hydraulic conductivity of the cell-to-cell path was determined separately, the hydraulic conductivity of the cell wall material could be evaluated as well (Lp_cw = 0.3 to 6.10⁻⁹ per meter per second per megapascal). Although, for radial water flow across the cortex and rhizodermis, the apoplasmic path was predominant, the contribution of the hydraulic conductance of the cell-to-cell path to the overall conductance increased significantly from the first layer of the cortex toward the inner layers from 2% to 23%. This change was mainly due to an increase of the hydraulic conductivity of the cell membranes which was Lp = 1.9.10⁻⁷ per meter per second per megapascal in the first layer and Lp = 14 to 9.10⁻⁷ per meter per second per megapascal in the inner layers of the cortex. The hydraulic conductivity of entire roots depended on whether hydrostatic or osmotic forces were used to induce water flows. Hydrostatic Lp_r was 1.2 to 2.3.10⁻⁷ per meter per second per megapascal and osmotic Lp_r = 1.6 to 2.8.10⁻⁸ per meter per second per megapascal. The apparent reflection coefficients of root cells (σ_s) of nonpermeating solutes (KCI, PEG 6000) decreased from values close to unity in the rhizodermis to about 0.7 to 0.8 in the cortex. In all cases, however, σ_s was significantly larger than the reflection coefficient of entire roots (σ_sr). For KCI and PEG 6000, σ_sr was 0.53 and 0.64, respectively. The results are discussed in terms of a composite membrane model of the root.     

Chemical composition of apoplastic transport barriers in relation to radial hydraulic conductivity of corn roots (Zea mays L.)

The hydraulic conductivity of roots (Lp_r) of 6- to 8-d-old maize seedlings has been related to the chemical composition of apoplastic transport barriers in the endodermis and hypodermis (exodermis), and to the hydraulic conductivity of root cortical cells. Roots were cultivated in two different ways. When grown in aeroponic culture, they developed an exodermis (Casparian band in the hypodermal layer), which was missing in roots from hydroponics. The development of Casparian bands and suberin lamellae was observed by staining with berberin-aniline-blue and Sudan-III. The compositions of suberin and lignin were analyzed quantitatively and qualitatively after depolymerization (BF₃/methanol-transesterification, thioacidolysis) using gas chromatography/mass spectrometry. Root Lp_r was measured using the root pressure probe, and the hydraulic conductivity of cortical cells (Lp) using the cell pressure probe. Roots from the two cultivation methods differed significantly in (i) the Lp_r evaluated from hydrostatic relaxations (factor of 1.5), and (ii) the amounts of lignin and aliphatic suberin in the hypodermal layer of the apical root zone. Aliphatic suberin is thought to be the major reason for the hydrophobic properties of apoplastic barriers and for their relatively low permeability to water. No differences were found in the amounts of suberin in the hypodermal layers of basal root zones and in the endodermal layer. In order to verify that changes in root Lp_r were not caused by changes in hydraulic conductivity at the membrane level, cell Lp was measured as well. No differences were found in the Lp values of cells from roots cultivated by the two different methods. It was concluded that changes in the hydraulic conductivity of the apoplastic rather than of the cell-to-cell path were causing the observed changes in root Lp_r. Received: 17 March 1999 / Accepted: 22 June 1999     

Effects of NaCl and CaCl(2) on Water Transport across Root Cells of Maize (Zea mays L.) Seedlings

