Understanding the Hydraulics of Porous Pipes: Tradeoffs Between Water Uptake and Root Length Utilization

The water uptake region in roots is several hundred times longer than the root diameter. The distributed nature of the uptake zone requires that the hydraulic design of roots be understood by analogy to flow through a “porous pipe.” Here we present results of an analytical and experimental investigation that allowed an in-depth analysis of root hydraulic properties. Measurements on nodal maize roots confirm the nonlinear distribution of water uptake predicted by the porous pipe model. The major design parameter governing the distribution of water uptake along a porous pipe is the ratio between its axial and radial hydraulic resistance. However, total flow is proportional to the pipe's overall resistance. These results suggest the existence of a tradeoff between the effective utilization of root length and the total capacity for water uptake.     

Root Water Uptake, Leaf Water Storage and Gas Exchange of a Desert Succulent: Implications for Root System Redundancy

A technique used for hydroponics was adapted to measure instantaneousroot water uptake from the soil for a leaf succulent CAM species,Agave deserti. Comparisons were made to previously modelledwater fluxes for A. deserti and to Encelia farinosa, a non-succulentC₃species. Net CO₂uptake and transpiration forA. deserti underwell-watered conditions occurred primarily at night whereasroot water uptake was relatively constant over 24 h. Leaf thicknessdecreased when transpiration commenced and then increased whenrecharge from the stem and soil occurred, consistent with previousmodels. A drought of 90 d eliminated net CO₂uptake and transpirationand reduced the water content of leaves by 62%. Rewetting theentire root system for 7 d led to a full recovery of leaf waterstorage but only 56% of maximal net CO₂uptake. Root water uptakewas maximal immediately after rewetting, which replenished rootwater content, and decreased to a steady rate by 14 d. Whenonly the distal 50% of the root system was rewetted, the timefor net CO₂uptake and leaf water storage to recover increased,but by 30 d gas exchange and leaf water storage were similarto 100% rewetting. Rewetting 10 or 20% of the root system resultedin much less water uptake; these plants did not recover leafwater storage or gas exchange by 30 d after rewetting. A redundancyin the root system of A. deserti apparently exists for dailywater uptake requirements under wet conditions but the entireroot system is required for rapid recovery from drought.Copyright1999 Annals of Botany Company Agave deserti Engelm., desert, drought, gas exchange, rewetting, roots, succulent, water uptake.     

Accumulation of Oxygen Gas in Excised Maize Root on Aging in Water, and Water Uptake by the Root

Transfer of excised maize root from wet sawdust to water causeda considerable reduction in the exudation rate of the root.After 1-day aging in water, the exudation rate increased about8-fold and the exudation continued for 3 days. Osmotic pressureof the exudate from the root decreased with time after excisionreaching almost zero in 2 days in spite of a high exudationrate. Concentrations of sugars, acids, Ca²⁺ and Mg²⁺ in theexudate decreased with the decrease of osmotic pressure, whilethe decrease in K⁺ concentration delayed and P₁ concentrationincreased. The gas content of the root, especially of O₂, increased duringaging in water. The accumulated O₂ gas may promote water uptake,because degasification of the root by evacuation induced a decreaseof water uptake. Also, the longitudinal gradient of the O₂ contentin the root coincides with the gradient of water uptake intensity. (Received February 7, 1982; Accepted July 2, 1983)     

Studies on the Genetic Model of Nitrogen Uptake by Maize Plants at Their Seedling Stage

1IntroductionThenitIOgenuptakebymaiZeplantsd~onthegeneticCharaCteristicsofwhettesUnderthecendnenvironmentcondition.AsanimPOratstageofrnabe,seedlingstagehasthehighestrateofnitrogenuptaketothedriedmatt.['J.InOrdertOprovidethetheoriticalbasisforplanningarationalschemeofbreeding,itisne~tostUdythegeneticangelofnit~nuptakebymaizeplantsattheirablingstage.2MaterialandMethodIn1994,11inbredswerechOSenastheParentstoprepucethe~of41geno~whichincludedthe6femaleparents,5maleparentSand3ocrewaccordingt…     

Water Uptake along the Length of Grapevine Fine Roots: Developmental Anatomy,Tissue-Specific Aquaporin Expression,and Pathways of Water Transport

