MAIN ROOT-LATERAL ROOT JUNCTIONS OF TWO DESERT SUCCULENTS: CHANGES IN AXIAL AND RADIAL COMPONENTS OF HYDRAULIC CONDUCTIVITY DURING DRYING

The influence of junctions between main roots and lateral roots on water flow was investigated for the desert succulents Agave deserti and Ferocactus acanthodes during 21 d of drying in soil. Under wet conditions, the junctions did not restrict xylem water flow from lateral roots to main roots, consistent with predictions of axial conductance based on vessel diameters. Embolism caused by drying reduced such axial conductance more at the junctions than in adjoining root regions. Connective tracheary elements at the junctions were abundantly pitted and had large areas of unlignified primary wall, apparently making them more susceptible to embolism than vessels or tracheids elsewhere in the roots. Unlike the decrease in axial conductance, the overall hydraulic conductivity of the junction increased during drying because of an increase in the conductivity of the radial pathway. Despite such increases, main roots may not lose substantial amounts of water to a dry soil during drought, initially because embolism at the junctions can limit xylem flow and later because soil hydraulic conductivity decreases. Moreover, the increased root hydraulic conductivity and a potentially rapid recovery from embolism by connective tracheary elements may favor water uptake near the junctions upon soil rewetting.     

Changes in root hydraulic conductivity for two tropical epiphytic cacti as soil moisture varies

The tropical epiphytic cacti Epiphyllum phyllanthus and Rhipsalis baccifera experience extreme variations in soil moisture due to limited soil volumes and episodic rainfalls. To examine possible root rectification, whereby water uptake from a wet soil occurs readily but water loss to a dry soil is minimal, responses of root hydraulic conductivity (L_p) to soil drying and rewetting were investigated along with the underlying anatomical changes. After 30 d of soil drying, L_p decreased 50%–70% for roots of both species, primarily because increased suberization of the periderm reduced radial conductivity. Sheaths composed of soil particles, root hairs, and mucilage covered young roots and helped reduce root desiccation. Axial (xylem) conductance increased during drying due to vessel differentiation and maturation, and drought-induced embolism was relatively low. Within 4 d of rewetting, L_p for roots of both species attained predrought values; radial conductivity increased for young roots due to the growth of new branch roots initiated during drying and for older roots due to the development of radial breaks in the periderm. The decreases in L_p during drought reduced plant water loss to a dry soil, and yet maximal water uptake and transpiration occurred within a few days of rewetting, helping these epiphytes to take advantage of episodic rainfalls in a moist tropical forest.     

Water uptake and structural plasticity along roots of a desert succulent during prolonged drought

Desert succulents resume substantial water uptake within 1–2 d of the cessation of drought, but the changes in root structure and hydraulic conductivity underlying such recovery are largely unknown. In the monocotyledonous leaf succulent Agave deserti Engelm. substantial root mortality occurred only for lateral roots near the soil surface; nearly all main roots were alive at 180 d of drought. New main roots were initiated and grew up to 320 mm at soil water potentials lower than – 5·0 MPa, utilizing water from the shoot. The hydraulic conductivity of distal root regions decreased 62% by 45 d of drought and 70% thereafter. After 7 d of rewetting, root hydraulic conductivity was restored following 45 d of drought but not after 90 and 180 d. The production of new lateral roots and the renewed apical elongation of main roots occurred 7–11 d after rewetting following 180 d of drought. Hydraulic conductivity was higher in the distal region than at midroot and often increased again near the root base, where many endodermal cells lacked suberin lamellae. Suberization and xylem maturation were influenced by the availability of moisture, suggesting that developmental plasticity along a root allows A. deserti to capitalize on intermittent or heterogeneous supplies of water.     

