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.     

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.     

The possibility of predicting solute uptake and plant growth response from independently measured soil and plant characteristics

Summary A model of the way the rate of growth of a plant may be affected by the level of supply of a nutrient is presented. Growth rate is linked to the nutrient level of the photosynthetic tissues, which is assumed to control changes in the net assimilation rate, the leaf area per unit shoot weight, the shoot: root ratio, the root surface area, and the distribution of nutrient between root and shoot. The uptake of nutrient depends on the concentration of nutrient at the root surface, the root surface area and its absorbing power. All these relationships may be determined in stirred solution culture. A method of applying this information to soil grown plants is suggested.Soil Science Laboratory, Department of Agricultural Science, University of Oxford     

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.     

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.     

Modelling the interactions between water and nutrient uptake and root growth

A model of three-dimensional root growth has been developed to simulate the interactions between root systems, water and nitrate in the rooting environment. This interactive behaviour was achieved by using an external-supply/internal-demand regulation system for the allocation of endogenous plant resources. Data from pot experiments on lupins heterogeneously supplied with nitrate were used to test and parameterise the model for future simulation work. The model reproduced the experimental results well (R ² = 0.98), simulating both the root proliferation and enhanced nitrate uptake responses of the lupins to differential nitrate supply. These results support the use of the supply/demand regulation system for modelling nitrate uptake by lupins. Further simulation work investigated the local uptake response of lupins when nitrate was supplied to a decreasing fraction of the root system. The model predicted that the nitrate uptake activity of lupin roots will increase as the fraction of root system with access to nitrate decreases, but is limited to an increase of around twice that of a uniformly supplied control. This work is the first example of a modelled root system responding plastically to external nutrient supply. This model will have a broad range of applications in the study of the interactions between root systems and their spatially and temporally heterogeneous environment.     

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.     

Effects of CO2, CH4 and N2 on growth and nutrition of rice seedlings

Summary Flooded soils, which accumulate gaseous products of anaerobic fermentation, are often associated with poor rice plant growth. In the present experiment the effects of CO₂, CH₄, N₂, and air on rice seedling growth and nutrition were evaluated. Nutrient culture techniques were used to avoid secondary soil effects normally experienced.Carbon dioxide gas in the root zone of rice reduced seedling growth significantly, whereas CH₄ and N₂ had no significant effect. Methane gave no stimulatory benefits, unlike results reported by some earlier workers. Of three major nutrient elements studied, P uptake was affected more than N or K. Phosphorus uptake was significantly reduced in leaves and sheaths by all three gases, but was significantly increased in roots. This suggests an immobilization mechanism affecting P in roots, and since CO₂, CH₄, and N₂ behaved similarly in contrast to air, a lack of oxygen in the root system is suspected as the causal mechanism rather than toxic effects of gases. Effects on N and K uptake were minimal and insignificant.Contribution from the Department of Agronomy and Range Science, University of California, Davis, California 95616.Contribution from the Department of Agronomy and Range Science, University of California, Davis, California 95616.     

An improved system of subjecting plants to water stress

An improved system of plant cultivation at stable and specific levels of polyethyleneglycol (PEG, mol. mass 1400–1600) — induced water stress has been described. To set up this system a perforated tubular glass vessel containing soil to support seedling growth was wrapped externally first with a layer of macroporous silica gel-G and then with three layers of a dialysis membrane of a lower exclusion limit (2000 mol. mass). Effects of 8 days of PEG — induced stress have been studied on uptake and translocation of N and P and growth of barley (Hordeum vulgare L. cv. KN 16) seedlings. Some of the noteworthy improvements of the system were exelusion of PEG from the plant consequent upon use of silica gel-membrane combination, shorter time (2 days) for the soil-plant-air continuum to attain steady state, and stability of the plant water potential over a period of a few days.     

