The supply of nutrient ions by diffusion to plant roots in soil

Summary Observation of soil grown roots of rye-grass shows that an approximately cylindrical volume of soil, the root hair cylinder, is densely occupied by root hairs. Estimates are given of the concentration of labile and solution potassium within the root hair cylinder during experiments measuring potassium uptake from two soils by single roots. Calculations, using a diffusion model, suggest that labile potassium concentrations may be reduced to between 99.3 and 53 per cent of the initial, depending on the diffusion characteristics of the soil and nutrient demand by the root. Of the total potassium absorbed by a root in 4 days, the proportion which is supplied from within the root hair cylinder is small (0.8 to 6.3 per cent) indicating that diffusion to the root from the soil outside the root hair cylinder is of paramount importance. When root demand is high, diffusion appears to limit uptake to between 71 and 59 per cent of that which roots of comparable physiology would be expected to absorb from stirred solution of the same concentration. Nevertheless, the presence of root hairs is calculated to have enhanced uptake by up to 77 per cent compared with roots without hairs because they virtually increase the root diameter. Diffusion does not appear to be a limiting factor when root demand is low and hairs can then add little to the efficiency of the root system in potassium absorption.     

The supply of nutrient ions by diffusion to plant roots in soil

Summary A single-root technique is used to measure the rate of supply of potassium by diffusion to 1-cm portions of cylindrical roots of onion and leek plants growing in soils containing different levels of exchangeable potassium. The relation between uptake and characteristics of the plant and soil is interpreted on the basis of a diffusion supply model. Uptake is accounted for in terms of the geometry of the absorbing root surface, the physiologically controlled absorbing power of the root, and the diffusion through the soil. The different uptakes of potassium by roots of comparable absorbing power from different soils can be predicted with some success from calculations using the root dimensions and either diffusion coefficients of potassium in soil, derived from flux to a cation exchange resin paper, or the form of the potassium scorption isotherm relating the concentration of labile ions to those in the soil solution. It is calculated that diffusion through the soil has reduced potassium uptake by the roots to between 87 and 39 per cent of that expected for roots of the same absorbing power in a stirred culture solution at the same initial soil solution concentration.     

Relationship of ion absorption to growth rate in taiga trees

Summary Rates of nutrient absorption were measured on excised roots of taiga tree seedlings grown in the laboratory. Phosphate and to a lesser extent ammonium (relatively immobile ions in the soil) were absorbed most rapidly by poplar and aspen, two species with rapid growth rates and most slowly by alder and/or black spruce, species with slow growth rates. In contrast, potassium (which is more mobile in soil) was absorbed most rapidly by slowly growing species. All species had low rates of nitrate and chloride absorption. Absorption rate of each ion was most temperature sensitive in those species that typically occupy the warmest soils (i.e. poplar and aspen). We suggest that in infertile soils a high capacity for uptake is an important component of root competition only in the case of mobile ions (e.g. potassium, nitrate), because only for these ions do diffusion shells of adjacent roots overlap; in contrast plants compete for immobile ions (e.g. phosphate) only by increasing absorptive surface via root growth or mycorrhizal association.     

Diffusion of phosphate to plant roots in soil

Summary Improved resolution in autoradiography, achieved by the use of the low energy isotope, P³³, as tracer for soil phosphorus, enables the exchangeable phosphorus in a soil block to be measured quantitatively. A technique is described for the autoradiography of the P-depletion zone around the roots growing in soil, from which the P gradients are measured by microdensitometry.The amounts of P taken up by rape (Brassica napus) on a P-treated Begbroke Sandy Loam compared well with that removed from the soil as measured from the autoradiograph of the depletion zone. The P gradient around the roots suggests intense root hair activity; but the zone of depletion extended well beyound the tips of root hairs. The experimentally observed gradient is much closer to the one predicted from diffusion theory considering uniform depletion from within the equivalent root hair cylinder, than to the one obtained assuming the root hairs are inactive.A rapid depletion of up to about 60 per cent of the exchangeable P was observed within the root hair cylinder during the initial 3 days of absorption. The corresponding concentration of P in solution within the cylinder determined from a desorption isotherm, is hence brought down to a low level very rapidly, and is held at or near this level at later periods. The amounts transferred into the root hair cylinder from outside as calculated from a diffusion model were lower than the experimental values. It is suggested that the discrepancy may lie in the calculation of the effective diffusion coefficients for P in the soil from a P-desorption isotherm, owing to difficulties involved in simulating the root environment in the desorption isotherm experimentSoil Science Laboratory, Department of Agricultural Science, University of Oxford     

