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.     

Roots regulate ion transport in the rhizosphere to counteract reduced mobility in dry soil

Diffusion of ions in the soil depends on soil moisture content. In a dry soil, transport of nutrients towards the root and the concomitant uptake could be reduced. However, pot and field experiments showed that this is not always the case. The objective of this paper was to investigate possible mechanisms of plants to counteract reduced nutrient supply due to water shortage. A split root system was used to investigate P and K inflow of oat and sugar beet at different soil moisture contents (Θ) without water shortage for the plant. The measured average P and K inflows were compared to model calculations considering diffusion, mass-flow, sorption and uptake processes. In the calculations, soil dryness impeded diffusion and decreased nutrient inflow as expected. Measured K inflow was decreased in a similar way indicating that Θ influences K diffusion. In contrast to this, measured P inflow was not influenced by Θ and under-estimated by the model. Low and high molecular exudates were collected at different water supply levels showing that exudation rate of both compounds was increased at water shortage. Especially the high molecular exudates (i.e. mainly mucilage) from water-stressed plants increased P concentration in soil solution under dry conditions in an incubation experiment. Calculated inflow considering this increased P concentration agreed well with measured P inflow indicating that exudation of mucilage could be a mechanism to overcome nutrient transport problems due to soil dryness.     

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

Summary Measurements were made of the diffusion of P³²-labelled phosphate to single roots of onion, leek and rye-grass growing in an Upper Greensand sandy loam (UGS) and a Coral Rag Clay (CRC) to which different amounts of phosphate had been added. Concentration-dependent diffusion coefficients for phosphate ions in the soils were calculated from phosphate desorption isotherms in calcium chloride. The experimental uptake by roots of known dimensions was compared with supply expected by diffusion to a cylindrical model root of the same dimensions. Allowance was made for absorption by the root hairs on rye-grass roots. Phosphate absorption by a cm length of intact root was found to continue for at least 16 days for onion, 10 days for leek and 5 days for rye-grass. Over a wide range of conditions (phosphate concentrations, soils, plant species), experimental uptake was close to the maximum calculated to be possible for the diffusion model except on one soil at a high level of phosphate. Although the concentration of phosphate in the soil solution at the root boundary appeared to be reduced to a small fraction of the initial concentration, because of the extreme non-linear form of the desorption isotherm less than 1/2 of the P³² exchangeable pool of P was considered to contribute to diffusion. Phosphate uptake by rye grass could only be accounted for if the root hairs were active. Although only a small fraction of the uptake is derived from inside the root hair cylinder, this increases the efficiency of the central root 2.3 fold by providing a zone close to the central root through which phosphate moves very readily.     

Uptake of solutes by multiple root systems from soil

Summary The analog described in Part I is used to investigate quantitatively the the effects of pattern and density on the uptake and uptake rate of nutrients which move to plant roots by diffusion. The uptake by two roots is considered first, to illustrate the competitive effect. The results for multiple root systems are given for a variety of different soil and plant parameters at different times and demonstrate the importance of pattern and density in the uptake of different plant nutrients in both competitive and non competitive situations. Pattern can decrease the uptake by root systems by at least 75 per cent, depending on the value of the diffusion coefficient, time, and root density. Graphs of two indices of dispersion against uptake are given so that the effect of any pattern can be estimated. A procedure is outlined which enables the uptake after any time by a developing root system to be predicted and compared with a theoretical maximum. If the uptake is known, then the graphs show whether soil or plant parameters are limiting uptake.     