The effect of salinity and calcium levels on water flows and on hydraulic parameters of individual cortical cells of excised roots of young maize (Zea mays L. cv Halamish) plants have been measured using the cell pressure probe. Maize seedlings were grown in one-third strength Hoagland solution modified by additions of NaCl and/or extra calcium so that the seedlings received one of four treatments: control; +100 millimolar NaCl; +10 millimolar CaCl₂; +100 millimolar NaCl + 10 millimolar CaCl₂. From the hydrostatic and osmotic relaxations of turgor, the hydraulic conductivity (Lp) and the reflection coefficient (σ_s) of cortical cells of different root layers were determined. Mean Lp values in the different layers (first to third, fourth to sixth, seventh to ninth) of the four different treatments ranged from 11.8 to 14.5 (Control), 2.5 to 3.8 (+NaCl), 6.9 to 8.7 (+CaCl₂), and 6.6 to 7.2 · 10⁻⁷ meter per second per megapascal (+NaCl + CaCl₂). These results indicate that salinization of the growth media at regular calcium levels (0.5 millimolar) decreased Lp significantly (three to six times). The addition of extra calcium (10 millimolar) to the salinized media produced compensating effects. Mean cell σ_s values of NaCl ranged from 1.08 to 1.16, 1.15 to 1.22, 0.94 to 1.00, and 1.32 to 1.46 in different root cell layers of the four different treatments, respectively. Some of these σ_s values were probably overestimated due to an underestimation of the elastic modulus of cells, σ_s values of close to unity were in line with the fact that root cell membranes were practically not permeable to NaCl. However, the root cylinder exhibited some permeability to NaCl as was demonstrated by the root pressure probe measurements that resulted in σ_sr of less than unity. Compared with the controls, salinity and calcium increased the root cell diameter. Salinized seedlings grown at regular calcium levels resulted in shorter cell length compared with control (by a factor of 2). The results demonstrate that NaCl has adverse effects on water transport parameters of root cells. Extra calcium could, in part, compensate for these effects. The data suggest a considerable apoplasmic water flow in the root cortex. However, the cell-to-cell path also contributed to the overall water transport in maize roots and appeared to be responsible for the decrease in root hydraulic conductivity reported earlier (Azaizeh H, Steudle E [1991] Plant Physiol 97: 1136-1145). Accordingly, the effect of high salinity on the cell Lp was much larger than that on root Lp_r.     

Effects of Salinity on Water Transport of Excised Maize (Zea mays L.) Roots

The root pressure probe was used to determine the effects of salinity on the hydraulic properties of primary roots of maize (Zea mays L. cv Halamish). Maize seedlings were grown in nutrient solutions modified by additions of NaCl and/or extra CaCl₂ so that the seedlings received one of four treatments: Control, plus 100 millimolar NaCl, plus 10 millimolar CaCl₂, plus 100 millimolar NaCl plus 10 millimolar CaCl₂. The hydraulic conductivities (Lp_r) of primary root segments were determined by applying gradients of hydrostatic and osmotic pressure across the root cylinder. Exosmotic hydrostatic Lp_r for the different treatments were 2.8, 1.7, 2.8, and 3.4·10⁻⁷ meters per second per megapascals and the endosmotic hydrostatic Lp_r were 2.4, 1.5, 2.7, and 2.3·10⁻⁷ meters per second per megapascals, respectively. Exosmotic Lp_r of the osmotic experiments were 0.55, 0.38, 0.68, and 0.60·10⁻⁷ meters per second per megapascals and the endosmotic Lp_r were 0.53, 0.21, 0.56, and 0.54·10⁻⁷ meters per second per megapascals, respectively. The osmotic Lp_r was significantly smaller (4-5 times) than hydrostatic Lp_r. However, both hydrostatic and osmotic Lp_r experiments showed that salinization of the growth media at regular (0.5 millimolar) calcium levels decreased the Lp_r significantly (30-60%). Addition of extra calcium (10 millimolar) to the salinized media caused ameliorative effects on Lp_r. The low Lp_r values may partially explain the reduction in root growth rates caused by salinity. High calcium levels in the salinized media increased the relative availability of water needed for growth. The mean reflection coefficients of the roots using NaCl were between 0.64 and 0.73 and were not significantly different for the different treatments. The mean values of the root permeability coefficients to NaCl of the different treatments were between 2.2 and 3.5·10⁻⁹ meters per second and were significantly different only in one of four treatments. Cutting the roots successively from the tip and measuring the changes in the hydraulic resistance of the root as well as staining of root cross-sections obtained at various distances from the root tip revealed that salinized roots had mature xylem elements closer to the tip (5-10 millimeters) compared with the controls (30 millimeters). Our results demonstrate that salinity has adverse effects on water transport and that extra calcium can, in part, compensate for these effects.     