Potassium uptake by plants,as characterized by root density,species and K/Rb ratio

Summary A small fraction of the plant K requirement is attained by root interception. The bulk of K has to be transported to the growing roots by mass-flow and diffusion in which diffusion mechanism plays the major role. Studies were undertaken to evaluate soil and plant parameters that might have influence on K supply mechanisms in soil and on plant uptake of K. Increasing wheat plant density led to competition for K absorption and resulted in lower K uptake by plant. In high plant density treatment, about 60% of the K requirement was met by diffusion process whereas in low plant density treatment mass-flow contributed most of the K demand. Solution diffusion and mass-flow were the major mechanisms of K supply to wheat roots. The mechanism of K supply to wheat root was compared with corn and onion. The major mechanism of K supply to corn and onion roots was exchange and solution diffusion. The mechanism of K supply to different crop species is attributable to differences in the K requirements, water flux rates and to the differences in root parameters.     

Proteoid root morphology and function inLupinus albus

Summary Current theories of phosphorus uptake by plants imply that they can augment diffusion to their root axes by the development of abundant root hairs or mycorrhizas. Some phosphorus efficient plants have root morphology with multi-branched roots and localised regions of densely packed root hairs, which we suggest is better suited to the retention of substances exuded by the roots than uptake of substances moving to the root by diffusion. Evidence of substantial exudation by the proteoid roots ofLupinus albus is presented.     

Root uptake and distribution of radiocaesium from contaminated soils and the enhancement of Cs adsorption in the rhizosphere

Uptake by roots from contaminated soil is one of the key steps in the entry of radiocaesium into the food chain. We have measured the uptake by roots of radiocaesium and its transfer to shoots of a heathland grass, sheep fescue (Festuca ovina L.) from two contrasting agricultural soils, a sandy podzol and a clayey calcareous soil. A culture device which keeps the roots separate from the soil was used thus allowing rhizosphere soil to be obtained easily and enhancing the effect of root action. Biomass production and ¹³⁷Cs in shoots and roots were recorded. Cs adsorption was studied on both the initial, nonrhizosphere soil and on rhizosphere soil in dilute soil suspension. Cs desorption was measured by resuspending subsamples of contaminated soil in solutions containing various concentrations of stable Cs. The proportion of Cs fixed, i.e. not readily desorbable, was calculated by comparison of the adsorption and desorption isotherms. Uptake by roots varied considerably between soils and did not appear to be diffusion limited. Root-to-shoot transfer did not differ for the two soils studied. Root action considerably enhanced Cs adsorption on the soils, particularly the in sandy podzol with a low Cs affinity. The value of K_d was increased by up to an order of magnitude. A large proportion of adsorbed Cs was found to be fixed, the K_d was up to seven times greater on desorption than adsorption, indicating that up to 80% of adsorbed Cs was not readily exchangeable. Root action had little effect on the fixed fraction. This revised version was published online in June 2006 with corrections to the Cover Date.     

Direct evidence on participation of root hairs in phosphorus (32P) uptake from soil

