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     

Pre-flowering Growth and Development of the Inflorescences of Maize: II. ACCUMULATION AND PARTITIONING OF DRY MATTER AND NITROGEN BY INFLORESCENCES

The accumulation and partitioning of dry matter and nitrogenwere examined in the developing tassel and two uppermost earshoots of field-grown maize under varying levels of appliednitrogen and times of sowing. Accumulation of dry matter and nitrogen within an axillary branchalways favoured the ear over the husk and shank. Dry matterand nitrogen accumulated faster in the first ear than in thetassel or second ear and the partitioning between inflorescencesof dry matter and nitrogen was not affected by treatment. Therelative rate of growth, RGR(dry matter), of the first and secondear shoots increased by up to 42% at high levels of appliednitrogen and with early sowing. In contrast, the relative rateof accumulation of nitrogen (RNAR) was not sensitive to N supply,although it was reduced, on average, by 22% at the late timeof sowing. We conclude that accumulation, but not partitioning, of drymatter and nitrogen between developing inflorescences of maizeare altered by nitrogen application, time of sowing, and positionof the inflorescence on the stem. Key words: Maize, N-application, partitioning, inflorescence, sowing time     

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.     

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     

Combined Effects of Partial Defoliation and Nutrient Availability on Cloned Betula pendula Saplings: I CHANGES IN GROWTH, PARTITIONING AND NITROGEN UPTAKE

The combined effects of partial defoliation and nutrient availabilityon dry matter accumulation and partitioning, and on nitrogenuptake and partitioning, were studied in cloned Betula pendulaRoth saplings. The saplings were randomly assigned to differentnutrient levels (5, 1·5 and 0·5 mol Nm^–3)in aerated nutrient culture and to the following defoliationtreatments: (1) control (no damage), (2)damage of the developingmain stem leaves (half of the leaf lamina removed), and (3)removalof the developing main stem leaves (entire leaf lamina removed).Measured in terms of cumulative whole-plant dry weight (includingremoved leaf tissue), the birch saplings were unable to compensatefor the loss of the developing leaves (treatment 3) during the14 d study period. In response to leaf removal (treatment 3)the mean final percentage reduction in whole-plant dry weightwas actually greater than the initial mean percentage reductioncaused by the removal itself; the magnitude of the final reductionwas independent of nutrient availability. After removal of thedeveloping leaves, branch growth was favoured at the expenseof the growth of the rest of the shoot; the relative branchgrowth was most pronounced at the highest nutrient level. Atthe two highest nutrient levels the nitrogen uptake of the saplingswith the developing leaves removed was less than that of undamagedsaplings. We suggest, however, that the incapacity of the saplingsfor compensatory growth after removal of the developing leaveswas primarily due to the decreased total carbon gain of thesaplings rather than to the decreased nitrogen uptake rate. Key words: Partial defoliation, nutrient availability, birch sapling, dry matter, nitrogen     

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     

Nitrogen Uptake and Plant Growth I. Effect of Nitrogen Removal on Growth of Polygonum cuspidatum

Polygonun cuspidatum was grown hydroponically to examine theeffect of nitrogen removal from the nutrient solution upon plantgrowth and the partitioning of dry matter and nitrogen amongorgans. Nitrogen removal reduced the growth rate mainly dueto the reduced growth of leaf area. Accelerated root growthwas observed only in plants which earlier had received highlevels of nitrogen. Nitrogen removal caused almost exclusiveallocation of available nitrogen to root growth. Nitrogen fluxfrom the shoot to the root occurred in plants which had receivedlow nitrogen. Not only was net assimilation rate (NAR) littleaffected by nitrogen removal, but it also was not correlatedwith the concentration of leaf nitrogen on an area basis. Light-saturatedCO₂ exchange rate (CER) was highly correlated with the concentrationof leaf nitrogen. Nitrogen use efficiency (NUE) in CER (CERdivided by leaf nitrogen) remained constant against leaf nitrogen,indicating efficient use of nitrogen under light saturation,while NUE in terms of NAR decreased with higher concentrationof leaf nitrogen. Polygonum cuspidatum Sieb. et Zuce., CO₂ exchange rate, growth analysis, leaf nitrogen, net assimilation rate, nitrogen use efficiency, partitioning of dry matter and nitrogen     

Effects of untreated two‐phase olive mill pomace on potted olive plantlets

Two‐phase pomace represents an important environmental problem in Mediterranean areas. With its high organic content, direct application of two‐phase pomace in the field is recommended to improve soil organic carbon levels and fertility. However, this does not consider any antagonistic effects that this application might have on root proliferation and biomass partitioning. We studied the effects of untreated two‐phase pomace on properties of growth substrate, and on shoot and root growth and biomass allocation of potted olive plantlets. A pot experiment was carried out in a greenhouse over 90 days, with five levels of two‐phase pomace and using two olive cultivars. The effects on shoot growth, leaf pigment content and maximum quantum efficiency of photosystem II (ΦPSII) were assessed each month. After 90 days, the shoot and root biomass of the olive plants was quantified, along with total organic matter, and carbon, nitrogen and polyphenol contents of the growth substrate and shoots, and the fine root nutritional status. Two‐phase pomace increased the total organic matter, total nitrogen and polyphenol contents of the growth substrate. It significantly altered biomass partitioning in the olive plantlets, with reduced shoot dry mass and leaf area, and new shoot formation. It also increased fine and total root dry mass for all two‐phase pomace levels except 40%. There were no significant differences in leaf pigment content and ΦPSII across the treatment levels. Therefore, application of untreated two‐phase pomace at more than 4% induces a severe imbalance in olive plantlet biomass partitioning, and shoot and root growth.     