To better understand water uptake patterns in root systems of woody perennial crops, we detailed the developmental anatomy and hydraulic physiology along the length of grapevine (Vitis berlandieri × Vitis rupestris) fine roots from the tip to secondary growth zones. Our characterization included the localization of suberized structures and aquaporin gene expression and the determination of hydraulic conductivity (Lp_r) and aquaporin protein activity (via chemical inhibition) in different root zones under both osmotic and hydrostatic pressure gradients. Tissue-specific messenger RNA levels of the plasma membrane aquaporin isogenes (VvPIPs) were quantified using laser-capture microdissection and quantitative polymerase chain reaction. Our results highlight dramatic changes in structure and function along the length of grapevine fine roots. Although the root tip lacked suberization altogether, a suberized exodermis and endodermis developed in the maturation zone, which gave way to the secondary growth zone containing a multilayer suberized periderm. Longitudinally, VvPIP isogenes exhibited strong peaks of expression in the root tip that decreased precipitously along the root length in a pattern similar to Arabidopsis (Arabidopsis thaliana) roots. In the radial orientation, expression was always greatest in interior tissues (i.e. stele, endodermis, and/or vascular tissues) for all root zones. High Lp_r and aquaporin protein activity were associated with peak VvPIP expression levels in the root tip. This suggests that aquaporins play a limited role in controlling water uptake in secondary growth zones, which contradicts existing theoretical predictions. Despite having significantly lower Lp_r, woody roots can constitute the vast majority of the root system surface area in mature vines and thus provide for significant water uptake potential.In woody perennial root systems, the majority of water uptake is often attributed to unsuberized fine roots (Kramer and Boyer, 1995), even though woody portions can constitute the vast majority of root surface area for these plants at maturity (Nightingale, 1934; Kramer and Bullock, 1966). This assumption has likely been reinforced by the fact that most studies investigating root water uptake have been done with herbaceous species, whose roots function more like the tips of woody perennials. Although unsuberized fine roots typically have a greater ability to absorb water (i.e. they are more conductive per unit of surface area), it has been shown that older suberized portions of woody taproots can still contribute significantly to root system water uptake (Kramer and Bullock, 1966; Queen, 1967; Chung and Kramer, 1975; MacFall et al., 1990, 1991). Despite this knowledge and the fact that unsuberized roots of many woody perennials are scarce or absent during periods of the growing season when peak transpiration requires much water (MacFall et al., 1991), we still know little about how suberized portions of perennial rooting systems contribute to radial water absorption across species.The composite transport model (Steudle, 2001) is a conceptual framework describing water transport into plant roots. This model posits that water is able to flow into the root via multiple parallel pathways, traveling either in the cell walls (apoplastic) and/or from cell to cell (symplastic and/or transcellular). Transport across the cell-to-cell pathway can involve water crossing plasma membranes; thus, the rate of water uptake can be influenced by the abundance and activity of aquaporins (i.e. water channels). The contribution of aquaporins to root water uptake has been the focus of numerous studies, and the absolute magnitude of this contribution appears to be highly variable, ranging from 20% to 90% across species (for review, see Javot and Maurel, 2002). Steudle (2000) suggested that radial water flow would be dominated by aquaporin regulation in heavily suberized roots, as flow through the apoplast would be minimized. The localization of aquaporins should play a critical role in defining their impact on radial water uptake across suberized and unsuberized roots. For herbaceous species, peak aquaporin mRNA and/or protein levels have been found in root tips and the endodermis, pericycle, phloem, and xylem tissues (Schäffner, 1998; Otto and Kaldenhoff, 2000; Suga et al., 2003; Fraysse et al., 2005; Knipfer et al., 2011). Few aquaporin localization studies have been conducted in woody perennials (Vandeleur et al., 2009). Recent work from our laboratory revealed a precipitous drop in aquaporin expression between the grapevine (Vitis spp. rootstocks) root tips and older root portions (Gambetta et al., 2012). These observations led to this study, where we explore patterns of aquaporin localization in Vitis species fine roots and how they intersect with the structural anatomy and patterns of suberization to affect water uptake along the root length.Hydraulic conductivity (Lp_r) of the apoplastic pathway can be altered through changes in cell wall chemistry, especially through the deposition of suberin. Suberized apoplastic barriers in plant roots include the Casparian band of the endodermis and the suberin lamella of the endodermis, exodermis, and periderm in woody species (Esau, 1977). Casparian bands and suberin lamella are solute impermeable (for review, see Peterson and Enstone, 1996), but across studies, the extent to which they impede the flow of water is highly variable (Peterson et al., 1993; Steudle et al., 1993; Peterson and Enstone, 1996; Schreiber et al., 2005). Regardless, studies support the idea that in roots there is always some flow across the cell-to-cell pathway due to apoplastic barriers and/or an osmotic component to the driving gradient (Steudle et al., 1993; Miyamoto et al., 2001; Knipfer and Fricke, 2011). In the cell-to-cell pathway, Lp_r can be altered by intrinsic plasma membrane properties, plasmodesmata (Oparka and Prior, 1992; Roberts and Oparka, 2003), and/or the abundance and activity of aquaporins. Changes in aquaporin gene expression and protein activity remain potentially dynamic and can occur within hours, while alterations of suberized apoplastic barriers are permanent and would manifest over longer developmental time frames.The total water potential gradient across a fine root can be composed of both osmotic (ΔΨOs) and hydrostatic (ΔΨHy) pressure gradients. A purely ΔΨOs requires that some portion of the pathway be cell to cell. A purely ΔΨHy should drive flow through both pathways, and the proportion of flow through the two pathways will be determined by their Lp_r. Experimentally, Lp_r generated under ΔΨHy is typically greater than Lp_r generated under ΔΨOs, typically ranging from 2-fold to more than 100-fold greater (Steudle et al., 1987; Hallgren et al., 1994; Miyamoto et al., 2001; Knipfer and Fricke, 2011). In some cases, Lp_r is nearly equal under both types of gradients (Miyamoto et al., 2001; Knipfer and Fricke, 2011). These results suggest that if Lp_r through the apoplast were to be reduced by the presence of an apoplastic barrier, this would force flow across a cell-to-cell pathway regardless of the driving gradient (Steudle, 2000).In this study, we sought to provide a more detailed understanding of the localization of aquaporin expression and its contribution to radial water uptake in different zones of grapevine fine roots, from the unsuberized actively growing root tip to portions of the fine root undergoing secondary growth and containing a developed periderm. We characterized the developmental anatomy along the length of the fine root, including the localization of suberized structures, and quantified tissue-specific mRNA levels of plasma membrane aquaporin isogenes via a combination of laser-capture microdissection (LCM) and quantitative PCR. Finally, we determined the Lp_r of root tips and secondary growth root zones under both ΔΨOs and ΔΨHy while investigating the contribution of aquaporin activity to Lp_r via chemical inhibition.     