Uptake of phosphorus and potassium in relation to root growth and root density

Summary This paper provides some quantitative data on the relationship between the rate of uptake of phosphorus and potassium from soil and the amount of root, root density and rate of root growth. Three experiments were conducted with winter wheat, all grown in the same soil. Root growth and density were manipulated in three ways: (1) by root pruning; (2) by a split-root technique; (3) by growing plants in different soil volumes. Root lengths as well as weights were determined.Potassium uptake per unit amount of root was generally lower the higher the root density, suggesting that roots were competing with each other for potassium even at the lowest density. In contrast, phosphorus uptake showed a good correlation with root growth irrespective of root density or plant age. Phosphorus uptake during a period was more closely and consistently correlated with root growth during that period than with the total amount of root on the plant. The results can be explained in terms of ion supply to the root surface, taking into account the diffusion coefficients of the ions and the approximate distances between neighbouring roots.Now Mrs. Watkins; address 39 Leach Heath Lane, Rubery, Birmingham.Now Mrs. Watkins; address 39 Leach Heath Lane, Rubery, Birmingham.     

Modelling potassium uptake by wheat (Triticum aestivum) crops

Spring wheat was grown in the field under deficient and sufficient levels of soil K and with high and low supplies of fertiliser nitrogen. Measurements were made of K uptake, soil nutrient supply parameters, root growth and, in solution culture, root influx parameters. Mechanistic models predicted uptake reasonably well under K-deficient conditions, but over-predicted uptake, by as much as 4 times, under K-sufficient conditions. The over-prediction was apparently due to poor characterisation of plant demand.     

Exploitation of K near roots of cotton,maize, upland rice,and soybean grown in an Ultisol

Native soil potassium (K) has received increased attention as a K source for plants to reduce fertiliser input. Our objective was to compare the ability of different crops to utilise native K. We also wanted to study the exploitation and transport pattern of soil K influenced by plant uptake. Cotton, maize, soybean, and upland rice were cultivated in rhizoboxes. The system permitted sampling of 1-mm-thin soil layers at increasing distances from the plant roots. Both exchangeable and nonexchangeable K was determined and compared with plant uptake of K. The upland rice was superior in K uptake, and took up some nonexchangeable K. Soybean and cotton grew poorly, and K was accumulated in the root zone due to excess supply by mass flow. The importance of mass flow over diffusion of K was verified by calculations and is contrary to accepted principles of K transport in soil. The reasons were high transpiration and restricted root growth. This indicates that mass flow of K in some situations is more important than generally assumed. Mass flow also caused the accumulation of Ca and Na in the root zone, especially that of rice. Accumulation of K in the root zone of rice did not cause K fixation, possibly due to an unknown K-releasing mechanism of upland rice. This revised version was published online in July 2006 with corrections to the Cover Date.     

Uptake of solutes by multiple root systems from soil

Summary A procedure is put forward for calculating the plant uptake of solutes supplied by diffusion and mass flow to the randomly dispersed roots of a developing root system. The model was tested as follows: (a) for a constant root density, and both transport processes—against a more accurate numerical solution of the same system (b) for an increasing root density, and for supply by diffusion only—by electrical simulation using the analog described in Part I. In both cases, results obtained by the two types of calculation were in close agreement. A less accurate method which includes both supply mechanisms and does not require a computer is presented, and compared with an electrical simulation when there is no mass flow. Agreement is within 20 per cent. The model should be useful for predicting plant nutrient uptake from soil, and may be of special interest to modellers of the whole plant system. re]19720905 Soil Science Laboratory, Department of Agricultural Science ,University of Oxford Present address: Department of Plant Sciences,University of Leeds     

Potassium and phosphorus uptake by corn genotypes grown in the field as influenced by root characteristics