Water transport in maize roots : measurement of hydraulic conductivity, solute permeability, and of reflection coefficients of excised roots using the root pressure probe

A root pressure probe has been used to measure the root pressure (P_r) exerted by excised main roots of young maize plants (Zea Mays L.). Defined gradients of hydrostatic and osmotic pressure could be set up between root xylem and medium to induce radial water flows across the root cylinder in both directions. The hydraulic conductivity of the root (Lp_r) was evaluated from root pressure relaxations. When permeating solutes were added to the medium, biphasic root pressure relaxations were observed with water and solute phases and root pressure minima (maxima) which allowed the estimation of permeability (P_Sr) and reflection coefficients (σ_sr) of roots. Reflection coefficients were: ethanol, 0.27; mannitol, 0.74; sucrose, 0.54; PEG 1000, 0.82; NaCl, 0.64; KNO₃, 0.67, and permeability coefficients (in 10⁻⁸ meters per second): ethanol, 4.7; sucrose, 1.6; and NaCl, 5.7. Lp_r was very different for osmotic and hydrostatic gradients. For hydrostatic gradients Lp_r was 1·10⁻⁷ meters per second per megapascal, whereas in osmotic experiments the hydraulic conductivity was found to be an order of magnitude lower. For hydrostatic gradients, the exosmotic Lp_r was about 15% larger than the endosmotic, whereas in osmotic experiments the polarity in the water movement was reversed. These results either suggest effects of unstirred layers at the osmotic barrier in the root, an asymmetrical barrier, and/or mechanical effects. Measurements of the hydraulic conductivity of individual root cortex cells revealed an Lp similar to Lp_r (hydrostatic). It is concluded that, in the presence of external hydrostatic gradients, water moves primarily in the apoplast, whereas in the presence of osmotic gradients this component is much smaller in relation to the cell-to-cell component (symplasmic plus transcellular transport).     

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     

Axial and Radial Hydraulic Resistance to Roots of Maize (Zea mays L.)

A root pressure probe was employed to measure hydraulic properties of primary roots of maize (Zea mays L.). The hydraulic conductivity (Lp_r) of intact root segments was determined by applying gradients of hydrostatic and osmotic pressure across the root cylinder. In hydrostatic experiments, Lp_r was constant along the segment except for an apical zone of approximately 20 millimeters in length which was hydraulically isolated due to a high axial resistance. In osmotic experiments, Lp_r decreased toward the base of the roots. Lp_r (osmotic) was significantly smaller than Lp_r (hydrostatic). At various distances from the root tip, the axial hydraulic resistance per unit root length (R_x) was measured either by perfusing excised root segments or was estimated according to Poiseuille's law from cross-sections. The calculated R_x was smaller than the measured R_x by a factor of 2 to 5. Axial resistance varied with the distance from the apex due to the differentiation of early metaxylem vessels. Except for the apical 20 millimeters, radial water movement was limiting water uptake into the root. This is important for the evaluation of Lp_r of roots from root pressure relaxations. Stationary water uptake into the roots was modeled using measured values of axial and radial hydraulic resistances in order to work out profiles of axial water flow and xylem water potentials.     

Mercurial-sensitive water transport in barley roots

An isolated barley root was partitioned into the apical and basal part across the partition wall of the double-chamber osmometer. Transroot water movement was induced by subjecting the apical part to a sorbitol solution, while the basal part with the cut end was in artificial pond water. The rate of transroot osmosis was first low but enhanced by two means, infilitration of roots by pressurization and repetition of osmosis. Both effects acted additively. The radial hydraulic conductivity (Lp_r) was calculated by dividing the initial flow rate with the surface area of the apical part of the root, to which sorbitol was applied, and the osmotic gradient between the apical and basal part of the root. Lp_r which was first 0.02–0.04 pm s⁻¹ Pa⁻¹ increased up to 0.25–0.4 pm s⁻¹ Pa⁻¹ after enhancement. Enhancement is assumed to be caused by an increase of the area of the plasma membrane which is avallable to osmotic water movement. The increased Lp_r is in the same order of magnitude as the hydraulic conductivity (Lp) of epidermal and cortical cells of barley roots obtained by Steudie and Jeschke (1983). HgCl₂, a potent inhibitor of water channels, suppressed Lp_r of non-infiltrated and infiltrated roots down to 17% and 8% of control values, respectively. A high sensitivity of Lp_r to HgCl₂ suggests that water channels constitute the most conductive pathway for osmotic radial water movement in barley roots.     