Upscaling from Rhizosphere to Whole Root System: Modelling the Effects of Phospholipid Surfactants on Water and Nutrient Uptake

While the rhizosphere presents a different chemical, physical and biological environment to bulk soil, most experimental and modelling investigations of plant growth and productivity are based on bulk soil parameters. In this study, water and nutrient acquisition by wheat (Triticum aestivum L.) roots was investigated using rhizosphere- and root-system-scale modelling. The physical and chemical properties of rhizosphere soil could be influenced by phospholipid surfactants in the root mucilage. Two models were compared: a 2-dimensional (2D) Finite Element Method rhizosphere model, and a 3-dimensional (3D) root architecture model, ROOTMAP. ROOTMAP was parameterised to reproduce the results of the detailed 2D model, and was modified to include a rhizosphere soil volume. Lecithin (a phospholipid surfactant) could be exuded into the rhizosphere soil volume, decreasing soil water content and hydraulic conductivity at any given soil water potential, and decreasing phosphate adsorption to soil particles. The rhizosphere-scale modelling (5 × 5 mm² soil area, 10 mm root length, uptake over 12 h) predicted a reduction in water uptake (up to 16% at 30 kPa) and an increase in phosphate uptake (up to 4%) with lecithin exudation into the rhizosphere, but little effect on nitrate uptake, with only a small reduction in dry soil (1.6% at 200 kPa). The 3D root model reproduced the water (y = 1.013x, R² = 0.996), nitrate (y = 1x, R² = 1) and phosphate (y = 0.978x, R² = 0.998) uptake predictions of the rhizosphere model, providing confidence that a whole root system model could reproduce the dynamics simulated by a Finite Element Method rhizosphere model. The 3D root architecture model was then used to scale-up the rhizosphere dynamics, simulating the effect of lecithin exudation on water, nitrate and phosphate acquisition by a wheat root system, growing over 41 d. When applied to growing and responsive roots, lecithin exudation increased P acquisition by up to 13% in nutrient-rich, and 49% in relatively nutrient-poor soil. A comparison of wheat (Triticum aestivum L.) and lupin (Lupinus angustifolius L.) root architectures, suggested an interaction between the P acquisition benefit of rhizosphere lecithin and root architecture, with the more highly-branched wheat root structure acquiring relatively more P in the presence of lecithin than the sparsely-branched lupin root system.     

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     

Focus Issue on Roots: Visualization of Root Water Uptake: Quantification of Deuterated Water Transport in Roots Using Neutron Radiography and Numerical Modeling