Root hairs substantially extend root surface for ion uptake. Although many reports suggest a relationship between root hairs and phosphorus (P) uptake of plants, the role of root hairs in phosphorus uptake from soils is still debated. We measured uptake of phosphorus from soil directly via root hairs. Root hairs only were allowed to penetrate through a tightly stretched nylon screen (53 µm) glued to the bottom of a PVC tube. The penetrating root hairs grew for 2 and 4 days in soil labelled with radioisotope phosphorus (P) tracer ³²P (185 kBq g^-1 dry soil) filled in another PVC tube. Transparent plastic rings of thickness ranging from 0.25 mm to 2.0 mm were inserted between the two PVC tubes. This provided slit width for microscopic observations in situ, which confirmed that only root hairs were growing into the ³²P labelled soil. In some cases no rings were inserted (slit width = 0) where both root hairs and root surface were in contact with the labelled soil (total ³²P uptake). The uptake of³² P from soil via the root hairs only was quantified by measuring activity of ³²P in the plant shoot (³²P uptake only via root hairs).The results showed that when 70 percent of the root hairs grew into the labelled soil, they contributed to 63 percent of the total P uptake. With decreasing number of root hairs growing into the ³²P labelled soil, the quantity of ³²P in the plant shoot decreased. In this study, P uptake via root hairs was measured in a soil-based system, where root hairs were the only pathway of ³²P from soil to the plant shoot. Therefore, this study provides a strong evidence on the substantial participation of root hairs in uptake of phosphorus from soil.     

Uptake of solutes by multiple root systems from soil

Summary The theoretical treatment of diffusion of solutes to a number of parallel, competing roots is difficult, but an electrical analog has been constructed which allows solute uptake by such a system to be simulated easily and rapidly. The construction, theory and operation of the analog are described. Differences in diffusion coefficients, dimensions, root size and uptake properties can all be dealt with. Approximate methods are available for simulating mass flow with diffusion, slow release of nutrients in the soil, the presence of root hairs, and incomplete root-soil contact.     

A root hairless barley mutant for elucidating genetic of root hairs and phosphorus uptake

This paper reports a new barley mutant missing root hairs. The mutant was spontaneously discovered among the population of wild type (Pallas, a spring barley cultivar), producing normal, 0.8 mm long root hairs. We have called the mutant bald root barley (brb). Root anatomical studies confirmed the lack of root hairs on mutant roots. Amplified Fragment Length Polymorphism (AFLP) analyses of the genomes of the mutant and Pallas supported that the brb mutant has its genetic background in Pallas. The segregation ratio of selfed F₂ plants, resulting from mutant and Pallas outcross, was 1:3 (–root hairs:+root hairs), suggesting a monogenic recessive mode of inheritance.In rhizosphere studies, Pallas absorbed nearly two times more phosphorus (P) than the mutant. Most of available inorganic P in the root hair zone (0.8 mm) of Pallas was depleted, as indicated by the uniform P depletion profile near its roots. The acid phosphatase (Apase) activity near the roots of Pallas was higher and Pallas mobilised more organic P in the rhizosphere than the mutant. The higher Apase activity near Pallas roots also suggests a link between root hair formation and rhizosphere Apase activity. Hence, root hairs are important for increasing plant P uptake of inorganic as well as mobilisation of organic P in soils.Laboratory, pot and field studies showed that barley cultivars with longer root hairs (1.10 mm), extracted more P from rhizosphere soil, absorbed more P in low-P field (Olsen P=14 mg P kg^–1 soil), and produced more shoot biomass than shorter root hair cultivars (0.63 mm). Especially in low-P soil, the differences in root hair length and P uptake among the cultivars were significantly larger. Based on the results, the perspectives of genetic analysis of root hairs and their importance in P uptake and field performance of cereals are discussed.     

Variation in phosphorus uptake efficiency by genotypes of cowpea (Vigna unguiculata) due to differences in root and root hair length and induced rhizosphere processes

The lengths of roots and root hairs and the extent of root-induced processes affect phosphorus (P) uptake efficiency by plants. To assess the influence of variation in the lengths of roots and root hairs and rhizosphere processes on the efficiency of soil phosphorus (P) uptake, a pot experiment with a low-P soil and eight selected genotypes of cowpea (Vigna unguiculata (L) WALP) was conducted. Root length, root diameter and root hair length were measured to estimate the soil volume exploited by roots and root hairs. The total soil P was considered as a pool of Olsen-P, extractable with 0.5 M NaHCO₃ at pH 8.5, and a pool of non-Olsen-P. Model calculations were made to estimate P uptake originated from Olsen-P in the root hair zone and the Olsen-P moving by diffusion into the root hair cylinder and non-Olsen-P uptake. The mean uptake rate of P and the mean rate of non-Olsen-P depletion were also estimated. The genotypes differed significantly in lengths of roots and root hairs, and in P uptake, P uptake rates and growth. From 6 to 85% of total P uptake in the soil volume exploited by roots and root hairs was absorbed from the pool of non-Olsen-P. This indicates a considerable activity of root-induced rhizosphere processes. Hence the large differences show that traits for more P uptake-efficient plants exist in the tested cowpea genotypes. This opens the possibility to breed for more P uptake-efficient varieties as a way to bring more sparingly soluble soil P into cycling in crop production and obtain capitalisation of soil P reserves.     