Root growth, developmental changes in the apex, and hydraulic conductivity for Opuntia ficus-indica during drought

Developmental changes in the root apex and accompanying changes in lateral root growth and root hydraulic conductivity were examined for Opuntia ficus-indica (L.) Miller during rapid drying, as occurs for roots near the soil surface, and more gradual drying, as occurs in deeper soil layers. During 7 d of rapid drying (in containers with a 3-cm depth of vermiculite), the rate of root growth decreased sharply and most root apices died; such a determinate pattern of root growth was not due to meristem exhaustion but rather to meristem mortality after 3 d of drying. The length of the meristem, the duration of the cell division cycle, and the length of the elongation zone were unchanged during rapid drying. During 14 d of gradual drying (in containers with a 6-cm depth of vermiculite), root mortality was relatively low; the length of the elongation zone decreased by 70%, the number of meristematic cells decreased 30%, and the duration of the cell cycle increased by 36%. Root hydraulic conductivity ( L _P) decreased to one half during both drying treatments; L _P was restored by 2 d of rewetting owing to the emergence of lateral roots following rapid drying and to renewed apical elongation following gradual drying. Thus, in response to drought, the apical meristems of roots of O. ficus-indica near the surface die, whereas deeper in the substrate cell division and elongation in root apices continue. Water uptake in response to rainfall in the field can be enhanced by lateral root proliferation near the soil surface and additionally by resumption of apical growth for deeper roots.     

Hydraulic conductance and mercury-sensitive water transport for roots of Opuntia acanthocarpa in relation to soil drying and rewetting

Drought-induced changes in root hydraulic conductance (LP) and mercury-sensitive water transport were examined for distal (immature) and mid-root (mature) regions of Opuntia acanthocarpa. During 45 d of soil drying, LP decreased by about 67% for distal and mid-root regions. After 8 d in rewetted soil, LP recovered to 60% of its initial value for both regions. Axial xylem hydraulic conductivity was only a minor limiter of LP. Under wet conditions, HgCl2 (50 microM), which is known to block membrane water-transport channels (aquaporins), decreased LP and the radial hydraulic conductance for the stele (L(R, S)) of the distal root region by 32% and 41%, respectively; both LP and L(R, S) recovered fully after transfer to 2-mercaptoethanol (10 mM). In contrast, HgCl2 did not inhibit LP of the mid-root region under wet conditions, although it reduced L(R, S) by 41%. Under dry conditions, neither LP nor L(R, S) of the two root regions was inhibited by HgCl2. After 8 d of rewetting, HgCl2 decreased LP and L(R, S) of the distal region by 23% and 32%, respectively, but LP and L(R, S) of the mid-root region were unaltered. Changes in putative aquaporin activity accounted for about 38% of the reduction in LP in drying soil and for 61% of its recovery for the distal region 8 d after rewetting. In the stele, changes in aquaporin activity accounted for about 74% of the variable L(R, S) during drought and after rewetting. Thus, aquaporins are important for regulating water movement for roots of O. acanthocarpa.     

Radial Hydraulic Conductivity of Individual Root Tissues of Opuntia ficus-indica (L.) Miller as Soil Moisture Varies

The constraints on water uptake imposed by individual root tissueswere examined forOpuntia ficus-indicaunder wet, drying, andrewetted soil conditions. Root hydraulic conductivity (L_P) andaxial conductance (K_h) were measured for intact root segmentsfrom the distal region with an endodermis and from midroot witha periderm;L_Pwas then measured for each segment with successivetissues removed by dissection. Radial conductivity (L_R) wascalculated fromL_PandK_hfor the intact segment and for the individualtissues by considering the tissue conductivities in series.Under wet conditions,L_Rfor intact distal root segments was lowestfor the cortex; at midroot, where cortical cells are dead,L_Rforthe cortex was higher and no single tissue was the predominantlimiter ofL_R.L_Rfor the endodermis and the periderm were similarunder wet conditions. During 30d of soil drying,L_Rfor the distalcortex increased almost three-fold due to the death of corticalcells, whereasL_Rfor the midroot cortex was unaffected;L_Rforthe endodermis and the periderm decreased by 40 and 90%, respectively,during drying. For both root regions under wet conditions, thevascular cylinder had the highestL_R, which decreased by 50–70%during 30d of soil drying. After 3d of rewetting, new lateralroots emerged, increasingL_Rfor the tissues outside the vascularcylinder as well as increasing uptake of an apoplastic tracerinto the xylem of both the roots and the shoot. The averageL_Rforintact root segments was similar under wet and rewetted conditions,but the conductivity of the tissues outside the vascular cylinderincreased after rewetting, as did the contribution of the apoplasticpathway to water uptake. Opuntia ficus-indica; prickly pear; root hydraulic conductivity; endodermis; periderm; apoplast; lateral root emergence     