Our understanding of soil and plant water relations is limited by the lack of experimental methods to measure water fluxes in soil and plants. Here, we describe a new method to noninvasively quantify water fluxes in roots. To this end, neutron radiography was used to trace the transport of deuterated water (D₂O) into roots. The results showed that (1) the radial transport of D₂O from soil to the roots depended similarly on diffusive and convective transport and (2) the axial transport of D₂O along the root xylem was largely dominated by convection. To quantify the convective fluxes from the radiographs, we introduced a convection-diffusion model to simulate the D₂O transport in roots. The model takes into account different pathways of water across the root tissue, the endodermis as a layer with distinct transport properties, and the axial transport of D₂O in the xylem. The diffusion coefficients of the root tissues were inversely estimated by simulating the experiments at night under the assumption that the convective fluxes were negligible. Inverse modeling of the experiment at day gave the profile of water fluxes into the roots. For a 24-d-old lupine (Lupinus albus) grown in a soil with uniform water content, root water uptake was higher in the proximal parts of lateral roots and decreased toward the distal parts. The method allows the quantification of the root properties and the regions of root water uptake along the root systems.Understanding how and where plant roots extract water from soil remains an open question for both plant and soil scientists. One of the open questions concerns the locations of water uptake along the root system (Frensch and Steudle, 1989; Doussan et al., 1998; Steudle, 2000; Zwieniecki et al., 2003; Javaux et al., 2008). A motivation of these studies is that a better prediction of root water uptake may help to optimize irrigation and identify optimal traits to capture water. Despite its importance, there is little experimental information on the spatiotemporal distribution of the uptake zone along roots growing in soil. The lack of experimental data is largely due to the technical difficulties in measuring water fluxes in soils and roots.Quantitative information on the rate and location of root water uptake along roots growing in soil is needed to better understand the function of roots in extracting water from the soil and tolerating drought events. Such information may show which parts of roots are more involved in water extraction and how root hydraulic properties change during root growth and exposure to water-limiting conditions. For instance, it is not clear how root anatomy and the hydraulic conductivity of roots change as the soil becomes dry or the transpiration demand increases. Quantitative information of the location of root water uptake can be used to estimate the spatial distribution of hydraulic conductivities along roots. This information is needed to parameterize the most recent and advanced models of root water uptake, such as those of Doussan et al. (1998) and Javaux et al. (2008).Most of the experimental information on the spatial distribution of water uptake is limited to roots grown in hydroponic and aeroponic cultures (Frensch and Steudle, 1989; Varney and Canny, 1993; Zwieniecki et al., 2003; Knipfer and Fricke, 2010a). These investigations substantially improved our knowledge of the mechanism of water transport in roots. However, roots grown in hydroponic and aeroponic cultures may have different properties than those of roots grown in soils. As the soil dries, the hydraulic conductivity of roots and of the root-soil interface changes and likely affects the profile of root water uptake (Blizzard and Boyer, 1980; Nobel and Cui, 1992; Huang and Nobel, 1993; McCully, 1995; North and Nobel, 1997; Carminati et al., 2011; Knipfer et al., 2011; McLean et al., 2011; Carminati, 2012).New advances in imaging techniques are opening new avenues for noninvasively studying water uptake by roots in soils (Doussan et al., 1998; Garrigues et al., 2006; Javaux et al., 2008; Pohlmeier et al., 2008; Moradi et al., 2011). Imaging methods such as x-ray computed tomography, light transmission imaging, NMR, and computed neutron radiography allow quantifying the changes of water content in the root zone with different accuracy and spatial resolution. However, due to the concomitant soil water redistribution, the local changes in soil water content are not trivially related to root uptake. Consequently, the estimation of root water uptake requires coupling the imaging methods with the modeling of water flow in the soil, which, in turn, requires accurate information on the hydraulic properties of soil and roots. An additional complexity is represented by the peculiar and only partly understood hydraulic properties of the soil in the vicinity of the roots, the so-called rhizosphere.The hydraulic properties of the rhizosphere are influenced by root and microorganism activity, soil compaction due to root growth, and the formation of air-filled gaps between soil and roots when roots shrink (Nye, 1994; North and Nobel, 1997; Carminati et al., 2010; Aravena et al., 2011; Moradi et al., 2011; Carminati, 2013; Zarebanadkouki and Carminati, 2014). To date, it has been technically difficult to quantify the hydraulic properties of the rhizosphere. Carminati et al. (2011) showed that the hydraulic properties of the first 1 to 2 mm near the root affect the profile of water content and water potential toward the root.Recently, we introduced a novel method to noninvasively trace the flow of water in soil and roots (Zarebanadkouki et al., 2012, 2013). The method combines neutron radiography and the injection of deuterated water (D₂O). Neutron radiography is an imaging technique that allows one to quantify the water distribution in thin soil samples with high accuracy and spatial resolution (Moradi et al., 2008). D₂O is an isotope of normal water. Its chemical and physical properties are similar to those of water, but in contrast to water, it is almost transparent in neutron transmission imaging (Matsushima et al., 2012). This property makes D₂O an excellent tracer for neutron imaging of water flow.In our previous experiments (Zarebanadkouki et al., 2012, 2013), D₂O