Direct evidence showing the effect of root surface iron plaque on arsenite and arsenate uptake into rice (Oryza sativa) roots

The present study aimed to investigate the effects of root surface iron plaque on the uptake kinetics of arsenite and arsenate by excised roots of rice (Oryza sativa) seedlings. The results demonstrated that the presence of iron plaque enhanced arsenite and decreased arsenate uptake. Arsenite and arsenate uptake kinetics were adequately fitted by the Michaelis-Menten function in the absence of plaque, but produced poor fits to this function in the presence of plaque. Phosphate in the uptake solution did not have a significant effect on arsenite uptake irrespective of the presence of iron plaque; however phosphate had a significant effect on arsenate uptake. Without iron plaque, phosphate inhibited arsenate uptake. The presence of iron plaque diminished the effect of phosphate on arsenate uptake, possibly through a combined effect of arsenate desorption from iron plaque.     

The phosphate uptake mechanism

The slow rate of diffusion of phosphate in soil results in a zone of depletion of phosphate ions in solution around the roots of plants in low phosphate soils. Transfer of phosphate to the site of uptake into the root symplasm limits phosphate uptake in such soils. This transfer involves movement across the depletion zone and through the root apoplasm. The apoplasm is made up of the cell walls of epidermal and cortical cells, together with the associated intercellular spaces. Although the pores in the open latticework of these cell walls permit movement of nutrients around cells, they increase the path length across which phosphate ions have to diffuse. The structural components and net negative charges of the cell walls also influence the effective concentrations of phosphate in the apoplasm. This concentration may be further modified by excreted organic compounds around cell walls and the presence of micro-organisms that use such compounds as carbon sources. A membrane on the inner surface of the cell wall, the plasmalemma, separates the apoplasm from the symplasm. Uptake of nutrients into the root symplasm occurs through transporter proteins embedded in this membrane. Understanding of the mechanisms by which phosphate is transported across the plasmalemma into the plant symplasm has advanced considerably over the past 4 years due to the application of molecular techniques. Genes encoding the transporters involved in this process have been isolated from a number of plant species. These transporters belong to a family of membrane proteins characterized by having 12 membrane-spanning domains arranged in a '6+6' configuration. H₂PO₄ ^– ions, together with protons, are transported through this protein. This transport process is driven by the potential across the membrane maintained by the action of a H⁺-ATPase, the `proton pump', that extrudes protons to the outer surface of the membrane. The expression of genes encoding high-affinity root phosphate transporters is regulated by the phosphorus (P) status of the plant. The transduction pathway involved in this regulation is not known at present. It is a systemic response rather than a localized response, however, the overall phosphate status of the plant being the controlling factor. Under phosphate stress, the expression of genes encoding these phosphate transporters is up-regulated. This results in a greater number of transporter proteins in the plasmalemma and enhanced phosphate uptake rates, if phosphate is available at the membrane surface. Uptake occurs around the root tip, into epidermal cells with their associated root hairs and into cells in the outer layers of the root cortex. Further back along the root axis, phosphate can also be taken up by transfer from mycorrhizal fungi to root cortical cells.Strategies for increasing nutrient uptake by overexpressing genes encoding high-affinity phosphate transporters are likely to be mainly applicable to situations where a reasonable phosphate concentration can be maintained at the outer surface of the plasmalemma. Maintaining such a concentration is a major problem in the phosphate deficient soils of the semi-arid tropics (SAT), so emphasis in these soils is on strategies to improve the movement of phosphate to the surface of the plasmalemma. There may be scope, however, for manipulating the expression of genes involved in the internal mobilisation of phosphate within the plant, thereby improving phosphate utilisation.     