Root attributes affecting water uptake of rice (Oryza sativa) under drought

Lowland rice roots have a unique physiological response to drought because of their adaptation to flooded soil. Rice root attributes that facilitate growth under flooded conditions may affect rice response to drought, but the relative roles of root structural and functional characteristics for water uptake under drought in rice are not known. Morphological, anatomical, biochemical, and molecular attributes of soil-grown rice roots were measured to investigate the genotypic variability and genotype×environment interactions of water uptake under variable soil water regimes. Drought-resistant genotypes had the lowest night-time bleeding rates of sap from the root system in the field. Diurnal fluctuation predominated as the strongest source of variation for bleeding rates in the field and root hydraulic conductivity (Lp (r)) in the greenhouse, and was related to expression trends of various PIP and TIP aquaporins. Root anatomy was generally more responsive to drought treatments in drought-resistant genotypes. Suberization and compaction of sclerenchyma layer cells decreased under drought, whereas suberization of the endodermis increased, suggesting differential roles of these two cell layers for the retention of oxygen under flooded conditions (sclerenchyma layer) and retention of water under drought (endodermis). The results of this study point to the genetic variability in responsiveness to drought of rice roots in terms of morphology, anatomy, and function.     

Changes in Structure and Hydraulic Conductivity for Root Junctions of Desert Succulents as Soil Water Status Varies

Variations in hydraulic conductivity (L_P) and the underlying anatomical and morphological changes were investigated for main root-lateral root junctions of Agave deserti and Ferocactus acanthodes under wet, dry, and rewetted soil conditions. During 21 d of drying, L_P and radial conductivity (L_R) increased threefold to fivefold at junctions of both species. The increase in L_R was accompanied by the formation of an apoplastic pathway for radial water movement from the surface of the junction to the stele for A. deserti and by the rupture of periderm by emerging primordia of secondary lateral roots for F. acanthodes. During 7 d of rewetting, L_R decreased for junctions of A. deserti, as apoplastic water movement was not apparent, but L_R was unchanged for F. acanthodes. Axial conductance (K_h) decreased during drying for both species, largely because of embolism related to the degradation of unlignified cell wall areas in tracheary elements at the root junction. The resulting apertures in the cell walls of such elements would admit air bubbles at pressure differences of only 0.12-0.19 MPa. Rewetting restored K_h for both species, but not completely, due to blockage of xylem elements by tyloses. About 40% of the primary lateral roots of the monocotyledon A. deserti abscised during 21 d of drying. For the dicotyledon F. acanthodes, which can form new conduits in its secondary xylem, only 10% of the primary lateral roots abscised during 21 d of drying, consistent with the much greater frequency of lateral roots that persist during drought in the field compared with the case for the sympatric A. deserti.     