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 common chemical extractant for estimating plant-available zinc in different soil types (peaty,red and alluvial)

Summary Relative efficiency of five chemical extractants for the extraction of available Zn in four different soils and its uptake by rice seedlings was studied in a pot culture experiment. The Zn extracted by dithizone-ammonium acetate showed a significant relationship with plant uptake whereas the values for other extractants except NH₄OAc (pH4.8) did not approach the level of significance. Among the soil properties studied, pH and CaCO₃ correlated negatively and organic matter and CEC positively with Zn uptake by rice plants.Contribution from the Soil Science and Agricultural Chemistry Dept., Banaras Hindu University, Varanasi-221005, India.Lecturer and Research Scholar of the Soil Science and Agricultural Chemistry Department, respectively.Lecturer and Research Scholar of the Soil Science and Agricultural Chemistry Department, respectively.     

Diffusion of phosphate to plant roots in soil

Summary Autoradiographs of rape (Brassica napus L.) seedlings growing in a Begbroke Sandy Loam treated to different P levels showed P accumulations near root apices of primary and lateral roots, without corresponding depletion from the adjacent soil, indicating marked translocation.Laterals less than 2 days old did not deplete the soil despite considerable P accumulations in them. Their growth and P uptake were enhanced when the growth of the primary root was checked. The length of root hairs decreased markedly with increasing P supply.The P depletion zones developed in the same way at all points along the primary axis (except for a short length behind the apex). At the highest P level the concentration of exchangeable P at the root surface was lowered by about 30% on day 2, by about 40% on day 4 and rose slowly after day 8.Whereas in P treated soils the depletion from within the root hair cylinder was fairly uniform, in the low P soil there was a continuous decrease in P concentrations toward the root surface, within the root hair zone.Soil Science Laboratory, Department of Agricultural Science, University of Oxford     

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.     

Numerical study of some age-dependent parameters in root nutrient uptake

Computer simulation of root nutrient uptake has become a very powerfull tool in the analysis of plant and soil characteristics. One of the shortcomings of earlier numerical models is the lack of a proper accounting for age-dependent root parameters. In this article we present an algorithm that not only allows for time-and/or space-varying root growth rates, convective moisture uptake, root density, initial distribution of nutrient, effective diffusion coefficients, and buffer power; but also accounts for time-varying root efflux, root absorption power and maximum nutrient influxive rate.Several elementary numerical examples of NH ₄ ⁺ , NO ₃ ^– , P and K uptake for roots with temporally varying characteristics are presented.Contribution from the Purdue Agric. Exp. Stn., W. Lafayette, IN. Journal Paper No. 9476.     

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.     

Root Morphology and Nitrogen Uptake of Maize Simultaneously Supplied with Ammonium and Nitrate in a Split-root System

We studied the response of maize (Zea mays L. cv. Anjou 256)to a simultaneous, but separated supply of ammonium and nitrate(localized supply, LS). A split-root system was used to supplyhalf of the roots with ammonium and the other half with nitrate.A homogeneously distributed supply of both nitrogen forms (HS)was the control treatment. Seedlings were grown for 12 d fromthe two-leaf to the three-leaf stage in hydroponics at threepH levels (4, 5·5 and 7). The total N concentration was3 mol m^-3. The split-root system was established by removingthe seminal root system and using only four nodal roots perplant. Total root length and root surface area were recordedautomatically with a modified Delta- T area meter. Other morphologicalroot traits (such as main axis length and diameter, number,density, and length of laterals) were recorded manually. Uptakeof ammonium and nitrate was measured by the depletion of thenutrient solution. As compared with LS, HS was superior in shootand root DM, total root length and root surface area, ammoniumand nitrate uptake and shoot nitrogen concentration, irrespectiveof pH level. This indicates that, also under field conditions,mixed ammonium and nitrate fertilization is only beneficialto plant growth if both N forms are evenly distributed in thesoil. At both HS and LS, ascending pH increased the ammonium:nitrateuptake ratio. At LS, declining pH induced a considerable shiftin the distribution of root DM, root length, and root surfacearea the nitrate-fed compartment.Copyright 1993, 1999 AcademicPress Maize, Zea may L., ammonium, nitrate, pH, root morphology, split-root     

The effects of root temperature and Ca supply on the growth and transpiration of cotton seedlings (Gossypium hirsutum,L.)