Summary Root parameters of three corn (Zea mays L.) genotypes influencing P and K uptake were investigated in solution culture and field experiments. The data for these parameters were used to simulate P and K uptake by plants grown in the field using the Claassen-Barber model⁵. Root characteristics for ion influx, maximum rate of influx,Imax; Michaelis-Menten constant,Km; and minimum concentration of solution below which no further net influx occurs,Cmin were determined in solution culture. These kinetic parameters varied 2 to 3 fold among genotypes. Variations among genotypes were different for K than for P.Three corn genotypes were grown in the field and harvested 47, 54 and 68 days after emergence. Yield and root surface per plant increased about 3 fold during this time. At 47 days, 2/3 of the total root surface was in the top soil whereas 3 weeks later, it was less than 50%. Genotypes differed in distribution of roots between the topsoil and subsoil as well as in root surface per unit of shoot.K uptake predicted by the Claassen-Barber model was 2 to 3 times the observed. The overprediction could be related to high root density (length of root per unit soil volume) which indicated that competition between roots occurred that was not considered in the simulation model. The predicted P uptake (y) was correlated (r=0.91) to observed uptake (x) byy=0.98+0.67x, indicating underprediction of P uptake. The presence of root hairs may have been the cause of the underprediction. The calculated contribution of the subsoil to the observed uptake was 10% for K and 1% in the case of P. It was concluded that the plant parameters used to simulate nutrient uptake were rated accurately when allowance was made for root competition and presence of root hairs.Journal Paper No. 7608. Purdue University, Agric. Exp. Station, West Lafayette, IN 47907. Contribution from the Department of Agronomy. This research was supported in part by the Tennessee Valley Authority and the Deutsche Forschungsgemeinschaft.     

Modelling diverse root density dynamics and deep nitrogen uptake—A simple approach

We present a 2-D model for simulation of root density and plant nitrogen (N) uptake for crops grown in agricultural systems, based on a modification of the root density equation originally proposed by Gerwitz and Page in J Appl Ecol 11:773–781, (1974). A root system form parameter was introduced to describe the distribution of root length vertically and horizontally in the soil profile. The form parameter can vary from 0 where root density is evenly distributed through the soil profile, to 8 where practically all roots are found near the surface. The root model has other components describing root features, such as specific root length and plant N uptake kinetics. The same approach is used to distribute root length horizontally, allowing simulation of root growth and plant N uptake in row crops. The rooting depth penetration rate and depth distribution of root density were found to be the most important parameters controlling crop N uptake from deeper soil layers. The validity of the root distribution model was tested with field data for white cabbage, red beet, and leek. The model was able to simulate very different root distributions, but it was not able to simulate increasing root density with depth as seen in the experimental results for white cabbage. The model was able to simulate N depletion in different soil layers in two field studies. One included vegetable crops with very different rooting depths and the other compared effects of spring wheat and winter wheat. In both experiments variation in spring soil N availability and depth distribution was varied by the use of cover crops. This shows the model sensitivity to the form parameter value and the ability of the model to reproduce N depletion in soil layers. This work shows that the relatively simple root model developed, driven by degree days and simulated crop growth, can be used to simulate crop soil N uptake and depletion appropriately in low N input crop production systems, with a requirement of few measured parameters.     

The role of soil hydraulic conductivity on the spatial and temporal variation of root water uptake in drip-irrigated corn

A multiplexed TDR system and a heat-pulse system for stem sap flow measurements were used to determine the spatial and temporal pattern of root water uptake in field-grown corn. The TDR probes, 0.15 and 0.30 m in length, were buried vertically in the soil profile to a depth of 0.95 m below the soil surface and heat-pulse sensors were installed on the plant base. Nocturnal readings from TDR probes were used successfully to differentiate the two components of moisture change: root uptake and net drainage. The instantaneous rate of water extraction by the plant measured by the heat-pulse system agreed well with the integrated rate of root water uptake measured frequently (at half-hour or hourly intervals) by the TDR probes. This agreement enabled further exploration into the cause of the evolution of the spatial and temporal patterns of root water uptake during a drying cycle. The results indicated that right after irrigation in the well-watered soil profile, it is the spatial distribution of the roots that mainly determines the typical pattern of root extraction, in addition to the fact that the roots near the plant base are more effective than those farther away. The higher density and effectiveness of the roots near the plant base dry the soil rapidly so that soil hydraulic conductivity soon becomes a limiting factor for water uptake. Further analysis revealed that a decrease in root uptake occurs near the plant base under a given atmospheric demand when the relative bulk soil hydraulic conductivity decreases to 0.002K _r. This suggests that low conductivity (high resistance) in the soil near the plant base is the initial cause for downward and lateral shifting of the root uptake pattern. Note that this critical value of hydraulic conductivity is not universal since it depends on the soil type and atmospheric water demand during the period under observation. Therefore, prior to the application of moisture content or suction head as measures of water availability or to control irrigation scheduling, it is suggested that these parameters be calibrated by the soil K() or K() curves, respectively, for the expected atmospheric water demand for the specific crop and growing period.     

