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     

A Shoot:Root Partitioning Model

A model for partitioning newly-synthesized structural dry matterbetween shoot and root is developed. It is based on a postulatedpartitioning function, which depends upon the relative levelsof carbon and nitrogen substrates, with parameters determiningthe control point and also the degree of control. The modelis used to investigate the relationships between plant specificgrowth rate, shoot:root ratio, and the specific activities ofshoot and root (which depend upon environment), during steady-stateexponential growth; the transient behaviour of the model isalso explored and oscillations in these quantities are obtained. Shoot:root ratio, specific growth rate, mathematical model, partition of assimilates     

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     

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     

Dynamic model of the response of a vegetative grass crop to light, temperature and nitrogen

Abstract A previously described growth model of the vegetative grass crop is extended to include a simple representation of the root system, uptake of nitrogen from a soil nitrogen pool, and response to fertilizer application. The model simulates the processes of light interception, photosynthesis, partitioning of new growth, leaf area expansion, growth and maintenance respiration, ageing of plant tissues, senescence, recycling of substrates from senescing tissues, nitrogen uptake by the plant, leaching, mineralization, and fertilizer application. A principal component of the model, nitrogen uptake, is assumed to depend positively on plant carbon substrate concentration and soil nitrogen concentration, and to be inhibited by plant nitrogen substrate concentration. The dynamic responses to different levels of soil nitrogen, of shoot and root growth, nitrogen uptake and root activity, carbon and nitrogen plant substrate concentrations, and the fraction of substrate carbon used by the shoots, are examined; realistic behaviour is observed. The model predicts nitrogen fertilizer responses of yield and plant nitrogen content, which are compared directly with experimental data; good agreement is obtained.     

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     

Modelling Water in Crops and Plant Ecosystems

A water submodel is described that is specifically designedfor use with plant growth simulators that represent internalplant substrates and variable shoot:root partitioning. The modelcalculates water flow from soil to root, root to shoot, andshoot to the atmosphere, for a closed-canopy situation. As presentedhere, the model has three state variables: the masses of waterin the soil, root and shoot, and represents the processes ofevapotranspiration, rainfall interception and evaporation fromthe canopy, and drainage. The Penman –Monteith equationis used for crop transpiration. The fluxes of water from soilto root, and root to shoot, are driven by water potential difference.Tissue water potential and its components are calculated fromtissue water content and other plant variables and parameters.The model is able to simulate osmoregulation and describes avariable relationship between tissue water potential, its componentsand relative water content, depending on growth conditions.The model has elsewhere been integrated with two plant ecosystemmodels: for grassland and forest. The specific implementationand simulations given are for the Hurley pasture model (Thornleyand Verberne, 1989), a temperate grass vegetative growth model.The model gives reasonable predictions for diurnal changes inwater potential, drying-down behaviour and other quantitieswithin the scope of the model. Simulation; model; water relations; crop growth; grass     

Simulating Growth and Root-shoot Partitioning in Prairie Grasses Under Elevated Atmospheric CO2and Water Stress

We constructed a model simulating growth, shoot-root partitioning,plant nitrogen (N) concentration and total non-structural carbohydratesin perennial grasses. Carbon (C) allocation was based on theconcept of a functional balance between root and shoot growth,which responded to variable plant C and N supplies. Interactionsbetween the plant and environment were made explicit by wayof variables for soil water and soil inorganic N. The modelwas fitted to data on the growth of two species of perennialgrass subjected to elevated atmospheric CO₂and water stresstreatments. The model exhibited complex feedbacks between plantand environment, and the indirect effects of CO₂and water treatmentson soil water and soil inorganic N supplies were important ininterpreting observed plant responses. Growth was surprisinglyinsensitive to shoot-root partitioning in the model, apparentlybecause of the limited soil N supply, which weakened the expectedpositive relationship between root growth and total N uptake.Alternative models for the regulation of allocation betweenshoots and roots were objectively compared by using optimizationto find the least squares fit of each model to the data. Regulationby various combinations of C and N uptake rates, C and N substrateconcentrations, and shoot and root biomass gave nearly equivalentfits to the data, apparently because these variables were correlatedwith each other. A partitioning function that maximized growthpredicted too high a root to shoot ratio, suggesting that partitioningdid not serve to maximize growth under the conditions of theexperiment.Copyright 1998 Annals of Botany Company plant growth model, optimization, nitrogen, non-structural carbohydrates, carbon partitioning, elevated CO₂, water stress,Pascopyrum smithii,Bouteloua gracilis, photosynthetic pathway, maximal growth     