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     

Do root hydraulic properties change during the early vegetative stage of plant development in barley (Hordeum vulgare)?

Background and Aims

As annual crops develop, transpirational water loss increases substantially. This increase has to be matched by an increase in water uptake through the root system. The aim of this study was to assess the contributions of changes in intrinsic root hydraulic conductivity (Lp, water uptake per unit root surface area, driving force and time), driving force and root surface area to developmental increases in root water uptake.

Methods

Hydroponically grown barley plants were analysed during four windows of their vegetative stage of development, when they were 9–13, 14–18, 19–23 and 24–28 d old. Hydraulic conductivity was determined for individual roots (Lp) and for entire root systems (Lp_r). Osmotic Lp of individual seminal and adventitious roots and osmotic Lp_r of the root system were determined in exudation experiments. Hydrostatic Lp of individual roots was determined by root pressure probe analyses, and hydrostatic Lp_r of the root system was derived from analyses of transpiring plants.

Key Results

Although osmotic and hydrostatic Lp and Lp_r values increased initially during development and were correlated positively with plant transpiration rate, their overall developmental increases (about 2-fold) were small compared with increases in transpirational water loss and root surface area (about 10- to 40-fold). The water potential gradient driving water uptake in transpiring plants more than doubled during development, and potentially contributed to the increases in plant water flow. Osmotic Lp_r of entire root systems and hydrostatic Lp_r of transpiring plants were similar, suggesting that the main radial transport path in roots was the cell-to-cell path at all developmental stages.

Conclusions

Increase in the surface area of root system, and not changes in intrinsic root hydraulic properties, is the main means through which barley plants grown hydroponically sustain an increase in transpirational water loss during their vegetative development.     

Responses of transpiration and hydraulic conductance to root temperature in nitrogen- and phosphorus-deficient cotton seedlings

Suboptimal N or P availability and cool temperatures all decrease apparent hydraulic conductance (L) of cotton (Gossypium hirsutum L.) roots. The interaction between nutrient status and root temperature was tested in seedlings grown in nutrient solutions. The depression of L (calculated as the ratio of transpiration rate to absolute value of leaf water potential [Ψ_w]) by nutrient stress depended strongly on root temperature, and was minimized at high temperatures. In fully nourished plants, L was high at all temperatures ≥20°C, but it decreased greatly as root temperature approached the chilling threshold of 15°C. Decreasing temperature lowered Ψ_w first, followed by transpiration rate. In N- or P-deficient plants, L approached the value for fully nourished plants at root temperatures ≥30°C, but it decreased almost linearly with temperature as roots were cooled. Nutrient effects on L were mediated only by differences in transpiration, and Ψ_w was unaffected. The responses of Ψ_w and transpiration to root cooling and nutrient stress imply that if a messenger is transmitted from cooled roots to stomata, the messenger is effective only in nutrient-stressed plants.     

Water and solute transport along developing maize roots

Hydraulic and osmotic properties were measured along developing maize (Zea mays L.) roots at distances between 15 and 465 mm from the root tip to quantify the effects of changes in root structure on the radial and longitudinal movement of water and solutes (ions). Root development generated regions of different hydraulic and osmotic properties. Close to the root tip, passive solute permeability (root permeability coefficient, P_sr) was high and selectivity (root reflection coefficient, _sr) low, indicative of an imperfect semipermeable root structure. Within the apical 100–150 mm, P_sr decreased by an order of magnitude and _sr increased significantly. Root hydraulic conductivity (Lp_r) depended on the nature of the force (hydrostatic and osmotic). Osmotic Lp_r was smaller by an order of magnitude than hydrostatic Lp_r and decreased with increasing distance from the root tip. Throughout the root, responses in turgor of cortical cells and late metaxylem to step changes in xylem pressure applied to the base of excised roots were measured at high spatial resolution. The resulting profiles of radial and longitudinal propagation of pressure showed that the endodermis had become the major hydraulic barrier in older parts of the root, i.e. at distances from the apex ä 150 mm. Other than at the endodermis, no significant radial hydraulic resistance could be detected. The results permit a detailed analysis of the root's composite structure which is important for its function in collecting and translocating water and nutrients.Abbreviations and Symbols CPP cell pressure probe - IT root segments with intact tips; - Lp_r root hydraulic conductivity - Lp_rh hydrostatic hydraulic conductivity of root - Lp_ro osmotic hydraulic conductivity of root - P_app hydrostatic pressure applied to cut end of root - P_c cell turgor - P_{c, cor} turgor of cortical cell - P_c,xyl turgor of late metaxylem vessel - P_ro stationary root pressure - P_r0,seal stationary root pressure of sealed root segment - P_sr solute permeability coefficient of root - RPP root pressure probe - TR root segments with tip removed - _sr reflection coefficient of root Dedicated to Professor Andreas Sievers on the occasion of his retirement     

