Compensatory Changes in the Partitioning of Dry Matter in Relation to Nitrogen Uptake and Optimal Variations in Growth

Equations are derived relating relative growth rate (RGR) toroot:shoot ratio, root length, nitrogen inflow rate, leaf area,photosynthesis and carbon and nitrogen concentrations in theplant. The extents to which changes in specific root lengthand root: shoot ratio can compensate for the effects of lowN availability upon RGR are examined. Such responses could haveseveral compensatory functions: maximizing RGR; maintaininggrowth in which the activities of root and shoot limit RGR equally;and maximizing the efficiency of increase in RGR. Growth, nitrogen, carbon, dry matter, partitioning, root:shoot ratio, relative growth rate     

A Model of Shoot: Root Partitioning with Optimal Growth

A shoot: root partitioning model is presented, which is a developmentof previous approaches in the area. The model incorporates asa physiologically reasonable apparent ‘goal’ forthe plant, the assumption that the partitioning of growth betweenthe shoot and root maximizes the plant specific growth ratein balanced exponential growth. The analysis is concerned principallywith plant growth being a function of carbon and nitrogen only,although it is indicated how other nutrients, or growth factors,may be incorporated. Plant growth is driven by the environmentalconditions, and partitioning is defined entirely in terms ofthe shoot: root ratio and carbon and nitrogen status of theplant. In its basic form the model requires the definition ofa single plant growth parameter, along with the shoot and rootspecific activities and structural composition. Shoot: root partitioning, specific growth rate, vegetative phase     

A Model of the Partitioning of Growth between the Shoots and Roots of Vegetative Plants

A model of the partitioning of new growth between the shootsand roots of vegetative plants is presented. There are two partitioningfunctions, involving one partitioning parameter, which describethe priorities for new growth in both the shoots and roots.The dynamic responses, to changes in the environment and toshoot defoliation, of shoot and root specific growth rates,shoot: root ratio, and carbon and nitrogen substrate levels,are examined; realistic behaviour is observed. Balanced exponentialgrowth solutions are also examined and it is concluded thatrelationships between some derived plant growth quantities maybe non-unique, thus emphasizing the need for a critical understandingof the underlying physiological processes involved in plantgrowth. Mathematical model, partitioning of assimilates, shoot: root ratio, specific growth rate, carbon and nitrogen substrate levels     

Nitrogen Uptake and Plant Growth II. An Empirical Model of Vegetative Growth and Partitioning

An empirical model of vegetative plant growth is presented.The model is based on experimental data on Polygonum cuspidatum,which showed (1) that the partitioning of dry matter and nitrogenamong organs was linearly related to the nitrogen concentrationof the whole plant and (2) that leaf thickness was negativelycorrelated with leaf nitrogen concentration. The model properlydescribes the behaviour of plants. Steady-state solutions ofthe model give the relative growth rate, specific leaf weight,and partitioning of dry matter and nitrogen among organs withthe net assimilation rate and the specific absorption rate asenvironmental variables. The effect of nitrogen removal on drymatter and nitrogen partitioning was examined as non-steady-statedynamic solutions of the model. The model predicted not onlyreduced leaf growth and enhanced root growth but also a fluxof nitrogen from the leaf to the root, which agreed with theexperimental results. Mathematical model, partitioning of dry matter and nitrogen, plant nitrogen, relative growth rate, shoot: root ratio, specific leaf weight     

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     

Effects of root pruning of wheat at different nitrogen levels

In two experiments, wheat plants growing in solutions of different nitrogen concentration were subjected to root pruning. In higher concentrations of nitrogen the growth rate was higher, and the proportional allocation of growth to shoot higher, but pruning did not affect the allocation of growth at either level of nitrogen. This result gives no support to Thornley's source-sink model of the control of shoot: root ratio.     