Drought-induced changes in soil contact and hydraulic conductivity for roots of Opuntia ficus-indica with and without rhizosheaths

Water movement between roots and soil can be limited by incomplete root–soil contact, such as that caused by air gaps due to root shrinkage, and can also be influenced by rhizosheaths, composed of soil particles bound together by root exudates and root hairs. The possible occurrence of air gaps between the roots and the soil and their consequences for the hydraulic conductivity of the root–soil pathway were therefore investigated for the cactus t Opuntia ficus-indica, which has two distinct root regions: a younger, distal region where rhizosheaths occur, and an older, proximal region where roots are bare. Resin-embedded sections of roots in soil were examined microscopically to determine root–soil contact for container-grown plants kept moist for 21 days, kept moist and vibrated to eliminate air gaps, droughted for 21 days, or droughted and vibrated. During drought, roots shrank radially by 30% and root–soil contact in the bare root region of nonvibrated containers was reduced from 81% to 31%. For the sheathed region, the hydraulic conductivity of the rhizosheath was the least limiting factor and the root hydraulic conductivity was the most limiting; for the bare root region, the hydraulic conductivity of the soil was the least limiting factor and the hydraulic conductivity of the root–soil air gap was the most limiting. The rhizosheath, by virtually eliminating root–soil air gaps, facilitated water uptake in moist soil. In the bare root region, the extremely low hydraulic conductivity of the root–soil air gap during drought helped limit water loss from roots to a drier soil.     

Location of the major barriers to water and ion movement in young roots of Zea mays L.

The main barriers to the movement of water and ions in young roots of Zea mays were located by observing the effects of wounding various cell layers of the cortex on the roots' hydraulic conductivities and root pressures. These parameters were measured with a root pressure probe. Injury to the epidermis and cortex caused no significant change in hydraulic conductivity and either no change or a slight decline in root pressure. Injury to a small area of the endodermis did not change the hydraulic conductivity but caused an immediate and substantial drop in root pressure. When large areas of epidermis and cortex were removed (15–38% of total root mass), the endodermis was always injured and root pressure fell. The hydraulic conductance of the root increased but only by a factor of 1.2–2.7. The results indicate that the endodermis is the main barrier to the radial movement of ions but not water. The major barrier to water is the membranes and apoplast of all the living tissue. These conclusions were drawn from experiments in which hydrostatic-pressure differences were used to induce water flows across young maize roots which had an immature exodermis and an endodermis with Casparian bands but no suberin lamellae or secondary walls. The different reactions of water and ions to the endodermis can be explained by the huge difference in the permeability of membranes to these substances. A hydrophobic wall barrier such as the Casparian band should have little effect on the movement of water, which permeates membranes and, perhaps, also the Casparian bands easily. However, hydrophobic wall depositions largely prevent the movement of ions. Several hours after wounding the endodermis, root pressure recovered to some extent in most of the experiments, indicating that the wound in the endodermis had been partially healed.Abbreviations Lp_r hydraulic conductivity of root; T_1/2 = half-time of water exchange between root xylem and external medium This research was supported by a grant from EUROSILVA (project no. 39473C) to E.S., and by a Bilateral Exchange Grant jointly funded by the Deutsche Forschungsgemeinschaft and the Natural Sciences and Engineering Research Council of Canada to C.A.P. We thank Mr. Burkhard Stumpf for his excellent technicial assistance.     

Aquaporins account for variations in hydraulic conductance for metabolically active root regions of Agave deserti in wet, dry, and rewetted soil

The importance of aquaporins for root hydraulic conductance (L_P) was investigated along roots of the desert succulent Agave deserti in wet, dry and rewetted soil. Water channel activity was inferred from HgCl₂‐induced reductions of L_P that were reversible by 2‐mercaptoethanol. Under wet conditions, HgCl₂ reduced L_P for the distal root region by 50% and for the root region near the shoot base by 36% but did not affect L_P for the mid‐root region. For all root regions, L_P decreased by 30–60% during 10 d in drying soil and was not further reduced by HgCl₂. After soil rewetting, L_P increased to pre‐drying values and was again reduced by HgCl₂ for the distal and the basal root regions but not the mid‐root region. For the distal region, water channels in the epidermis/exodermis made a disproportionately large contribution to radial hydraulic conductance of the intact segment; for the basal region, water channel activity was highest in the cortex and endodermis. The role of water channels was greatest in tissues in which cells were metabolically active both in the distal root region, where new apical growth occurs in wet soil, and in the basal region, which is the most likely root region to intercept light rainfall.     