was injected next to selected roots and its transport was monitored using time-series neutron radiography with a spatial resolution of 150 μm and a temporal resolution of 10 s for a duration of 2 h. We grew lupine (Lupinus albus) in aluminum containers (width of 25 cm, height of 30 cm, and thickness of 1 cm) filled with a sandy soil. The soil was partitioned into different compartments with a 1-cm layer of coarse sand acting as a capillary barrier (three vertical and four horizontal layers placed at regular intervals). The capillary barriers limited the transport of D₂O into a given region of soil and facilitated the quantification of D₂O transport into the roots. Figure 1 shows selected neutron radiographs of D₂O injection during the day and night. This figure is modified from Zarebanadkouki et al. (2013). The radiographs show that (1) the radial transport of D₂O into the roots was faster during the day than during the night and (2) the axial transport of D₂O along the roots was visible only during the day, while it was negligible at night. The differences between nighttime and daytime measurements were caused by the net flow of water induced by transpiration.Open in a separate windowFigure 1.Neutron radiographs of two samples after injection of 4 mL of D₂O during the day (A and B) and during the night (C and D). D₂O was injected in one compartment during the nighttime and in two compartments during the daytime. The images show the differences between the actual radiographs at time t and the radiograph before injection (t = 0). Brighter colors indicate lower neutron attenuation and higher D₂O-water ratio. The images show that (1) the transport of D₂O was faster during the day than during the night and (2) D₂O moved axially beyond the capillary barrier toward the shoot only during the day. Images are closeups of the original field of view of 15.75 × 15.75 cm showing the distribution of D₂O in the soil and root after D₂O injection. Figures are extracted from Zarebanadkouki et al. (2013). (A neutron radiograph of the whole sample used for daytime measurement is given in Figure 9.) [See online article for color version of this figure.]The interpretation of tracing experiments with D₂O in which water and D₂O are mixed is not straightforward (Carminati and Zarebanadkouki, 2013; Warren et al., 2013a, 2013b). To determine the convective fluxes from the radiographs, Zarebanadkouki et al. (2012, 2013) introduced a diffusion-convection model of D₂O transport in roots. The model was solved analytically. The model described the increase of the average D₂O concentration in the root with a double-exponential equation, in which the rate constants of the first and second phases were related to the transport of D₂O into the cortex and the stele of the roots. Although the model included important details of the root structure, such as different pathways of water across the root tissue, the diffusion of D₂O across the root tissue was strongly simplified. In particular, our previous model assumed that as soon as the roots were immersed in D₂O, the apoplastic free space of the root cortex was instantaneously saturated with D₂O. In other words, we assumed that all cortical cells and the root endodermis were simultaneously immersed in an identical concentration of D₂O equal to that of the soil. Additionally, we assumed that D₂O concentration inside the cortical cell and the root stele was uniform (well-stirred compartment).Although the radiographs clearly showed a significant axial transport of D₂O beyond the capillary barrier during the daytime (Fig. 1B), the model of Zarebanadkouki et al. (2013) was not capable of simulating it appropriately. Indeed, our previous model could only simulate the changes in D₂O concentration in the root segments immersed in D₂O. Since the concentration of D₂O in the root segment beyond the capillary barrier carries additional information on the axial and radial fluxes along the roots, we decided to modify our model to include such information.Another approximation of the previous model was the assumption that the radial water flow to the root was uniform along the root segment immersed in D₂O. However, Zarebanadkouki et al. (2013) found significant variations in root water uptake along the roots and suggested that root water uptake should be measured with a better spatial resolution.The objective of this study was to provide an adequate model to interpret tracing experiments with D₂O. We developed two different models to describe the transport of D₂O into roots. (1) In the first model, we described the transport of D₂O into the roots by taking into account the different pathways of water across the root tissue (i.e. the apoplastic and the cell-to-cell pathways). Although this model captures the complexity of the root structure, it requires several parameters, such as the ratio of the water flow in the apoplast over the water flow in the cell-to-cell pathway. We refer to this model as the composite transport model. (2) In the second model, we simplified the root tissue into a homogenous flow domain comprising both pathways. The latter model is a simplification of the complex root anatomy, but it has the advantage of requiring fewer parameters. We refer to this model as the simplified model.In the next sections, we introduce the two modeling approaches and run a sensitivity analysis to test whether the transport of D₂O into roots is sensitive to the parameters of the composite transport model. The question was, do we need the composite transport model to accurately estimate the water flow into the roots based on the experiments with neutron radiography? Or alternatively, can we use the simplified model to estimate the fluxes without the need of introducing several parameters?Our final goal was to develop a numerical procedure to extract quantitative information on the water fluxes and the root hydraulic properties based on the tracing experiments with neutron radiography. Based on the results of the sensitivity analysis, we chose the simplified model to simulate the experiments. By fitting the observed D₂O transport into the roots, we calculated the profiles of water flux across the roots of a 24-d-old lupine as well as the diffusion permeability of its roots.     