A Review of Evidence on the Control of Shoot: Root Ratio, in Relation to Models

Four basic models exist for the control of shoot: root ratio(S: R): (a) allometric models, proposing a fixed ratio of shootgrowth rate to root growth rate; (b) functional equilibriummodels, based on the ratio of shoot activity to root activity;(c) the Thornley model, based on carbon and nitrogen uptakeand transport, (d) hormone models, generally suggesting theroot produces a hormone that controls the shoot and vice versa.Models (a) and (b) are empirical, and therefore provide no testof the processes operating. Ontogenetic changes in S: R for fibrous-rooted herbs could befitted by a modified Thornley model. Ontogenetic effects mustbe excluded in judging other effects. Responses of S: R to deficits of water, major inorganic nutrients,light and carbon dioxide, and to defoliation and root pruning,usually conform to Thornley's model. With current knowledgeThornley's model cannot usefully be applied to minor nutrients,nutrient toxicity or temperature differences. S: R changes atreproduction usually conform to Thornley's model if it is assumedthat young reproductive structures are a strong sink, but thisbegs the question of what determines sink strength. There areapparent exceptions to most of these responses, which shouldbe studied further. Phytohormones can influence S: R, but may not be the controloperating in the normal, intact plant. Most of the availableevidence is compatible with a source-sink model of Thornley'stype, and therefore does not demand a hormonal theory of S:R control. There is a need for more critical tests. Shoot: root ratio, strategy, partitioning, models     

Analysis of Dry Matter Partitioning in Dactylis glomerata during Vegetative Growth Using a Carbon Budget Model

The dry matter partitioning in vegetative plants of Dactylisglomerata was studied from experiments performed in controlledenvironments. Plants were grown hydroponically in growth chambers,at two constant temperatures (17 and 25 °C). In both experimentsthe root fraction decreased regularly with time, an effect thatwas more accentuated in the higher temperature regime. In orderto explain the change in dry matter partitioning, the experimentalshoot and root growth were analysed using a carbon budget modelwhich includes shoot and root maintenance requirements. Themodel predicts a relationship between the root specific growthrate and the product of shoot specific growth rate and shootto root dry weight ratio. In the range of experimental accuracy,this relationship was found to be linear at both temperatures,which should indicate that the partitioning coefficients andthe root maintenance coefficient remained constant during vegetativegrowth. The effect of temperature on the value of these coefficientscan be specified from a linear regression analysis. Between17 and 25 °C, the root maintenance coefficient increasedby about a factor of two, whereas the partitioning coefficientsdid not vary significantly. On the basis of these results, itwas shown that the decrease in root fraction during vegetativegrowth should be mainly attributed to the decrease in net specificactivity of shoots.Copyright 1994, 1999 Academic Press Dactylis glomerata L., vegetative growth, model, partitioning, root:shoot ratio, shoot specific activity, maintenance requirements     

Simulation of Rhythmic Tree Growth under Constant Conditions

The observed rhythmic growth of trees under relatively uniform environmental conditions has been ascribed by some authors to endogenous factors, by others to slight fluctuations of environmental factors. A model for the simulation of rhythmic growth was developed based on the assumption that endogenous rhythms can result from feedback interaction between two potentially continuous processes, like shoot and root growth, if the slower process is rate limiting for the faster one. Rhythmic growth in trees would be the consequence of feedback mechanisms needed for maintaining a constant shoot: root ratio. Period length of the rhythms depends upon the rates of the growth processes involved. Environmental factors modify period length through affecting growth rates. Growth patterns predicted by the model compare well with growth measurements of tropical trees. The transition from intermittent to continuous growth, as observed under certain conditions, can be simulated by varying a single parameter in the model.     

The Rate of Growth and Partitioning of Assimilates in Young Grass Plants: A Mathematical Model

A model describing the increase in weight with time of younggrass plants is formulated. The parameters are the relativegrowth rates of the root and shoot systems; k, the ratio ofthe relative growth rate of the root system relative to thatof the shoot system; b, the weight of the root system when thatof the shoot system is unity, and u the rate of increase inweight of the whole plant per unit of shoot system per unitof time, k and b are the constants in the allometric formula,r = bs^k where r and are the weights of the root and shoot systems.The model enables the effect of changes in the distributionof assimilates between the root and shoot systems upon the rateof growth of the plant to be assessed. Data from a number ofexperiments are analysed in this manner and the significanceof the results discussed.     