Loss of water transport capacity due to xylem cavitation in roots of two CAM succulents

Loss of axial hydraulic conductance as a result of xylem cavitation was examined for roots of the Crassulacean acid metabolism (CAM) succulents Agave deserti and Opuntia ficus-indica. Vulnerability to cavitation was not correlated with either root size or vessel diameter. Agave deserti had a mean cavitation pressure of -0.93 ± 0.08 MPa by both an air-injection and a centrifugal method compared to -0.70 ± 0.02 MPa by the centrifugal method for O. ficus-indica, reflecting the greater tolerance of the former species to low water potentials in its native habitat. Substantial xylem cavitation would occur at a soil water potential of -0.25 MPa, resulting in a predicted 22% loss of conductance for A. deserti and 32% for O. ficus-indica. For an extended drought of 3 mo, further cavitation could cause a 69% loss of conductance for A. deserti and 62% for O. ficus-indica. A model of axial hydraulic flow based upon the cavitation response of these species predicted that water uptake rates are far below the maximum possible, owing to the high root water potentials of these desert succulents. Despite various shoot adaptations to aridity, roots of A. deserti and O. ficus-indica are highly vulnerable to cavitation, which partially limits water uptake in a wet soil but helps reduce water loss to a drying soil.     

Water Transport in Onion (Allium cepa L.) Roots (Changes of Axial and Radial Hydraulic Conductivities during Root Development)

The hydraulic architecture of developing onion (Allium cepa L. cv Calypso) roots grown hydroponically was determined by measuring axial and radial hydraulic conductivities (equal to inverse of specific hydraulic resistances). In the roots, Casparian bands and suberin lamellae develop in the endodermis and exodermis (equal to hypodermis). Using the root pressure probe, changes of hydraulic conductivities along the developing roots were analyzed with high resolution. Axial hydraulic conductivity (Lx) was also calculated from stained cross-sections according to Poiseuille's law. Near the base and the tip of the roots, measured and calculated Lx values were similar. However, at distances between 200 and 300 mm from the apex, measured values of Lx were smaller by more than 1 order of magnitude than those calculated, probably because of remaining cross walls between xylem vessel members. During development of root xylem, Lx increased by 3 orders of magnitude. In the apical 30 mm (tip region), axial resistance limited water transport, whereas in basal parts radial resistances (low radial hydraulic conductivity, Lpr) controlled the uptake. Because of the high axial hydraulic resistance in the tip region, this zone appeared to be "hydraulically isolated" from the rest of the root. Changes of the Lpr of the roots were determined by measuring the hydraulic conductance of roots of different length and referring these data to unit surface area. At distances between 30 and 150 mm from the root tip, Lpr was fairly constant (1.4 x 10-7 m s-1 MPa-1). In more basal root zones, Lpr was considerably smaller and varied between roots. The low contribution of basal zones to the overall water uptake indicated an influence of the exodermal Casparian bands and/or suberin lamellae in the endodermis or exodermis, which develop at distances larger than 50 to 60 mm from the root tip.     