A Relationship between Phosphate Uptake and Membrane Electrical Potential in Excised Roots of Helianthus annuus

Phosphate uptake by excised roots of sunflower (Helianthus annuus)was determined by the disappearance of phosphate from the externalsolution and by the accumulation of phosphate labelled with³²P. Over a 24 h period it was observed that net phosphate uptakedeclined to zero whilst uptake of ³²P continued unabated. Theelectrical PD of the cortical cell membranes declined in parallelwith net phosphate uptake and it was found that both could berestored by creating a pH gradient across the plasmalemma. Itwas concluded that net phosphate uptake was responsible fora component of the membrane PD of the root cortical cells. Key words: Roots, Phosphate, Membranes     

Phosphate absorption and root respiration of different plant growth forms from northern Alaska

Rate of phosphate absorption measured on field-collected, excised roots of eight species from a tundra site and a taiga forest was more strongly correlated with growth form traits than with type of community. In the taiga forest phosphate absorption rate ranged tenfold among species and was higher in graminoids than in shrubs. In both sites deciduous shrubs tended to have higher rates than evergreens. Phosphate absorption rate was not strongly reduced at low temperature in any species examined and there was a temperature optimum of about 20°C in Eriophorum vaginatum . Phosphate absorption rates were higher in lateral than primary roots, although both root types had substantial respiration rates. At one tundra site, calculations suggest that roots of E. vaginatum require 30–40 d to absorb a quantity of phosphorus equivalent to that required to synthesize the root and another 5–6 d to provide the net phosphorus requirements for the remainder of the plant.     

Role of root hairs in phosphorus depletion from a macrostructured soil

One rape (Brassica napus cv. Wesroona) plant and four cotton (Gossypium hirsutum cv. Sicot 3) plants were grown in plastic cells containing soil labelled with 407 kBq of³³P g⁻¹ soil. After 5–8 days of growth, the³³P depletion zones of all plants were autoradiographed and³³P uptake by plants was measured. The autoradiographs were scanned with a microdensitometer and the optical densities at several places within the³³P depletion zones of roots were obtained. The volume of soil explored by root hairs was estimated from measurements of root diameters and lengths of roots and root hairs. About half of the total³³P depleted by cotion roots came from outside the root hair cylinder whereas most of³³P taken up by rape was from within the root hair cylinder. Plants grown in a macrostructured soil may have roots growing in voids, within aggregates or on the surfaces of aggregates. The results of this study demonstrate that root hairs have a strong influence on the accessibility of phosphorus to roots in such a soil, and thus on the phosphorus nutrition of plants.     

A dual porosity model of nutrient uptake by root hairs

? The importance of root hairs in the uptake of sparingly soluble nutrients is understood qualitatively, but not quantitatively, and this limits efforts to breed plants tolerant of nutrient-deficient soils. ? Here, we develop a mathematical model of nutrient uptake by root hairs allowing for hair geometry and the details of nutrient transport through soil, including diffusion within and between soil particles. We give illustrative results for phosphate uptake. ? Compared with conventional 'single porosity' models, this 'dual porosity' model predicts greater root uptake because more nutrient is available by slow release from within soil particles. Also the effect of soil moisture is less important with the dual porosity model because the effective volume available for diffusion in the soil is larger, and the predicted effects of hair length and density are different. ? Consistent with experimental observations, with the dual porosity model, increases in hair length give greater increases in uptake than increases in hair density per unit main root length. The effect of hair density is less in dry soil because the minimum concentration in solution for net influx is reached more rapidly. The effect of hair length is much less sensitive to soil moisture.     