Nitrogen, phosphorus and light effects on growth and allocation of biomass and nutrients in wild rice

Separating plastic from ontogenetic and growth-limiting responses of plants to changes in resource availability can be challenging because there are a total of eight combinations of these three types of responses. These can, however, be uniquely distinguished on plots of root:shoot ratios against total biomass through time. We used this approach to separate ontogenetic, plastic, and growth-limiting responses of wild rice (Zizania palustris L.) to changes in nitrogen, phosphorus, and light availabilities. Relative growth rate was limited primarily by nitrogen but responded to increased light and phosphorus after nitrogen limitations were alleviated. Nitrogen addition increased relative growth rate because it simultaneously increased unit leaf rate, specific leaf area, and leaf weight ratio. Increased light did not change relative growth rate because decreased specific leaf area and leaf weight ratio compensated the increased unit leaf rate. Phosphorus did not change either relative growth rate or its underlying components. Plants responded ontogenetically to increased nitrogen and light availabilities by accelerating their developmental rate, and plastically by decreasing or increasing their root:shoot ratios, respectively. Plants did not respond either ontogenetically or plastically to increased phosphorus availability. Ontogenetic changes in growth can be separated from plastic and growth-limiting responses by plotting root:shoot ratio against total biomass in the context of the eight possible responses identified above, and also by examining how the underlying components of relative growth rate respond.     

营养生长期植物冠根比及其对环境因子的响应

将光强、温度和土壤水势等环境因子对植物光合、呼吸、同化物运输及生长等生理过程的影响结合起来,并考虑到各生理过程之间的交互作用,建立了一个营养生长期内植物条根比变化及对环境因子综合响应的模型。模型的运行结果表明,该模型与许多实验现象均相吻合。     

Accumulation of nitrate in the shoot acts as a signal to regulate shoot-root allocation in tobacco†

Mutants and transformants of tobacco (Nicotiania tabacum L. cv Gatersleben 1) with decreased expression of nitrate reductase have been used to investigate whether nitrate accumulation in the shoot acts as a signal to alter allocation between shoot and root growth. (a) Transformants with very low (1–3% of wild-type levels) nitrate reductase activity had growth rates, and protein, amino acid and glutamine levels similar to or slightly lower than a nitrate-limited wild-type, but accumulated large amounts of nitrate. These plants should resemble a nitrate-limited wild-type, except in responses where nitrate acts as a signal. (b) Whereas the shoot:root ratio decreases from about 3.5 in a well-fertilized wild-type to about 2 in a nitrate-limited wild-type, the transformants had a very high shoot:root ratio (8–10) when they were grown on high nitrate. When they were grown on lower nitrate concentrations their shoot:root ratio declined progressively to a value similar to that in nitrate-limited wild-types. Mutants with a moderate (30–50%) decrease of nitrate reductase also had a small but highly significant increase of their shoot:root ratio, compared to the wild-type. The increased shoot:root ratio in the mutants and transformants was due to a stimulation of shoot growth and an inhibition of root growth. (c) There was a highly significant correlation between leaf nitrate content and the shoot:root ratio for eight genotypes growing at a wide range of nitrate supply. (d) A similar increase of the shoot:root ratio in nitrate reductase-deficient plants, and correlation between leaf nitrate content and the shoot:root ratio, was found in plants growing on ammonium nitrate. (f) Split-root experiments, in which the transformants were grown with part of their root system in high nitrate and the other part in low nitrate, showed that root growth is inhibited by the accumulation of nitrate in the shoot. High concentrations of nitrate in the rooting medium actually stimulate local root growth. (g) The inhibition of root growth in the transformants was relieved when the transformants were grown on limiting phosphate, even though the nitrate content of the root remained high. This shows that the nitrate-dependent changes in allocation can be overridden by other signals that increase allocation to root growth. (h) The reasons for the changed allocation were investigated in transformants growing normally, and in split-root culture. Accumulation of nitrate in the shoot did not lead to decreased levels of amino acids or protein in the roots. However, it did lead to a strong inhibition of starch synthesis and turnover in the leaves, and to decreased levels of sugars in the root. The rate of root growth was correlated with the root sugar content. It is concluded that these changes of carbon allocation could contribute to the changes in shoot and root growth.     