Carbon allocation to shoots and roots in relation to nitrogen supply is mediated by cytokinins and sucrose: Opinion

In this paper we firstly show some general responses of biomass partitioning upon nitrogen deprivation. Secondly, these responses are explained in terms of allocation of carbon and nitrogen, photosynthesis and respiration, using a simulation model. Thirdly, we present a hypothesis for the regulation of biomass partitioning to shoots and roots.Shortly after nitrogen deprivation, the relative growth rate (RGR) of the roots generally increases and thereafter decreases, whereas that of the shoot decreases immediately. The increased RGR of the root and decreased RGR of the shoot shortly after a reduction in the nitrogen supply, cause the root weight ratio (root weight per unit plant weight) to increase rapidly.We showed previously that allocation of carbon and nitrogen to shoots and roots can satisfactorily be described as a function of the internal organic plant nitrogen concentration. Using these functions in a simulation model, we analyzed why the relative growth rate of the roots increases shortly after a reduction in nitrogen supply. The model predicts that upon nitrogen deprivation, the plant nitrogen concentration and the rate of photosynthesis per unit plant weight rapidly decrease, and the allocation of recently assimilated carbon and nitrogen to roots rapidly increases. Simulations show that the increased relative growth rate of the root upon nitrogen deprivation is explained by decreased use of carbon for root respiration, due to decreased carbon costs for nitrogen uptake. The stimulation of the relative growth rate of the root is further amplified by the increased allocation of carbon and nitrogen to roots. Using the simple relation between the plant nitrogen concentration and allocation, the model describes plant responses quite realistically.Based on information in the literature and on our own experiments we hypothesize that allocation of carbon is mediated by sucrose and cytokinins. We propose that nitrogen deprivation leads to a reduced cytokinin production, a decreased rate of cytokinin export from the roots to the shoot, and decreased cytokinin concentrations. A reduced cytokinin concentration in the shoot represses cell division in leaves, whereas a low cytokinin concentration in roots neutralizes the inhibitory effect of cytokinins on cell division. A reduced rate of cell division in the leaves leads to a reduced unloading of sucrose from the phloem into the expanding cells. Consequently, the sucrose concentration in the phloem nearby the expanding cells increases, leading to an increase in turgor pressure in the phloem nearby the leaf's division zone. In the roots, cell division continues and no accumulation of sugars occurs in dividing cells, leading to only marginal changes in osmotic potential and turgor pressure in the phloem nearby the root's cell division zone. These changes in turgor pressure in the phloem of roots and sink leaves affect the turgor pressure gradients between source leaf-sink leaf and source leaf-root in such a way that relatively more carbohydrates are exported to the roots. As a consequence RWR increases after nitrogen deprivation. This hypothesis also explains the strong relationship between allocation and the plant nitrogen status.     

Dry Matter Partitioning in a Determinate and an Indeterminate Cultivar of Vicia faba L. Under Contrasting Plant Distributions and Densities