Characteristics of Aluminum-Phosphate-Adapted Carrot Cells: Uptake and Utilization of the Phosphate

To elucidate the mechanism responsible for the superior growthof a selected line of carrot cells (Daucus carota L. cv MS Yonsun)in medium that contained AIPO₄, kinetic studies of the uptakeof phosphate and the efficiency of utilization of phosphatewere performed with the selected cells and the wild-type cells.When the two cell lines were grown in a medium with adequatesoluble phosphate (2 mM), there was no difference between theirgrowth rates. Rates of increase in fresh weight as a functionof increasing concentration of phosphate in the medium werealso identical between the cell lines, indicating that the efficiencyof utilization of phosphate by the selected cell line was similarto that by the wild-type cells. However, rate of uptake of phosphateby the selected cells under phosphate limited conditions (20µMNaH₂PO₄ at pH 5.6) was about 5-fold higher than that by thewild-type cells. Apparent K_m values for the uptake of phosphatewere calculated to be 13.6 and 9.1 µM for the selectedand the wild-type cells, respectively. The V_max valuewas estimatedto be 88.8 nmol per g fresh weight per min for the selectedcells and 28.2 for thewild-type cells. Thus, the selected cellshas an enhanced system for uptake of phosphate wherebytherewas an increase in the rate of the uptake without any dramaticchange in the affinity for phosphateions. (Received September 21, 1991; Accepted December 25, 1991)     

Na~ and Water Uptake in Relation to the Radial Reflection Coefficient of Root in Arrowleaf Saltbush Under Salt Stress

The response of halophyte arrowleaf saltbush(Atriplex triangularis Willd)plants to a gradient of salt stress were investigatedwith hydroponically cultured seedlings.Under salt stress,both the Na~ uptake into root xylem and negative pressures inxylem vessels increased with the elevation of salinity(up to 500 mol/m~3)in the root environment.However,the increment innegative pressures in root xylem far from matches the decrease in the osmotic potential of the root bathing solutions,evenwhen the osmotic potential of xylem sap is taken into consideration.The total water potential of xylem sap in arrowleafsaltbush roots was close to the osmotic potential of root bathing solutions when the salt stress was low,but a progressivelyincreased gap between the water potential of xylem sap and the osmotic potential of root bathing solutions was observedwhen the salinity in the root environment was enhanced.The maximum gap was 1.4 MPa at a salinity level of 500 mol/m~3without apparent dehydration of the tested plants.This discrepancy could not be explained with the current theories inplant physiology.The radial reflection coefficient of root in arrowleaf saltbush decreased with the enhanced salt stress wasand accompanied by an increase in the Na~ uptake into xylem sap.However,the relative Na~ in xylem exudates based onthe corresponding NaCl concentration in the root bathing solutions showed a tendency of decrease.The results showedthat the reduction in the radial reflection coefficient of roots in the arrowleaf saltbush did not lead to a mass influx of NaClinto xylem when the radial reflection coefficient of the root was considerably small;and that arrowleaf saltbush could usesmall xylem pressures to counterbalance the salt stresses,either with the uptake of large amounts of salt,or with thedevelopment of xylem pressures dangerously negative.This strategy could be one of the mechanisms behind the highresistance of arrowleaf saltbush plants to salt stress.     

The Effects of Nutrient Deficiency and Rust Infection on the Relationship Between Root Dry Weight and Length in Groundsel (Senecio vulgaris L.)

Groundsel (Senecio vulgaris L.) was grown in sand culture ata range of nutrient concentrations. Except when nutrient deficiencywas severe, infection by the rust fungus Puccinia lagenophoraeCooke substantially reduced root dry weight but had little effecton root length. Thus, specific root length (SRL, cm root mg^–1d. wt) was significantly increased in rust-infected plants.The inhibition of root dry weight caused by rust infection wasmost pronounced late in development, especially after floweringwhen, in control plants, root elongation but not dry weightaccumulation ceased. In rusted plants, and in all plants subjectedto severe nutrient deficiency, dry weight accumulation in theroots ceased concurrently with root elongation. Late in developmentat high nutrient concentration adventitious roots with low SRLswere produced. However, infection did not modify the productionof such roots and increases in SRL could not be attributed tochanges in any single type of root. There was an inverse relationship between SRL and root diameter.This relationship was unaffected by rust infection whilst nutrientdeficiency changed only its intercept: at a given SRL rootsof nutrient stressed plants were thinner than those of plantswith adequate nutrient supply. Thus, the smaller diameter ofroots of nutrient-stressed plants occurred independently ofmeasured changes in SRL but, in the absence of nutrient stress,the decrease in root diameter caused by rust was closely relatedto increases in SRL. Changes in the root: length relationships in rusted plants mayhave important implications for root activity in the field.In view of the reported changes in SRL, inhibition of root growthin terms of dry weight may be a poor indicator of potentialchanges in activity. Senecio vulgaris, rust infection, nutrient deficiency, root weight: length ratio, root diameter     