Hydraulic redistribution in a stand of <Emphasis Type="Italic">Artemisia tridentata</Emphasis>: evaluation of benefits to transpiration assessed with a simulation model

The significance of soil water redistribution facilitated by roots (an extension of "hydraulic lift", here termed hydraulic redistribution) was assessed for a stand of Artemisia tridentata using measurements and a simulation model. The model incorporated water movement within the soil via unsaturated flow and hydraulic redistribution and soil water loss from transpiration. The model used Buckingham-Darcy's law for unsaturated flow while hydraulic redistribution was developed as a function of the distribution of active roots, root conductance for water, and relative soil-root (rhizosphere) conductance for water. Simulations were conducted to compare model predictions with time courses of soil water potential at several depths, and to evaluate the importance of root distribution, soil hydraulic conductance and root xylem conductance on transpiration rates and the dynamics of soil water. The model was able to effectively predict soil water potential during a summer drying cycle, and the rapid redistribution of water down to 1.5 m into the soil column after rainfall events. Results of simulations indicated that hydraulic redistribution could increase whole canopy transpiration over a 100-day drying cycle. While the increase was only 3.5% over the entire 100-day period, hydraulic redistribution increased transpiration up to 20.5% for some days. The presence of high soil water content within the lower rooting zone appears to be necessary for sizeable increases in transpiration due to hydraulic redistribution. Simulation results also indicated that root distributions with roots concentrated in shallow soil layers experienced the greatest increase in transpiration due to hydraulic redistribution. This redistribution had much less effect on transpiration with more uniform root distributions, higher soil hydraulic conductivity and lower root conductivity. Simulation results indicated that redistribution of water by roots can be an important component in soil water dynamics, and the model presented here provides a useful approach to incorporating hydraulic redistribution into larger models of soil processes.     

Transport of Water and Solutes across Maize Roots Modified by Puncturing the Endodermis (Further Evidence for the Composite Transport Model of the Root)

The effects of puncturing the endodermis of young maize roots (Zea mays L.) on their transport properties were measured using the root pressure probe. Small holes with a diameter of 18 to 60 [mu]m were created 70 to 90 mm from the tips of the roots by pushing fine glass tubes radially into them. Such wounds injured about 10-2 to 10-3% of the total surface area of the endodermis, which, in these hydroponically grown roots, had developed a Casparian band but no suberin lamellae. The small injury to the endodermis caused the original root pressure, which varied from 0.08 to 0.19 MPa, to decrease rapidly (half-time = 10-100 s) and substantially to a new steady-state value between 0.02 and 0.07 MPa. The radial hydraulic conductivity (Lpr) of control (uninjured) roots determined using hydrostatic pressure gradients as driving forces was larger by a factor of 10 than that determined using osmotic gradients (averages: Lpr [hydrostatic] = 2.7 x 10-7 m s-1 MPa-1; Lpr [osmotic] = 2.2 x 10-8 m s-1 MPa-1; osmotic solute: NaCl). Puncturing the endodermis did not result in measurable increases in hydraulic conductivities measured by either method. Thus, the endodermis was not rate-limiting root Lpr: apparently the hydraulic resistance of roots was more evenly distributed over the entire root tissue. However, puncturing the endodermis did substantially change the reflection ([sigma]sr) and permeability (Psr) coefficients of roots for NaCl, indicating that the endodermis represented a considerable barrier to the flow of nutrient ions. Values of [sigma]sr decreased from 0.64 to 0.41 (average) and Psr increased by a factor of 2.6, i.e. from 3.8 x 10-9 to 10.1 x 10.-9 m s-1(average). The roots recovered from puncturing after a time and regained root pressure. Measurable increases in root pressure became apparent as soon as 0.5 to 1 h after puncturing, and original or higher root pressures were attained 1.5 to 20 h after injury. However, after recovery roots often did not maintain a stable root pressure, and no further osmotic experiments could be performed with them. The Casparian band of the endodermis is discontinuous at the root tip, where the endodermis has not yet matured, and at sites of developing lateral roots. Measurements of the cross-sectional area of the apoplasmic bypass at the root tip yielded an area of 0.031% of the total surface area of the endodermis. An additional 0.049% was associated with lateral root primordia. These areas are larger than the artificial bypasses created by wounding in this study and may provide pathways for a "natural bypass flow" of water and solutes across the intact root. If there were such a pathway, either in these areas or across the Casparian band itself, roots would have to be treated as a system composed of two parallel pathways (a cell-to-cell and an apoplasmic path). It is demonstrated that this "composite transport model of the root" allows integration of several transport properties of roots that are otherwise difficult to understand, namely (a) the differences between osmotic and hydrostatic water flow, (b) the dependence of root hydraulic resistance on the driving force or water flow across the root, and (c) low reflection coefficients of roots.     