Summary Cotton seedlings were germinated in either tap water or a CaSO₄ solution and then grown for two days in nutrient solutions containing 0.1, 1.0, or 10.0 me Ca/1. They were then transferred to cultures having the same Ca variables and the roots subjected to temperatures of 26, 18, 15 and 12°C for four days. The fresh weight of all plant parts and leaf area increased with increasing root temperatures and with increasing Ca levels, the effect of Ca being most pronounced at the higher temperatures. The dry weight was increased by increasing root temperature; the effect of Ca was rather small. Water use increased with increasing temperature and was higher for the lowest Ca level than for the higher levels. The results emphasize the overriding effect of low temperature on water uptake. Calcium had little effect on the growth depression resulting from low root temperatures. Joint contribution from the Agronomy Department, Mississippi Agricultural Experiment Station, State College and the Soil Science Department, North Carolina Agricultural Experiment Station, Raleigh. Published with the approval of the Directors of Research as Journal Contribution No.1530 and Paper No.2389 of the Journal Series, respectively. This is a report of research conducted while the author was on sabbatical leave at North Carolina State University, February 1–September 10, 1965.     

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.     

Nutrient Supply and the Growth of the Seminal Root System in Barley: II. LOCALIZED, COMPENSATORY INCREASES IN LATERAL ROOT GROWTH AND RATES OP NITRATE UPTAKE WHEN NITRATE SUPPLY IS RESTRICTED TO ONLY PART OF THE ROOT SYSTEM

The development of the seminal root system, its ability to absorbnitrate, and effects on shoot growth were studied in barleyplants in nutrient solution. The roots received either a uniformsupply of 1.0 mM nitrate (controls), or a supply of the samesolution restricted to a 4-cm length of only one of the mainseminal roots (axes) on each plant, the remainder of the rootsystem receiving a solution containing a low concentration (0.01mM). Marked increases took place in both the growth of lateralroots and the absorption and transport of ¹⁵N-labelled nitrate(per unit root weight) from the zone locally supplied with 1.0mM nitrate. These effects appear largely to compensate for thedeficient supply of nitrate to the remainder of the root system,since after 14 d the relative growth rate (g g^–1d^–1)of the total plant equals that of the controls. Rates of ¹⁵N-nitrateuptake (per unit root weight) remain relatively uniform throughoutthe 29-d experiment, during which root axes develop from theirinitial unbranched form to a complex system of laterals. Theresults are discussed in relation to possible mechanisms bywhich coordination is maintained between root growth, ion uptake,and shoot growth.     

Nitrogen-tillage-residue management

Summary NCSWAP (nitrogen and carbon cycling in soil, water and plant) is a simulation model of the soil-crop-water system which integrates water flow dynamics, crop growth, N transformations, tillage and residue effects, soil temperature, and solute transport. A small plot field study was initiated in May of 1980 to determine the effects of N rate (2 or 20 g N/m²), tillage (rototill or no-till), and residue management system (residue return or noresidue) on soil parameters, and maize (Zea mays L.) production.Significant differences due to treatments (N rate, tillage, and residue) were not detected in 1981 for the measured soil-plant parameters including soil moisture, yield, and N uptake. Therefore, two representative treatment combinations (N rates of 2 or 20 g N/m²-tilled-no residue) characterized the field research data. Calculated and observed data sets were compared for several parameters including: (1) soluble NO₃–N, (2) N leaching losses (3) plant total-N and¹⁵N, (4) root growth, (5) soil moisture, and (6) fertilizer efficiency.The objectives of this study were to initiate the validation process of the model NCSWAP, and to illustrate how NCSWAP can be used as a research tool to infer operational characteristics of the N cycle.Contribution of the Soil and Water Management Research Unit, USDA-ARS, and the Department of Soil Science, University of Minnesota, St. Paul, MN 55108. Minn Agric. Exp. Sta., Sci. J., Ser. Paper 13907.Senior Laboratory Technician; Research Chemist, USDA-ARS and Professor; Professor of Soil Microbiology; and Soil Scientist, USDA-ARS and Assistant Professor; all Department of Soil Science, University of Minnesota, respectively. Inquiries about NCSWAP should be sent to J. A. E. Molina.