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.     

Root growth and functioning under atmospheric CO2 enrichment

This paper examines the extent to which atmospheric CO₂ enrichment may influence growth of plant roots and function in terms of uptake of water and nutrients, and carbon allocation towards symbionts. It is concluded that changes in dry matter allocation greatly depend on the experimental conditions during the experiment, the growth phase of the plant, and its morphological characteristics. Under non-limiting conditions of water and nutrients for growth, dry matter partitioning to the root is not changed by CO₂ enrichment. The increase in root/shoot ratio, frequently observed under limiting conditions of water and/or nutrients, enables the plant to explore a greater soil volume, and hence acquire more water and nutrients. However, more data on changes in dry matter allocation within the root due to atmospheric CO₂ are needed. It is concluded that nitrogen fixation is favored by CO₂ enrichment since nodule mass is increased, concomitant with an increase in root length. The papers available so far on the influence of CO₂ enrichment on mycorrhizal functioning suggest that carbon allocation to the roots might be increased, but also here more experiments are needed.Abbreviations LAR leaf area ratio - LWR leaf weight ratio - SWR stem weight ratio - RGR relative growth rate - R/S root/shoot - RWR root weight ratio     

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     

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.     

A Transport-resistance Model of Forest Growth and Partitioning

The transport-resistance approach to dry-matter partitioningis used to construct a model of forest growth The model is atthe stand level for a monoculture of identical trees of thesame age There are five major organ compartments in the modelfoliage, branches, stem, coarse roots, and fine roots and mycorrhizasThe matter in each compartment is further subdivided into menstem,structure, carbon substrate, and nitrogen substrate The modelis driven by daily radiation including day length, ambient CO₂concentration, and daily means of air and soil temperature Thefine roots are provided with constant values of soil mineralnitrogen pools (ammonium and nitrate) from which uptake occursGrowth over about 100 years is simulated for various environmentalconditions and soil mineral nitrogen levels, thinning is alsosimulated Natural tree death occurs within the model Particularattention is paid to dry matter partitioning patterns, and tothe dry matter per stem when death occurs The model is robustand responsive, and provides a framework for further developmentand application to many ecological and environmental scenarios,as well as to some forest management problems Model, forest, growth, partitioning     

Effects of nitrogen forms on dry matter partitioning and nitrogen metabolism in two contrasting genotypes of Catasetum fimbriatum (Orchidaceae)

The effects of either organic (urea and glutamine) or inorganic nitrogen forms (nitrate and ammonium) on dry matter accumulation in shoots and roots and on nitrogen assimilatory enzyme activities were studied in two Catasetum fimbriatum genotypes. Both genotypes, which had inverse patterns of dry matter partitioning between shoots and roots, were aseptically incubated in gelled culture media containing 6 mol m⁻³ of nitrogen and incubated in growth chamber for 30 and 60 days. In vivo nitrate reductase, glutamine synthetase, glutamate dehydrogenase activities as well as free ammonium contents were determined in shoots and roots of plants grown in four different nitrogen sources. Nitrogen assimilatory enzyme activities showed the highest values in the genotype that accumulated dry matter predominantly in the shoots. The nitrogen sources supplied affected dry matter accumulation in shoots and roots of both C. fimbriatum genotypes; however, they were not enough to change the characteristic pattern of dry matter partitioning of each genotype. On the other hand, the differences in the root/shoot ratio found among nitrogen treatments were relatively higher in the genotype that directed dry matter mainly to roots than in the genotype that allocates biomass to shoots. Our results suggest that NADH-dependent glutamate dehydrogenase plays an important role in ammonium assimilation in C. fimbriatum plants, particularly in the root system. Nitrogen metabolism and the dry matter partitioning of the two genotypes are discussed.     