Aluminium and the Hydraulic Conductivity of Quercus rubra L. Root Systems

Two hydroponic experiments were conducted to determine the effectsof brief and prolonged AI³⁺ exposures on the hydraulic conductivity(Lp) of northern red oak (Quercus rubra L.) root systems. RootLp was determined using the pressure chamber method of Fiscus(1977). In the first experiment, 28- to 40-d-old seedlings weretreated for 4 d with complete nutrient solutions containingone of three Al concentrations (0.04, 1.85 or 3.71 mol m^–3)and either 0 or 50 mmol m^–3 P. Neither Lp nor daily transpirationwas affected by treatment. In Experiment II, seedlings were grown for 48–63 d incomplete solutions containing one of three Al concentrations(0, 0.75 or 2.00 mol m^–3) and either 10 or 250 mmol m^–3Ca. Lp and leaf area to root length ratio (LA/RL) were reducedwhen (AI³⁺/ Ca²⁺), the solution activity ratio, was 2.9 andhigher. Lp and LA/RL were also negatively correlated with Alconcentration and Al/Ca concentration ratio in the roots. Lpwas positively correlated with LA/RL in both experiments. Itis unclear whether Lp in the second experiment was reduced directlyby solution and root chemistry or whether Lp changed in responseto altered leaf/root balance. Key words: Al phytotoxicity, Al x Ca interaction, Quercus rubra, root hydraulic conductivity     

Roles of Morphology, Anatomy, and Aquaporins in Determining Contrasting Hydraulic Behavior of Roots

The contrasting hydraulic properties of wheat (Triticum aestivum), narrow-leafed lupin (Lupinus angustifolius), and yellow lupin (Lupinus luteus) roots were identified by integrating measurements of water flow across different structural levels of organization with anatomy and modeling. Anatomy played a major role in root hydraulics, influencing axial conductance (L_ax) and the distribution of water uptake along the root, with a more localized role for aquaporins (AQPs). Lupin roots had greater L_ax than wheat roots, due to greater xylem development. L_ax and root hydraulic conductance (L_r) were related to each other, such that both variables increased with distance from the root tip in lupin roots. L_ax and L_r were constant with distance from the tip in wheat roots. Despite these contrasting behaviors, the hydraulic conductivity of root cells (Lp_c) was similar for all species and increased from the root surface toward the endodermis. Lp_c was largely controlled by AQPs, as demonstrated by dramatic reductions in Lp_c by the AQP blocker mercury. Modeling the root as a series of concentric, cylindrical membranes, and the inhibition of AQP activity at the root level, indicated that water flow in lupin roots occurred primarily through the apoplast, without crossing membranes and without the involvement of AQPs. In contrast, water flow across wheat roots crossed mercury-sensitive AQPs in the endodermis, which significantly influenced L_r. This study demonstrates the importance of examining root morphology and anatomy in assessing the role of AQPs in root hydraulics.     

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.     

Hydraulic Resistance of Rough Lemon Roots

A pressure chamber technique was used to estimate hydraulic root resistance in rough lemon (Citrus jambhiri Lush.) seedlings. The effect of previous water stress on hydraulic root resistance was evaluated. A factorial 3 × 3 design with four replications was established with potted rough lemon seedlings in a growth chamber. Three water-stress treatments were applied by watering at intervals of 1, 2 and 3 days, and root resistance measurements were made after 6, 12 and 18 days of treatment. Plants that had experienced mild and severe water stress (irrigation interval of 2 and 3 days, respectively) had higher hydraulic root resistances after several drying cycles than those plants irrigated daily. Additional cycles had no significant effect. The increase in root resistance was not due to decreased root growth but apparently to changes in the permeability of the root cell membranes or to increased suberin deposition in the cell walls of the cortical cells. In a short-term experiment (1 h), temperature strongly affected water flow through rough lemon roots in the range 5 to 35°C. Temperature influenced root membrane permeability, since reduced blow could not be explained by changes in water viscosity.     