Gating of aquaporins by low temperature in roots of chilling-sensitive cucumber and chilling-tolerant figleaf gourd

Effects of low temperature (8 degrees C) on the hydraulic conductivity of young roots of a chilling-sensitive (cucumber, Cucumis sativus L.) and a chilling-resistant (figleaf gourd, Cucurbita ficifolia Bouche) crop have been measured at the levels of whole root systems (root hydraulic conductivity, Lp(r)) and of individual cortical cells (cell hydraulic conductivity, Lp). Exposure of roots to low temperature (LRT) for up to 6 d caused a stronger suberization of the endodermis in cucumber compared with figleaf gourd, but no development of exodermal Casparian bands in either species. Changes in anatomy after 6 d of LRT treatment corresponded with a reduction in hydrostatic root Lp(r) of cucumber roots by a factor of 24, and by a factor of 2 in figleaf gourd. In figleaf gourd, there was a reduction only in hydrostatic Lp(r) but not in osmotic Lp(r) suggesting that the activity of water channels was not much affected by LRT treatment in this species. Changes in cell Lp in response to chilling and recovery were similar to the root levels, although they were more intense at the root level. Activation energies (E(a)) and Q10 of water flow as measured at the cell level were high in cucumber (E(a)=109+/-13 kJ mol(-1); Q(10)=4.8+/-0.7; n=6-10 cells), but small in figleaf gourd (E(a)=11+/-2 kJ mol(-1); Q10=1.2+/-0.1; n=6-10 cells). Roots of figleaf gourd recovered better from LRT treatment than those of cucumber. In figleaf gourd, recovery (at both the root and cell level) often resulted in Lp and Lp(r) values which were even bigger than the original, i.e. there was an overshoot in hydraulic conductivity. These effects were larger for osmotic (representing the cell-to-cell passage of water) than for hydrostatic Lp(r). After a short-term (1 d) exposure to 8 degrees C followed by 1 d at 20 degrees C, hydrostatic Lp(r) of cucumber nearly recovered and that of figleaf gourd still remained higher due to the overshoot. By contrast, osmotic Lp(r) and cell Lp in both species remained high by a factor of 3 compared with the control, possibly due to an increased activity of water channels. After preconditioning of roots at LRT, increased hydraulic conductivity was completely inhibited by HgCl2 at both the root and cell levels. Different from figleaf gourd, recovery from chilling was not complete in cucumber after longer exposure to LRT. It is concluded that at LRT, both changes in the activity of aquaporins (AQPs) and alterations of root anatomy determine the water uptake in both species. The high temperature dependence of cell Lp in cucumber suggests conformational changes of AQPs during LRT treatment which result in channel closure and in a strong gating of AQP activity by low temperature. This mechanism is thought to be different from that in figleaf gourd where AQPs reacted in the conventional way, i.e. low temperature affected the mobility of water molecules in AQPs rather than their open/closed state, and Q(10) was low.     

Water uptake by seminal and adventitious roots in relation to whole-plant water flow in barley (Hordeum vulgare L.)

Prior to an assessment of the role of aquaporins in root water uptake, the main path of water movement in different types of root and driving forces during day and night need to be known. In the present study on hydroponically grown barley (Hordeum vulgare L.) the two main root types of 14- to 17-d-old plants were analysed for hydraulic conductivity in dependence of the main driving force (hydrostatic, osmotic). Seminal roots contributed 92% and adventitious roots 8% to plant water uptake. The lower contribution of adventitious compared with seminal roots was associated with a smaller surface area and number of roots per plant and a lower axial hydraulic conductance, and occurred despite a less-developed endodermis. The radial hydraulic conductivity of the two types of root was similar and depended little on the prevailing driving force, suggesting that water uptake occurred along a pathway that involved crossing of membrane(s). Exudation experiments showed that osmotic forces were sufficient to support night-time transpiration, yet transpiration experiments and cuticle permeance data questioned the significance of osmotic forces. During the day, 90% of water uptake was driven by a tension of about -0.15 MPa.     