Transcriptional regulation and functional properties of <Emphasis Type="Italic">Arabidopsis</Emphasis> Pht1;4, a high affinity transporter contributing greatly to phosphate uptake in phosphate deprived plants

Phosphate mobilization into the plant is a complex process requiring numerous transporters for absorption and translocation of this major nutrient. In the genome of Arabidopsis thaliana, nine closely related high affinity phosphate transporters have been identified but their specific roles remain unclear. Here we report the molecular, histological and physiological characterization of Arabidopsis pht1;4 high affinity phosphate transporter mutants. Using GUS-gene trap and in situ hybridization, Pht1;4 was found mainly expressed in inorganic phosphate (Pi) limiting medium in roots, primarily in the epidermis, the cortex and the root cap. In addition to this, expression was also observed at the lateral root branch points on the primary root and in the stele of lateral roots, suggesting a role of Pht1;4 in phosphate absorption and translocation from the growth medium to the different parts of the plant. Pi-starved pht1;4 plantlets exhibited a strong reduction of phosphate uptake capacity (40). This phenotype appears only related to the pht1;4 mutation as there were no obvious changes in the expression of other Pht1 family members in the mutants background. However, after 10 days of growth on phosphate deficient or sufficient medium, the Pi content in the mutants was not significantly different from that of the corresponding wild type controls. Furthermore, the mutants did not display any obvious growth defects or visible phenotypes when grown on a low phosphate containing medium. The work described here offers a first step in the complex genetic dissection of the phosphate transport system in planta.     

STUDIES OF THE BIOLOGY OF SCLEROTIUM CEPIVORUM BERK.

Sclerotia from 6-week-old pure cultures of Sclerotium cepivorum germinated immediately in soil only after abrasion of their rinds, but after burial in soil for a month or more, unabraded sclerotia became capable of germination.
Marked stimulation of germination occurred in the presence of host plants (onion, leek and shallot). Little or no germination occurred in soil alone or in the presence of non-host plants (barley, cabbage, carrot and white clover). Sclerotial germination was observed in a number of soils of widely differing pH and over a wide range of soil water content. Germination of sclerotia on uninjured onion roots was greatest at the tip region. On artificially injured roots sclerotial germination was enhanced but the effect of position was lost.
Sclerotial germination was independent of contact between roots and sclerotia. It was induced by root extracts of all Allium spp. tested, but of no other plants. Boiling or autoclaving root extracts did not destroy the active principle and it is concluded that under field conditions sclerotia are induced to germinate by a thermostable chemical substance from Allium roots.
The process of germination of sclerotia is described.     

Phosphorus nutrition of barley, buckwheat and rape seedlings. II. Influx and efflux of phosphorous by intact roots of different P status

Seedlings of barley (Hordeum vulgare L. cvs Salka and Zita), buckwheat (Fagopyrum esculentum Moench) and rape (Brassica napus L. ssp. napus cv. Line) were raised at 8 or 10 different extenral P concentrations in the range 0–2000 μM. Apart from P, the nutrient solutions were complete. Phosphate influx in roots of different P status was determined by use of a nutrient solution containing 0.1 mM³²P-labelled phosphate. A double labelling technique was used for simultaneous determination of influx (³³P) and efflux (³²P) of phosphorus by roots of barley and rape with three selected P levels. Flux determinations were also done in presence of a metabolic uncoupler (2,4-dinitrophenol) and a protein synthesis inhibitor (cycloheximide). Influx of phosphate was maximal at a certin optimal P level of the roots and decreased at both lower and higher P levels. Maximum phosphate influex [μmol (g root)-^?1 h^?1] were: rape 4,4, buckwheat 2.2, barley cv. Salka 1.6, barley cv. Zita 1.5. Both Hill plots and plots of the untransformed decreasing phosphate influx vs root P concentrations above the optimal were linear and had high correlation coefficients. The Hill coefficient varied between -3.1 and -4.2. The decrease of phosphate influx from the maximum to the lowest value at the highest P concentration of the root was 60–70%. Hence, phosphate influex appeared to be regulated through negative feedback by the internal level of phosphorous in the roots. The regulation mechanism seems bascially similar for the three species and may be of an allosteric type. P efflux from roots of low and optimal (with regard to P influx) P status was 15–20% of the simultaneous P influx. Contary to P influx, P efflux increased at high P status and almost eliminated (barley) or halved (rape) net P uptake. 2,4-Dinitrophenol reduced both P influx and P efflux by low P roots and gave linearly increasing P efflux with increasing root P status. This indicates that P efflux partly occurred by counter transport and ion exchange at the uptake sites, partly by passive P efflux along an electrochemical potential gradient. Phosphate influx was not affected by inhibition of barley root growth with cycloheximide, but P efflux increased considerably.