Models for root: shoot, vegetative: generative and stem: leafpartitioning are presented to quantify dry matter partitioningof two contrasting genotypes of Vicia faba, an indeterminateand a ‘topless’. A third plant type, a determinateof which two to three inflorescences had been removed, was alsoincluded but behaved similarly to the intact plant. The root:shoot partitioning model predicts linear relationships betweenroot proportion and the product of air vapour pressure deficitand relative growth rate. Data from field experiments were consistentwith model prediction and coefficients estimated were similarfor both genotypes. Partitioning into pods was modelled as proportionalto the number of actively growing pods younger than 1000°Cd. Coefficients estimated were similar in both genotypes butdifferent between densities and years. In the indeterminategenotype, stem: leaf ratio was allometric throughout, whereasleaf growth ceased but stem growth continued in the determinateafter formation of the terminal inflorescence. Relatively moredry matter was allocated to stems than to leaves in high thanin low densities. In conclusion, the main differences in drymatter partitioning between genotypes concern leaf: stem partitioningduring early pod filling and pod partitioning due to pod numberdifferences. Partitioning, root, shoot, leaf, stem, Vicia faba L.     

Response of Dry Matter Partitioning, Growth, and Carbon and Nitrogen Levels in the Tomato Plant to Changes in Root Temperature: Experiment and Theory

Growth, dry matter partitioning, and the levels of various substancesin tomato plants grown in liquid nutrient culture at four roottemperatures (15, 20, 25, and 30°C) in controlled environmentcabinets were measured. In order to understand the role of roottemperature in growth and development, the partitioning modelof Thornley (1972) was used to try and describe the observedbehaviour. The steady-state behaviour could be simulated reasonablyby the model, although certain aspects of the time-course datapresent problems. It was observed that relative growth rateis proportional to the product of the carbon and nitrogen levels(one of the two principal assumptions of the model). The modelalso made realistic predictions about the influence of roottemperature on the fraction of the plant dry matter in the root,and on the carbon and nitrogen levels. The effect of root temperaturewas incorporated into the model by assuming that certain valuesof specific root activity correspond to the different temperatures.In addition the model predicted an experimentally reasonablerelation between net assimilation rate and relative growth rate.     

Relative Nitrogen Limitation at Steady-state Nutrition as a Determinant of Plasticity in Five Grassland Plant Species

Partitioning of biomass between roots and different shoot partshas often been used to explain the response of plants to variationsin resource availability. There are still many uncertaintiesin the importance of this trait for plant performance, and clearguidelines on how partitioning should be quantified in relationto growth rate and resource supply are of fundamental importancefor such an understanding. This paper reports an attempt toshow how plant nitrogen status relates to root:shoot partitioningand other plastic responses, in a manner that can be used forquantitative predictions. The reactions to nitrogen limitationof five grassland plant species, with different ecological demands,were compared. The species used were the forbs Polygala vulgarisand Crepis praemorsa, and the grasses Danthonia decumbens, Agrostiscapillaris and Dactylis glomerata. The experiment was conductedin a climate chamber where the plants were grown hydroponically(1) under non-limiting nutrient conditions and (2) at a steady-statenitrogen limitation, which enabled the plants to express halfof their growth potential. The relative growth rate (RGR) ofthe species was strongly related to plant nitrogen concentration(PNC) and leaf area ratio (LAR), whereas the effects on netassimilation rate (NAR) were very small. Despite large differencesin maximum relative growth rate, the species showed remarkablesimilarities in dry matter partitioning between root and shoot.It is concluded that root:shoot partitioning can be treatedas a direct function of the relative resource limitation ofthe plant. The difficulty of attaining well-defined levels ofresource limitation in soil, other solid substrates and manyhydroponic systems may be the most important reason for thedivergent results in earlier studies. Better knowledge of soil-rootinteractions, and plant responses to the whole span of resource-supplylevels, is required for a thorough understanding of how nutrientslimit growth. Copyright 1999 Annals of Botany Company Growth rate, plant strategies, plasticity, partitioning, biomass, nitrogen, nutrient limitation, grassland.     