Determination of the Relationship between Nutrient Uptake Rate and Solution Concentration at the Root Surface under Field Conditions: 32P-Orthophosphate Uptake by Apple Roots

The relationships between the rates of uptake of ³²P-labelledorthophosphate per unit length, surface area and volume of rootand the concentration of ³²P-orthophosphate in solution at theroot surface were determined in a solution depletion experimentconducted in a root laboratory, using a part of the currentseason's roots of a 3.5-year-old composite apple tree growingunder field conditions. The results are compared with thoseof a previous experiment on young M.9 rootstocks grown in controlledenvironment. The rate of P uptake per unit root of the field-grown tree increasedapproximately linearly with P concentration in the externalsolution over the range 0.8–10.0 mmol m^–3, confirmingthe results of the growth cabinet experiment However, at anygiven external concentration, uptake rate per unit root of thefield-grown tree was lower than that observed in the growthcabinet experiment. Possible reasons for this difference arediscussed.     

玉米根冠细胞的脱落及其对根与根际关系的影响

以光学与电子显微镜技术观察了玉米幼苗根冠细胞脱落进程与胞间关系的变化。发现随着外沿细胞的脱离,固有的胞间连丝渐次伸展、孔径扩展,进而断裂为半连丝。半连丝内侧端仍与质膜保持固有的连接;外侧端向壁外伸展,直接与根际介质沟通,从而在根冠共质体与根际质外体之间建立了动态性的单向共质联系。以成列囊泡在半连丝中存在与由断口端溢出、半连丝呈现活跃ＡＴＰ酶活性、外源不透膜示踪物向胞内原生质掺入以及原生质的穿壁等实例证明,半连丝仍具生理活性,可作为根冠共质体与根际介质间物质与信息交流的直接通道。     

过表达VHA-c4和VHA-c5基因对拟南芥根长的影响

为研究液泡H+-ATPase c亚基VHA-c4和VHA-c5基因在植物生长发育过程中的作用,本研究构建了拟南芥VHA-c4和VHA-c5过表达载体并转化野生型拟南芥,分别获得9个和7个T2代转基因纯合体株系。采用半定量RT-PCR方法对过表达VHA-c4和VHA-c5的转基因纯合体进行阳性鉴定,发现其mRNA表达量均高于对照。对转基因纯合体进行暗培养和正常光照培养,结果显示,黑暗条件下,所有VHA-c4转基因株系的主根变短,而在正常光照下,所有VHA-c5转基因株系的主根变短,推测VHA-c4和VHA-c5分别在黑暗和光照条件下影响植物根的生长。用ABA和糖(葡萄糖和蔗糖)处理转基因纯合体,结果显示它们与野生型的表型无明显差异,表明VHA-c4和VHA-c5基因的过表达没有影响拟南芥对ABA和糖的响应。     

低磷条件下不同基因型小麦幼苗对磷的吸收和利用效率及水分的影响

 选用在土壤磷水平为5～7mgP·kg-1土的条件下筛选出来的不同磷效率的4个冬小麦品种,采用盆栽试验研究了有效磷为3．2mgP·kg-1土时的磷效率、磷吸收效率、磷利用效率及土壤水分对这些指标的影响。结果表明：在有效磷很低的土壤上,“磷高效”品种小偃54和81(85)5—3—3—3在幼苗期并未表现出较高的磷效率。尽管这两个品种的磷吸收效率显著地高于NC37和京411,但由于它们的磷利用效率相对低于京411,从而使磷效率并未显著地提高。土壤水分对4个品种的磷吸收效率和利用效率均有显著影响。     

利用蚕豆根尖微核技术监测韶关市区河段水质的研究

利用蚕豆根尖细胞微核技术监测韶关市区河段水质污染状况,测定6个断面水样的蚕豆根尖细胞微核千分率(MCN‰)及污染指数(PI),并进行F检验。结果表明,各断面水样均能引起微核率升高,孟洲坝、曲江桥、武江桥、高桥、长坝和十里亭断面的微核率分别为10.73‰、7.94‰、5.98‰、5.85‰、5.78‰、4.45‰,PI分别为3.04、2.25、1.69、1.66、1.64、1.26。按照污染指数划分标准,十里亭断面基本没有污染,武江桥、高桥和长坝断面均为轻度污染,孟洲坝和曲江桥断面属于中度污染。表明除十里亭断面外,其余各断面均有不同程度的污染。     