Root architecture and hydraulic conductance in nutrient deprived Pistacia lentiscus L. seedlings

Plants respond to low nutrient availability by modifying root morphology and root system topology. Root responses to nitrogen (N) and phosphorus (P) limitation may affect plant capacity to withstand water stress. But studies on the effect of nutrient availability on plant ability to uptake and transport water are scarce. In this study, we assess the effect of nitrogen and phosphorus limitation on root morphology and root system topology in Pistacia lentiscus L seedlings, a common Mediterranean shrub, and relate these changes to hydraulic conductivity of the whole root system. Nitrogen and phosphorus deprivation had no effect on root biomass, but root systems were more branched in nutrient limited seedlings. Total root length was higher in seedlings subjected to phosphorus deprivation. Root hydraulic conductance decreased in nutrient-deprived seedlings, and was related to the number of root junctions but not to other architectural traits. Our study shows that changes in nutrient availability affect seedling water use by modifying root architecture. Changes in nutrient availability should be taken into account when evaluating seedling response to drought.     

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.     

Strabismus regulates asymmetric cell divisions and cell fate determination in the mouse brain

The planar cell polarity (PCP) pathway organizes the cytoskeleton and polarizes cells within embryonic tissue. We investigate the relationship between PCP signaling and cell fate determination during asymmetric division of neural progenitors (NPs) in mouse embryos. The cortex of Lp/Lp (Loop-tail) mice deficient in the essential PCP mediator Vangl2, homologue of Drosophila melanogaster Strabismus (Stbm), revealed precocious differentiation of neural progenitors into early-born neurons at the expense of late-born neurons and glia. Although Lp/Lp NPs were easily maintained in vitro, they showed premature differentiation and loss of asymmetric distribution of Leu-Gly-Asn–enriched protein (LGN)/partner of inscuteable (Pins), a regulator of mitotic spindle orientation. Furthermore, we observed a decreased frequency in asymmetric distribution of the LGN target nuclear mitotic apparatus protein (NuMa) in Lp/Lp cortical progenitors in vivo. This was accompanied by an increase in the number of vertical cleavage planes typically associated with equal daughter cell identities. These findings suggest that Stbm/Vangl2 functions to maintain cortical progenitors and regulates mitotic spindle orientation during asymmetric divisions in the vertebrate brain.     

Apoplastic barriers,aquaporin gene expression and root and cell hydraulic conductivity in phosphate-limited sheepgrass plants

Mineral nutrient supply can affect the hydraulic property of roots. The aim of the present work on sheepgrass (Leymus chinensis L.) plants was to test whether any changes in root hydraulic conductivity (Lp; exudation analyses) in response to a growth-limiting supply of phosphate (P) are accompanied by changes in (1) cell Lp via measuring the cell pressure, (2) the aquaporin (AQP) gene expression by performing qPCR and (3) the formation of apoplastic barriers, by analyzing suberin lamella and Casparian bands via cross-sectional analyses in roots. Plants were grown hydroponically on complete nutrient solution, containing 250 µM P, until they were 31–36 days old, and then kept for 2–3 weeks on either complete solution, or transferred on solution containing 2.5 µM (low-P) or no added P (no-P). Phosphate treatments caused significant decreases in root and cell-Lp and AQP gene expression, while the formation of apoplastic barriers increased, particularly in lateral roots. Experiments using the AQP inhibitor mercury (Hg) suggested that a significant portion of radial root water uptake in sheepgrass occurs along a path involving AQPs, and that the Lp of this path is reduced under low- and no-P. It is concluded that a growth-limiting supply of phosphate causes parallel changes in (1) cell Lp and aquaporin gene expression (decrease) and (2) apoplastic barrier formation (increase), and that the two may combine to reduce root Lp. The reduction in root Lp, in turn, facilitates an increased root-to-shoot surface area ratio, which allocates resources to the root, sourcing the limiting nutrient.