Inward-Rectifying K+ Channels in Root Hairs of Wheat (A Mechanism for Aluminum-Sensitive Low-Affinity K+ Uptake and Membrane Potential Control)

K+ is the most abundant cation in cells of higher plants, and it plays vital roles in plant growth and development. Extensive studies on the kinetics of K+ uptake in roots have shown that K+ uptake is mediated by at least two transport mechanisms, one with a high and one with a low affinity for K+. However, the precise molecular mechanisms of K+ uptake from soils into root epidermal cells remain unknown. In the present study we have pursued the biophysical identification and characterization of mechanisms of K+ uptake into single root hairs of wheat (Triticum aestivum L.), since root hairs constitute an important site of nutrient uptake from the soil. These patch-clamp studies showed activation of a large inward current carried by K+ ions into root hairs at membrane potentials more negative than -75 mV. This K+ influx current was mediated by hyperpolarization-activated K+-selective ion channels, with a selectivity sequence for monovalent cations of K+ > Rb+ [almost equal to] NH4+ >> Na+ [almost equal to] Li+ > Cs+. Kinetic analysis of K+ channel currents yielded an apparent K+ equilibrium dissociation constant (Km) of [almost equal to]8.8 mM, which closely correlates to the major component of low-affinity K+ uptake. These channels did not inactivate during prolonged stimulation and would thus enable long-term K+ uptake driven by the plasma membrane proton-extruding pump. Aluminum, which is known to inhibit cation uptake at the root epidermis, blocked these inward-rectifying K+ channels with half-maximal current inhibition at [almost equal to]8 [mu]M free Al3+. Aluminum block of K+ channels at these Al3+ concentrations correlates closely to Al3+ phytotoxicity. It is concluded that inward-rectifying K+ channels in root hairs can function as both a physiologically important mechanism for low-affinity K+ uptake and as regulators of membrane potential. The identification of this mechanism is a major step toward a detailed molecular characterization of the multiple components involved in K+ uptake, transport, and membrane potential control in root epidermal cells.     

Simulation of nutrient uptake by a growing root system considering increasing root density and inter-root competition

A simulation model is presented which describes uptake of a growth limiting nutrient from soil by a growing root system. The root surface is supposed to behave like a zero-sink. Uptake of the nutrient is therefore determined by the rate of nutrient supply to the root surface by mass flow and diffusion. Inter-root competition and time dependent root density are accounted for by assigning to each root a finite cylindrical soil volume that delivers nutrients. The radius of these cylinders declines with increasing root density. Experiments with rape plants grown on quartz sand were used to evaluate the model. Simulated nitrogen uptake agreed well with observed uptake under nitrogen limiting conditions. In case no nitrogen limitation occurred nitrogen uptake was overestimated by the model, probably because the roots did not behave like a zero-sink any more.     