Controls of biomass partitioning between roots and shoots: Atmospheric CO2 enrichment and the acquisition and allocation of carbon and nitrogen in wild radish

Summary The effects of CO₂ enrichment on plant growth, carbon and nitrogen acquisition and resource allocation were investigated in order to examine several hypotheses about the mechanisms that govern dry matter partitioning between shoots and roots. Wild radish plants (Raphanus sativus × raphanistrum) were grown for 25 d under three different atmospheric CO₂ concentrations (200 ppm, 330 ppm and 600 ppm) with a stable hydroponic 150 mol 1^–1 nitrate supply. Radish biomass accumulation, photosynthetic rate, water use efficiency, nitrogen per unit leaf area, and starch and soluble sugar levels in leaves increased with increasing atmospheric CO₂ concentration, whereas specific leaf area and nitrogen concentration of leaves significantly decreased. Despite substantial changes in radish growth, resource acquisition and resource partitioning, the rate at which leaves accumulated starch over the course of the light period and the partitioning of biomass between roots and shoots were not affected by CO₂ treatment. This phenomenon was consistent with the hypothesis that root/shoot partitioning is related to the daily rate of starch accumulation by leaves during the photoperiod, but is inconsistent with hypotheses suggesting that root/shoot partitioning is controlled by some aspect of plant C/N balance.     

Transport Exchange of Carbon,Nitrogen and Water in the Context of Whole Plant Growth and Functioning — Case History of a Nodulated Annual Legume

The economy of carbon, nitrogen and water during growth of nodulated, nitrogen-fixing plants of white lupin (Lupinus albus L.) was studied by measuring C, N and H₂O content of plant parts, concentrations of C and N in bleeding sap of xylem and phloem, transpirational losses of whole shoots and shoot parts, and daily exchanges of CO₂ between shoot and root parts and the surrounding atmosphere. Relationships were studied between water use and dry matter accumulation of shoot and fruits, and between net photosynthesis rate and leaf area, transpiration rate and nitrogen fixation. Conversion efficiencies were computed for utilization of net photosynthate for nitrogen fixation and for production of dry matter and protein in seeds. Partitioning of the plant's intake of C, N and H₂O was described in terms of growth, transpiration, and respiration of plant parts. An empirically-based model was developed to describe transport exchanges in xylem and phloem for a 10-day interval of growth. The model depicted quantitatively the mixtures of xylem and phloem streams which matched precisely the recorded amounts of C, N and H₂O assimilated, absorbed or consumed by the various parts of the plant. The model provided information on phloem translocation of carbon and nitrogen to roots from shoots, the cycling of carbon and nitrogen through leaves, the relationship between transpiration and nitrogen partitioning to shoot organs through the xylem, the relative amount of the plant's water budget committed to phloem translocation, and the significance of xylem to phloem transfer of nitrogen in stems as a means of supplying nitrogen to apical regions of the shoot.     

Effect of Root and Shoot Meristem Temperature on Shoot to Root Dry Matter Partitioning and the Internal Concentrations of Nitrogen and Carbohydrates in Maize and Wheat

Maize (Zea mays L.) and spring wheat (Triticum aestivum L.)were grown in nutrient solution at uniformly high air temperature(20 °C), but different root zone temperatures (RZT 20, 16,12 °C). To manipulate the ratio of shoot activity to rootactivity, the plants were grown with their shoot base includingthe apical meristem either above (i.e. at 20 °C) or withinthe nutrient solution (i.e. at 20, 16 or 12 °C). In wheat, the ratio of shoot:root dry matter partitioning decreasedat low RZT, whereas the opposite was true for maize. In bothspecies, dry matter partitioning to the shoot was one-sidedlyincreased when the shoot base temperature, and thus shoot activity,were increased at low RZT. The concentrations of non-structuralcarbohydrates (NSC) in the shoots and roots were higher at lowin comparison to high RZT in both species, irrespective of theshoot base temperature. The concentrations of nitrogen (N) inthe shoot and root fresh matter also increased at low RZT withthe exception of maize grown at 12 °C RZT and 20 °Cshoot base temperature. The ratio of NSC:N was increased inboth species at low RZT. However this ratio was negatively correlatedwith the ratio of shoot:root dry matter partitioning in wheat,but positively correlated in maize. It is suggested that dry matter partitioning between shoot androots at low RZT is not causally related to the internal nitrogenor carbohydrate status of the plants. Furthermore, balancedactivity between shoot and roots is maintained by adaptationsin specific shoot and root activity, rather than by an alteredratio of biomass allocation between shoot and roots.Copyright1994, 1999 Academic Press Wheat, Triticum aestivum, maize, Zea mays, root temperature, shoot meristem temperature, biomass allocation, shoot:root ratio, carbohydrate status, nitrogen status, functional equilibrium     