The Effects of Infection by Rust (Puccinia lagenophorae Cooke) on the Growth of Groundsel (Senecio vulgaris L.) Cultivated under a Range of Nutrient Concentrations

Groundsel (Senecio vulgaris L.), healthy or infected with therust fungus Puccinia lagenophorae Cooke, was grown at a rangeof nutrient concentrations in sand culture. There were statisticallysignificant interactions between the effects of infection andnutrient supply upon the dry weights of stems, leaves, rootsand reproductive tissues, leaf area and cumulative capitulumproduction. This interaction occurred since infection causedsignificant inhibitions of growth only at moderate or high nutrientconcentrations. At low concentrations rusted plants were similarto or slightly larger than controls. Both in controls and rustedplants root: shoot ratios increased as nutrient supply declined.The ratio of root: shoot dry weight was consistently reducedby infection whilst root length: leaf area ratio was relativelyunchanged. More detailed investigations confirmed that infection had littleeffect on plant growth under nutrient deficient conditions despitesuppression of the host's ability to increase root: shoot ratiosin response to nutrient stress. This reflected the inhibitionof relative growth rates in rusted plants at high but not lownutrient concentrations, which in turn reflected reduced netassimilation rates (NAR). Increases in leaf-area ratio (LAR)often ameliorated the decline in NAR in rusted plants. Senecio vulgaris L., Puccinia lagenophorae Cooke, nutrient deficiency, growth, root: shoot ratio     

The Effect of Root Temperature on Growth and Uptake of Ammonium and Nitrate by Brassica napus L. in Flowing Solution Culture: I. GROWTH

Macduff, J. H., Hopper, M. J. and Wild, A. 1987. The effectof root temperature on growth and uptake of ammonium and nitrateby Brassica napus L. in flowing solution culture. I. Growth.—J.exp. Bot. 38: 42–52 Oilseed rape (Brassica napus L. cv. Bien venu) was grown for49 d in flowing nutrient solution at pH 6?0 with root temperaturedecrementally reduced from 20?C to 5?C; and then exposed todifferent root temperatures (3, 5, 7, 9, 11, 13,17 or 25?C)held constant for 14 d. The air temperature was 20/15?C day/nightand nitrogen was supplied automatically to maintain 10 mmolm^–3 NH₄NO₃ in solution. Total dry matter production wasexponential with time and similar at all root temperatures givinga specific growth rate of 0?0784 g g^–1 d^–1. Partitioningof dry matter was influenced by root temperature; shoot: rootratios increased during treatment at 17?C and 25?C but decreasedafter 5 d at 3?C and 5?C. The ratio of shoot specific growthrate: root specific growth rate increased with the ratio ofwater soluble carbohydrates (shoot: root). Concentrations ofwater soluble carbohydrates in shoot and root were inverselyrelated to root temperature; at 3, 5 and 7?C they increasedin stem + petioles throughout treatment, coinciding with a decreasein the weight of tissue water per unit dry matter. These resultssuggest that the accumulation of soluble carbohydrates at lowtemperature is the result of metabolic imbalance and of osmoticadjustment to water stress. Key words: Brassica napus, oilseed rape, root temperature, specific growth rate     