The role of plant species and soil condition in the structural development of the rhizosphere

Roots naturally exert axial and radial pressures during growth, which alter the structural arrangement of soil at the root–soil interface. However, empirical models suggest soil densification, which can have negative impacts on water and nutrient uptake, occurs at the immediate root surface with decreasing distance from the root. Here, we spatially map structural gradients in the soil surrounding roots using non‐invasive imaging, to ascertain the role of root growth in early stage formation of soil structure. X‐ray computed tomography provided a means not only to visualize a root system in situ and in 3‐D but also to assess the precise root‐induced alterations to soil structure close to, and at selected distances away from the root–soil interface. We spatially quantified the changes in soil structure generated by three common but contrasting plant species (pea, tomato, and wheat) under different soil texture and compaction treatments. Across the three plant types, significant increases in porosity at the immediate root surface were found in both clay loam and loamy sand soils and not soil densification, the currently assumed norm. Densification of the soil was recorded, at some distance away from the root, dependent on soil texture and plant type. There was a significant soil texture × bulk density × plant species interaction for the root convex hull, a measure of the extent to which root systems explore the soil, which suggested pea and wheat grew better in the clay soil when at a high bulk density, compared with tomato, which preferred lower bulk density soils. These results, only revealed by high resolution non‐destructive imagery, show that although the root penetration mechanisms can lead to soil densification (which could have a negative impact on growth), the immediate root–soil interface is actually a zone of high porosity, which is very important for several key rhizosphere processes occurring at this scale including water and nutrient uptake and gaseous diffusion.     

Nutrient availability moderates transpiration in Ehrharta calycina

Transpiration-driven 'mass-flow' of soil-water can increase nutrient flow to the root surface. Here it was investigated whether transpiration could be partially regulated by nutrient status. Seeds of Ehrharta calycina from nine sites across a rainfall gradient were supplied with slow-release fertilizer dibbled into the sand surrounding the roots and directly available through interception, mass-flow and diffusion (dubbed 'interception'), or sequestered behind a 40-microm mesh and not directly accessible by the roots, but from which nutrients could move by diffusion or mass-flow (dubbed 'mass-flow'). Although mass-flow plants were significantly smaller than interception plants as a consequence of nutrient limitation, they transpired 60% faster, had 90% higher photosynthesis relative to transpiration (A/E), and 40% higher tissue P, Ca and Na concentrations than plants allowed to intercept nutrients directly. Tissue N and K concentrations were similar for interception and mass-flow plants. Transpiration was thus higher in the nutrient-constrained 'mass-flow' plants, increasing the transport of nutrients to the roots by mass-flow. Transpiration may have been regulated by N availability, resulting in similar tissue concentration between treatments. It is concluded that, although transpiration is a necessary consequence of photosynthetic CO(2) uptake in C(3) plants, plants can respond to nutrient limitation by varying transpiration-driven mass-flow of nutrients.     

Modelling zinc uptake by rice crop using a Barber-Cushman approach

The Barber-Cushman mechanistic nutrient uptake model which has been utilized extensively to describe and predict nutrient uptake by crop plants at different stages of crop growth was evaluated for its ability to predict the Zn uptake by rice seedlings. Uptake of the nutrient is, therefore, determined by the rate of nutrient supply to the root surface by mass flow and diffusion. Inter root competition and time dependent root density are accounted for by soil volume that delivers nutrients. The radii of these cylinders decline with increasing density. Since mass flow and diffusion each supply zinc to the root, the process can be described mathematically using the model of Barber-Cushman (1984). The 11 parameters of the model for the uptake by rice cultivars were measured by established experimental techniques. Zinc uptake at different growth stages predicted by the model was compared to measured zinc uptake by rice cultivars grown on sandy loam soil in a green house. Predicted zinc uptake was significantly correlated with observed uptake r ²=0.99^**. Sensitivity analysis was also used to investigate the impact of changes in soil nutrient supply, root morphological and root uptake kinetic parameters on simulated nutrient uptake. Overall results of sensitivity analysis indicate that the half distance between root axes, rate of root growth and water flux affect the uptake of zinc particularly at their higher values rather than at lower values and DaZn is the most sensitive parameter for zinc uptake at its lower values.     