Regulation of a plant aquaporin by a Casparian strip membrane domain protein‐like

The absorption of soil water by roots allows plants to maintain their water status. At the endodermis, water transport can be affected by initial formation of a Casparian strip and further deposition of suberin lamellas and regulated by the function of aquaporins. Four Casparian strip membrane domain protein‐like (CASPL; CASPL1B1, CASPL1B2, CASPL1D1, and CASPL1D2) were previously shown to interact with PIP2;1. The present work shows that CASPL1B1, CASPL1B2, and CASPL1D2 are exclusively expressed in suberized endodermal cells, suggesting a cell‐specific role in suberization and/or water transport regulation. When compared with wild‐type plants, and by contrast to caspl1b1*caspl1b2 double loss of function, caspl1d1*caspl1d2 double mutants showed, in some control or NaCl stress experiments and not upon abscisic acid (ABA) treatment, a weak enlargement of the continuous suberization zone. None of the mutants showed root hydraulic conductivity (Lp_r) phenotype, whether in control, NaCl, or ABA treatment conditions. The data suggest a slight negative role for CASPL1D1 and CASPL1D2 in suberization under control or salt stress conditions, with no major impact on whole root transport functions. At the molecular level, CASPL1B1 was able to physically interact with PIP2;1 and potentially could influence the regulation of aquaporins by acting on their phosphorylated form.     

Spatio-temporal Variations in Axial Conductance of Primary and First-order Lateral Roots of a Maize Crop as Predicted by a Model of the Hydraulic Architecture of Root Systems

Rates at which water can be transported along plant roots (axial pathway) vary through time, in part depending on xylem maturation. Because of experimental constraints, the dynamics of root functional heterogeneity under field conditions remains mostly uncharted territory. Recent advances in mechanistic modelling offer opportunities to bypass such experimental limitations. This paper examines the dynamics of local variations in axial conductance of primary and first-order lateral roots of a maize crop using the architecture-based modelling approach developed by Doussan et al. (Annals of Botany: 81, 213–223, 1998). Specifically, we hypothesised that points of major resistance to long distance water transfers could arise from discrepancies between the hydraulic maturity (or water carrying capacity) of main axes and branch roots. To test this assumption, spatial distributions of root axial conductance were tested after 30, 60 and 100 days at soil depths of 10, 50 and 100 cm under a maize (Zea mays L.) crop sown at a density of 8 plants m⁻². As the crop developed, the corresponding root populations encompassed ever increasing amounts of hydraulically mature first-order laterals (branch roots): after a 100-day growth period, the vast majority of laterals had reached their maximum axial conductance at all soil depths down to 100 cm. In contrast, the axial conductance of a large proportion of main axes (primary roots) remained low, even at shallow soil depths and after 100 days of growth. The imbalance between the hydraulic maturity of primary and lateral roots was most conspicuous at soil depths of 100 cm, where ~10% only of the former compared to ~80% of the latter, had reached their maximum axial conductance after a 100-day growth period.     

植物抗旱性中的补偿效应及其在农业节水中的应用

在论述植物补偿效应存在类型和研究范畴的基础上,详细评述了植物抗旱性中根系形态结构功能及地上部干物质积累、产量和水分利用效率方面的补偿效应及其影响因素,并对植物抗旱作用中补偿生长的可能生理学机制作了探讨。同时,对补偿效应在提高农业水分利用效率中的应用进行了讨论