Optimization of Plant Root: Shoot Ratios and Internal Nitrogen Concentration

A general theoretical approach is developed to analyze the morphologicaland physiological responses of plants to nitrogen availability.The optimal leaf-nitrogen concentration and corresponding optimalroot: shoot ratio which maximize relative growth rate are foundquantitatively as a function of root—specific activitywhich is assumed to be a function of soil nitrogen availability.The cost of increasing tissue nitrogen concentration is foundto be primarily related to an increase in allocation to roots.Predictions of the analysis are consistent with previous theoriesand general empirical findings, suggesting that plants respondoptimally to soil nitrogen. Relative growth rate is predictedto be a nearly linear function of whole-plant nitrogen concentrationand shoot fraction is a monotonically increasing function oftissue nitrogen concentration when plants respond optimallyto soil nitrogen availability. Plant growth, root:shoot ratios, biomass allocation, nitrogen productivity, optimization     

The nitrogen handling characteristics of white clover (Trifolium repens L.) cultivars and a perennial ryegrass (Lolium perenne L.) cultivar

Ryegrass (Lolium perenne L.) and white clover (Trifolium repens L.) have contrasting responses to soil mineral N availability and clover has the ability to fix atmospheric N(2) symbiotically. It has been hypothesized that these differences are the key to understanding grass-clover coexistence and vegetative dynamics in pastures. However, the whole plant response of clover and ryegrass to mineral N availability has not been fully characterized and inter-cultivar variability in the N-handling dynamics of clover has not been assessed. A detailed experimental study to address these issues was undertaken. For all clover cultivars and ryegrass, mass specific mineral N uptake rates (of whole plants) were similar saturating functions of mineral N availability. For all clover cultivars total N assimilation rates, whole plant C : N ratios and root : shoot ratios were independent of mineral N availability. Clover growth rates were also independent of mineral N availability except for a slight (<10%) reduction at very low N availability levels. Specific N(2) fixation rate (whole plant) was precisely controlled to ensure fixation balanced the deficit between mineral N uptake and the total N assimilation required to maintain constant whole plant C : N ratio. There was always a deficit between N uptake and the total N assimilation required to maintain C : N ratio. Consequently, some N(2) fixation remained engaged even at high mineral N availability levels. All inter-cultivar variation in N(2) fixation dynamics could be attributed to variations in growth rate. Clover mass specific growth rate declined as plant size increased. Ryegrass specific growth rate, whole plant C : N ratio and root : shoot ratio were dependent on N availability. Increased N availability led to increased growth rate and decreased C : N and root : shoot ratios. Specific growth rate was also dependent on plant size, growth rate declining as plant size increased. It is concluded that clover inter-cultivar variation in field performance is unlikely to be a consequence of variation in N-handling characteristics. Inter-cultivar differences in growth rate are likely to be a much more important source of variation.     

A Balanced Quantitative Model for Root: Shoot Ratios in Vegetative Plants

The vegetative growth of a two-component plant consisting ofroot and shoot only is considered in terms of the transportand utilization of two required substrates, one providing carbonand the other providing nitrogen. The model provides a quantitativescheme for examining how root: shoot ratios depend upon thespecific activities of root and shoot and hence environment.It has been shown that the total shoot activity is proportionalto the total root activity in a plant undergoing steady-stategrowth.     

Growth, Nitrogen Uptake and Flow in Maize Plants Affected by Root Growth Restriction

The objective of the present study was to investigate the influence of a reduced maize root-system size on root growth and nitrogen (N) uptake and flow within plants. Restriction of shoot-borne root growth caused a strong decrease in the absorption of root: shoot dry weight ratio and a reduction in shoot growth. On the other hand, compensatory growth and an increased N uptake rate in the remaining roots were observed. Despite the limited long-distance transport pathway in the mesocotyl with restriction of shoot-borne root growth, N cycling within these plants was higher than those in control plants, implying that xylem and phloem flow velocities via the mesocotyl were considerably higher than in plants with an intact root system. The removal of the seminal roots in addition to restricting shoot-borne root development did not affect whole plant growth and N uptake, except for the stronger compensatory growth of the primary roots. Our results suggest that an adequate N supply to maize plant is maintained by compensatory growth of the remaining roots, increased N uptake rate and flow velocities within the xylem and phloem via the mesocotyl, and reduction in the shoot growth rate.