Root proliferation,nitrate inflow and their carbon costs during nitrogen capture by competing plants in patchy soil

The responses of roots to nitrogen- and phosphorus-rich patches of soil include proliferation of laterals and stimulation of nutrient inflow (uptake rate per unit root length) within the patch. Nitrate uptake from an N-rich patch is thereby maximised and, perhaps, compensates for an uneven supply of nitrate to the whole root system. Paradoxically, the often weak correlation between root length density and N uptake found in experiments on single plants and crop monocultures suggests that root proliferation in patches has only a minor compensatory influence on N capture. This paradox was resolved when it was realised that localised root proliferation during inter-specific competition for nitrate can lead to a strong association between root length density and nitrate capture. Here, a simple model of inter-specific competition is used to estimate the stimulation in inflow required in one plant to match the N capture of a competitor that responds only by root proliferation, and to estimate associated carbon costs. The model predicts that nitrate inflow must increase proportionally more than root length density to achieve the same N capture. For example, the N capture possible with a 10% increase in root length density can be matched by increasing N inflow by anything from 20% to 20-fold, depending on the initial conditions: the faster the rate of change in root length density, the greater the required relative increase in inflow. In those terms, proliferation would seem the better option, but one that may be more costly in terms of its carbon requirement.     

The relative efficiency of zinc carriers on growth and zinc nutrition of corn

Summary A comparison of different zinc carriers showed that application of Zn-DTPA, Zn-EDTA, Zn-fulvate and ZnSO₄ significantly increased the dry matter yield and zinc uptake by corn over the control treatment where no zinc was applied. The chelates in particular enhanced to a greater extent the uptake of both native and applied sources than that observed with ZnSO₄ as the zinc carrier. Both the dry matter yield and zinc uptake by corn showed a positive and significant relationship with self-diffusion coefficient of zinc showing thereby that diffusion contributed mainly the supply of Zn from the ambient soil matrix to plant roots. The effectiveness of the chelates varied depending on their capacity to retain Zn in a soluble form in the soil solution.It is evident that zinc nutrition of plants in alkaline and calcareous soils can be more effectively regulated by both synthetic and natural chelates or organic manures which contain substantial amount of complexed zinc.Journal Paper No. 1 from the Department of Soil Science and Agric. Chemistry, Tirhut College of Agriculture, Dholi, Muzaffarpur, Bihar, India.     

Performance of seminal and nodal roots of wheat in stagnant solution: K+ and P uptake and effects of increasing O2 partial pressures around the shoot on nodal root elongation

Roots of intact wheat plants were grown for 7-12 d in stagnant nutrient solution, containing 0.1% agar, to mimic the lack of convection in waterlogged soil. Net K+ and P uptakes by seminal and nodal roots were measured separately using a split root system. For seminal roots in stagnant solution, net uptakes as a percentage of aerated roots were between 0% and 16% for P, while K+ ranged between 15% uptake and 54% loss. For the more waterlogging-tolerant nodal roots, net uptakes in stagnant nutrient solution, as a percentage of aerated roots, were 31-73% for P and 69-115% for K+. Elongation rates of nodal roots in stagnant nutrient were about 35-43% of those for roots in aerated solution. This partial inhibition occurred in these nodal roots despite their 15% porosity (v/v). Elevation of O2 partial pressures around the shoots to 40 kPa and then to 80 kPa substantially accelerated nodal root elongation in stagnant solution, demonstrating that most of the inhibition seen with ambient O2 around the shoots was associated with a restricted O2 supply to these nodal roots. Thus, in wheat nodal roots, with a partial pressure of 20 kPa O2 around the shoots, O2 diffusion from the shoots did not completely relieve the restrictions on elongation resulting from stagnancy in the nutrient solution. These results contrast with those in the literature for rice, in which roots function efficiently in stagnant solutions (0.1% agar). So, when wheat roots are aerenchymatous there are still restrictions to O2 diffusion in the gas space continuum between the atmosphere and the functional tissues of the roots. This poor acclimation must have been due to inefficiency of the aerenchymatous axes, which may include persistence of anoxic steles, and/or restricted O2 diffusion in other parts of the gas space continuum, in either the shoots and shoot-root junction or in the root tip.