Modulation of carbon and nitrogen allocation in Urtica dioica and Plantago major by elevated CO2: Impact of accumulation of nonstructural carbohydrates and ontogenetic drift

Doubling the atmospheric CO² concentration from 350 to 700 μ1 1^?1 increased the relative growth rate (RGR) of hydroponically grown Urtica dioica L. and Plantago major ssp. pleiosperma Pilger only for the first 10–14 days. Previous experiments with P. major led to the conclusion that RGR did not respond in proportion to the rate of photosynthesis. The present paper is focussed on the analysis of the impact of changes in leaf morphology, dry matter partitioning, dry matter chemical composition and ontogenetic drift on this discrepancy. Soon after the start of the treatment, carbohydrate concentrations were higher at elevated CO²: a reaction that was largely due to starch accumulation. An increase in the percentage of leaf dry matter and decreases in the specific leaf area (SLA) and the shoot nitrogen concentration were correlated with an increase in the total nonstructural carbohydrate concentration (TNC). A combination of accumulation of soluble sugars and starch and ontogenetic drift explains the decrease in SLA at the elevated CO² level. A similar ontogenetic effect of elevated CO² was observed on the specific root length (SRL). Other variables such as shoot nitrogen concentration and percentage leaf dry matter were not affected by correction of data for TNC levels. The net diurnal fluctuation of the carbohydrate pool in P. major was equal for both CO² concentrations, indicating that the growth response to elevated CO² may be ruled by variables other than photosynthesis, as for instance sink strength. Elevated CO² did not greatly influence the partitioning of nitrogen between soluble and insoluble, reduced N and nitrate, nor the allocation of dry matter between leaf. stem and root. The finding that the root to shoot ratio (R/S) was not affected by elevated CO² implies that, in order to maintain a balanced activity between roots and shoot, no shift in partitioning of dry matter upon doubling of the atmospheric CO² concentration is required. Our data on R/S are in good agreement with the response of R/S to high CO² predicted by models based on such a theorem.     

油菜地上部干物质分配与产量形成模拟模型

利用油菜器官生长与发育进程及环境因子之间的定量关系,构建了基于分配指数的油菜地上部器官干物质分配动态模拟模型.各器官干物质分配指数随着生理发育时间而变化,基因型、播期、氮素及水分水平影响各器官干物质在地上部分配的大小.其中,氮素营养水平对绿色叶片干物质分配影响最大,氮素营养水平越高,绿色叶片分配指数越大;播期影响角果分配指数,晚播的角果分配指数高于早播.模型引入氮素营养指数、水分及播期影响因子来定量油菜各器官在实际生产条件下的分配强度,同时考虑了品种遗传特性的影响.通过不同品种氮肥处理试验建立模型,利用不同品种播期试验资料对模型进行了初步检验,表明模型具有较好的预测性和适用性.     

Partitioning and Utilization of Carbon and Nitrogen in Nodulated Roots and Nodules of Chickpea (Cicer arietinum) Grown at Two Moisture Levels

The partitioning and utilization of carbon (C) and nitrogen(N) in nodulated roots and nodules of chickpea (Cicer arietinumL.) was studied at two moisture levels at 10-d intervals 40–140d after sowing (DAS). More C was used in respiration and lessin growth of nodulated roots and nodules under water stresscompared to controls during all growth stages except at theearly vegetative stage. Similarly, less nitrogen was investedin dry matter of both nodules and nodulated roots under stress,except during the vegetative stage where more nitrogen was used.Calculated over the entire growth period, as much as 14 and20% of the total nitrogen and 3 and 4% of the total carbon fixedby the plant was lost to the rooting medium under controlledand stressed conditions, respectively. The efficiency of nitrogen fixation with respect to net C utilizationwas maximal during seed filling under both control and stressconditions However, the efficiency of nitrogen fixation wasalways greater under drier conditions. Carbon, chickpea (Cicer arietinum L.), nitrogen, nitrogen fixation, partitioning     

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     

Growth and Biomass Allocation, with Varying Nitrogen Availability, of Near-isogenic Pea Lines with Differing Foliage Structure

The single-gene mutation afila in pea (Pisum sativum L.) resultsin the replacement of proximal leaflets with branched tendrils,thereby reducing leaf area. This study investigated whethertheafila line could adjust biomass partitioning when exposedto varying nutrient regimes, to compensate for reduced leafarea, compared with wild-type plants. Wild-type and afila near-isogeniclines were grown in solution culture with nitrate-N added toinitially N-starved seedlings at relative addition rates (R_N)of 0.06, 0.12, 0.15 and 0.50 d^-1. The relative growth rate (R_W)of the whole plants closely matched R_Nat 0.06 and 0.12 d^-1,but higher R_Nresulted in a slightly higher growth rate. At agiven R_N, the wild-type line had lower plant nitrogen statusthan the afila line. R_Wof the roots of the afila line was lessthan R_Wof the roots of the wild-type at the three higher ratesof N supply despite a greater accumulation of N in the rootsof the afila plants. Consequently, plant nitrogen productivity(growth rate per unit nitrogen) was lower for afila. Dry matterallocation was strongly influenced by nitrogen status, but nodifferences in shoot–root dry matter allocation were foundbetween wild-type and afila with the same plant N status. Theseresults imply that decreased leaf area as a result of the single-genemutation afila affects dry matter allocation, but only accordingto its effect on the nitrogen status. Copyright 2000 Annalsof Botany Company Pisum sativum, pea, nitrogen limitation, growth, shoot–root allocation, relative growth rate, nitrogen productivity, isolines