3种湿地植物对锌的吸收分配及其与根表铁氧化物胶膜的关系

在室内培养条件下,以灯心草、茭白和美人蕉3种湿地植物为材料,研究了湿地植物对锌的吸收分配能力与根表铁氧化物胶膜之间的关系.结果表明:(1)3种湿地植物积累锌的总量大小顺序为:茭白>美人蕉>灯心草,茭白积累锌的总量是灯心草的1.79倍;它们根表铁氧化物胶膜含量表现为灯心草>茭白>美人蕉,且其间存在显著差异(P<0.05).(2)锌在湿地植物中分配比例表现为;根中锌量>地上部分锌量>根表铁氧化物胶膜上吸附锌量;锌主要积累在湿地植物根中,地上部分和根表铁氧化物胶膜上吸附的锌量无显著差异.(3)湿地植物根表铁氧化物胶膜上吸附锌的数量与湿地植物地下部分锌含量呈极显著正相关(r=0.983 5**),增加根表铁氧化物胶膜上锌的数量就能明显提高地下部分锌含量;每千克土壤加入1 g FeSO4后,3种湿地植物积累锌的总量平均增加了21%.可见,湿地植物根表铁氧化物胶膜对锌的吸附也是湿地植物固定或积累锌的重要途径之一.     

Na^＋ and Water Uptake in Relation to the Radial Reflection Coefficient of Root in Arrowleaf Saltbush Under Salt Stress

The response of halophyte arrowleaf saltbush (Atriplex triangularis Willd) plants to a gradient of salt stress were investigated with hydroponically cultured seedlings. Under salt stress, both the Na+ uptake into root xylem and negative pressures in xylem vessels increased with the elevation of salinity (up to 500 mol/m3) in the root environment. However, the increment in negative pressures in root xylem far from matches the decrease in the osmotic potential of the root bathing solutions, even when the osmotic potential of xylem sap is taken into consideration. The total water potential of xylem sap in arrowleaf saltbush roots was close to the osmotic potential of root bathing solutions when the salt stress was low, but a progressively increased gap between the water potential of xylem sap and the osmotic potential of root bathing solutions was observed when the salinity in the root environment was enhanced. The maximum gap was 1.4 MPa at a salinity level of 500 mol/m3 without apparent dehydration of the tested plants. This discrepancy could not be explained with the current theories in plant physiology. The radial reflection coefficient of root in arrowleaf saltbush decreased with the enhanced salt stress was and accompanied by an increase in the Na+ uptake into xylem sap. However, the relative Na+ in xylem exudates based on the corresponding NaCl concentration in the root bathing solutions showed a tendency of decrease. The results showed that the reduction in the radial reflection coefficient of roots in the arrowleaf saltbush did not lead to a mass influx of NaCl into xylem when the radial reflection coefficient of the root was considerably small; and that arrowleaf saltbush could use small xylem pressures to counterbalance the salt stresses, either with the uptake of large amounts of salt, or with the development of xylem pressures dangerously negative. This strategy could be one of the mechanisms behind the high resistance of arrowleaf saltbush plants to salt stress.     

树木叶片面积与叶柄长、节间长和叶倾角的关系初探

树木的叶片面积与叶柄长、节间长之间有着明显的内在联系．本文分析研究了这个联系,在适当的理想假设下通过数学建模得到了反映树木叶片面积与叶柄长、节间长和叶倾角之间的关系模型．接着,文章对模型的不合理之处进行了分析与改进,并得到叶片面积与节间长、心茎距之间的关系表达式．最后,本文利用分别采集到的13种互生和对生树木叶子上的相关数据对模型进行了检验．检验结果表明建立的模型是正确的,能够反映树木的叶片面积与叶柄长、节间长和叶倾角之间的相互数量关系．     

Root Morphology and Zn2+ Uptake Kinetics of the Zn Hyperaccumulator of Sedum alfredii Hance

Root morphology and Zn2 uptake kinetics of the hyperaccumulating ecotype (HE) and nonhyperaccumulating ecotype (NHE) of Sedum alfredii Hance were investigated using hydroponic methods and the radiotracer flux technique. The results indicate that root length, root surface area, and root volume of NHE decreased significantly with increasing Zn2 concentration in growth media, whereas the root growth of HE was not adversely affected, and was even promoted, by 500 μmol/L Zn2 . The concentrations of Zn2 in both ecotypes of S. alfredii were positively correlated with root length, root surface area and root volumes, but no such correlation was found for root diameter. The uptake kinetics for 65Zn2 in roots of both ecotypes of S. alfredii were characterized by a rapid linear phase during the first 6 h and a slower linear phase during the subsequent period of investigation. The concentration-dependent uptake kinetics of the two ecotypes of S. alfredii could be characterized by the Michaelis-Menten equation, with the Vmax for 65Zn2 influx being threefold greater in HE compared with NHE, indicating that enhanced absorption into the root was one of the mechanisms involved in Zn hyperaccumulation. A significantly larger Vmax value suggested that there was a higher density of Zn transporters per unit membrane area in HE roots.