Shoot and Root Activities During Steady-state Plant Growth

A simple model for steady-state plant growth is described. Thegrowth constant, measured during steady-state exponential growth,is related to the specific activities of the shoot and the root,enabling the effects of certain environmental variables (light,carbon dioxide and nitrogen) on the growth constant to be examined.The model is used to interpret data on the growth kinetics ofwheat (Macdowall, 1972a, b, c).     

Effects of CO2 enrichment and intraspecific competition on biomass partitioning, nitrogen content and microbial biomass carbon in soil of perennial ryegrass and white clover

Seedlings of perennial ryegrass (Lolium perenne L. cv. Parcour)and white clover (Trifolium repens L. cv. Karina) grown at fivedifferent plant densities were exposed to ambient (390 ppm)and elevated (690 ppm) CO₂ concentrations. After 43 d the effectsof CO₂ enrichment and plant density on growth of shoot and root,nitrogen concentration of tissue, and microbial biomass carbon(C_mic) in soil were determined. CO₂ enrichment of Lolium perenneincreased shoot growth on average by 17% independent of plantdensity, while effects on root biomass ranged between -4% and+ 107% due to an interaction with plant density. Since tilernumber per plant was unaffected by elevated CO₂, the small responseof shoot growth to CO₂ enrichment was atributed to low sinkstrength. A significant correlation between nitrogen concentrationof total plant biomass and root fraction of total plant drymatter, which was not changed by CO₂ enrichment, indicates thatnitrogen status of the plant controls biomass partitioning andthe effect of CO₂ enrichment on root growth. Effects of elevatedCO₂ and plant density on shoot and root growth of Trifoliumrepens were not significantly interacting and mean CO₂-relatedincrease amounted to 29% and 66%, respectively. However, growthenhancement due to elevated CO₂ was strongest when leaf areaindex was lowest. Total amounts of nitrogen in shoots and rootswere bigger at 690 ppm than at 390 ppm CO₂. There was a significantincrease in C_mic in experiments with both species whereas plantdensity had no substantial effect. Key words: CO₂ enrichment, intraspecific competition, biomass partitioning, Lolium perenne, Trifolium repens, grassland     

Ara-Rhizotron: An Effective Culture System to Study Simultaneously Root and Shoot Development of Arabidopsis

Studying Arabidopsis thaliana (L.) Heynh. root development in situ at the whole plant level without affecting shoot development has always been a challenge. Such studies are usually carried out on individual plants, neglecting competition of a plant population, using hydroponic systems or Agar-filled Petri dishes. Those both systems, however, present some limitations, such as difficulty to study precisely root morphogenesis or time-limited culture period, respectively. In this paper, we present a method of Arabidopsis thaliana (L.) Heynh. cultivation in soil medium, named “Ara-rhizotron”. It allows the non-destructive study of shoot and root development simultaneously during the entire period of vegetative growth. In this system, roots are grown in 2D conditions, comparable to other soil cultures. Moreover, grouping several Ara-rhizotrons in a box enables the establishment of 3D shoot competition as for plants grown in a population. In comparison to a control culture grown in pots in the same environmental conditions, the Ara-rhizotron resulted in comparable shoot development in terms of dry mass, leaf area, number of leaves and nitrogen content. We used this new culture system to study the effect of irrigation modalities on plant development. We found that irrigation frequency only affected root partitioning in the soil and shoot nitrogen content, but not shoot or root growth. These effects appeared at the end of the vegetative growth period. This experiment highlights the opportunity offered by the Ara-rhizotron to point out tardy effects, affecting simultaneously shoot development and root architecture of plants grown in a population. We discuss its advantages in relation to root development and physiology, as well as its possible applications.