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.     

Toward an allocation scheme for global terrestrial carbon models

The distribution of assimilated carbon among the plant parts has a profound effect on plant growth, and at a larger scale, on terrestrial biogeochemistry. Although important progress has been made in modelling photosynthesis, less effort has been spent on understanding the carbon allocation, especially at large spatial scales. Whereas several individual-level models of plant growth include an allocation scheme, most global terrestrial models still assume constant allocation of net primary production (NPP) among plant parts, without any environmental coupling. Here, we use the CASA biosphere model as a platform for exploring a new global allocation scheme that estimates allocation of photosynthesis products among leaves, stems, and roots depending on resource availability. The philosophy underlying the model is that allocation patterns result from evolved responses that adjust carbon investments to facilitate capture of the most limiting resources, i.e. light, water, and mineral nitrogen. In addition, we allow allocation of NPP to vary in response to changes in atmospheric CO₂. The relative magnitudes of changes in NPP and resource-use efficiency control the response of root:shoot allocation. For ambient CO₂, the model produces realistic changes in above-ground allocation along productivity gradients. In comparison to the CASA standard estimate using fixed allocation ratios, the new allocation scheme tends to favour root allocation, leading to a 10% lower global biomass. Elevated CO₂, which alters the balance between growth and available resources, generally leads to reduced water stress and consequently, decreased root:shoot ratio. The major exception is forest ecosystems, where increased nitrogen stress induces a larger root allocation.     

Physiological and growth responses of Centaurea maculosa (Asteraceae) to root herbivory under varying levels of interspecific plant competition and soil nitrogen availability

Summary Centaurea maculosa seedlings were grown in pots to study the effects of root herbivory by Agapeta zoegana L. (Lep.: Cochylidae) and Cyphocleonus achates Fahr. (Col.: Curculionidae), grass competition and nitrogen shortage (each present or absent), using a full factorial design. The aims of the study were to analyse the impact of root herbivory on plant growth, resource allocation and physiological processes, and to test if these plant responses to herbivory were influenced by plant competition and nitrogen availability. The two root herbivores differed markedly in their impact on plant growth. While feeding by the moth A. zoegana in the root cortex had no effect on shoot and root mass, feeding by the weevil C. achates in the central vascular tissue greatly reduced shoot mass, but not root mass, leading to a reduced shoot/root ratio. The absence of significant effects of the two herbivores on root biomass, despite considerable consumption, indicates that compensatory root growth occurred. Competition with grass affected plant growth more than herbivory and nutrient status, resulting in reduced shoot and root growth, and number of leaves. Nitrogen shortage did not affect plant growth directly but greatly influenced the compensatory capacity of Centaurea maculosa to root herbivory. Under high nitrogen conditions, shoot biomass of plants infested by the weevil was reduced by 30% compared with uninfested plants. However, under poor nitrogen conditions a 63% reduction was observed compared with corresponding controls. Root herbivory was the most important stress factor affecting plant physiology. Besides a relative increase in biomass allocation to the roots, infested plants also showed a significant increase in nitrogen concentration in the roots and a concomitant reduction in leaf nitrogen concentration, reflecting a redirection of the nitrogen to the stronger sink. The level of fructans was greatly reduced in the roots after herbivore feeding. This is thought to be a consequence of their mobilisation to support compensatory root growth. A preliminary model linking the effects of these root herbivores to the physiological processes of C. maculosa is presented.     

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。     

Seedling survival of Mediterranean shrubland species in relation to root:shoot ratio, seed size and water and nitrogen use

1. We hypothesized that in Mediterranean plant communities seedling survival of different species during the drought period would be related to their ability to use below-ground resources, particularly water and nitrogen.
2. For 5 years we studied under field conditions the summer seedling survival of 11 dominant species of a Spanish Mediterranean shrubland. We related seedling survival of the different species to their seed size and root allocation estimated as the slope of the function root biomass vs log shoot biomass. We used δ¹³C for the estimation of water-use efficiency and δ¹⁵N and nitrogen concentration to determine the sources of nitrogen utilized. We correlated these variables with root allocation.
3. Seedling survival of the different species was positively correlated with root allocation and seed size. Root allocation was also positively correlated with seed size. δ¹⁵N and nitrogen concentration were also positively correlated with root allocation, but δ¹³C was not.
4. Under the relatively moist conditions occurring during summer 1996, higher root allocation was associated with the use of nitrogen from more nitrogen saturated microsites.     

Seedling vigour and the early growth of transplanted rice (Oryza sativa)

Previous studies suggest that the positive response of transplanted rice (Oryza sativa L.) to nursery fertiliser application was due to increased seedling vigour or possibly to increased nutrient content. This paper presents results of two glasshouse experiments designed to test the hypothesis that seedling vigour was responsible for the response of transplanted seedlings to nursery treatments. The aim of the present study was to explore the concept of seedling vigour of transplanted rice and to determine what plant attributes conferred vigour on the seedlings. Seedling vigour treatments were established by subjecting seedlings to short-term submergence (0, 1 and 2 days/week) in one experiment and to leaf clipping or root pruning and water stress in another to determine their effect on plant growth after transplanting. Submerging seedlings increased plant height but depressed shoot and root dry matter and root:shoot ratio of the seedling at 28 days after sowing. After transplanting these seedlings, prior submergence depressed shoot dry matter at 40 days. Nursery nutrient application increased plant height, increased root and shoot dry matter, but generally decreased root:shoot ratio. Pruning up to 60% of the roots at transplanting decreased shoot and root dry matter, P concentration in leaves at panicle initiation (PI) and straw dry matter and grain yield at maturity. By contrast, pruning 30% of leaves depressed shoot and root dry matter by 30% at PI, and root dry matter and straw and grain yield by 20% at maturity. The combined effects of leaf clipping and root pruning on shoot, root and straw dry matter were largely additive. It is concluded that the response of rice yield to nursery treatments is largely due to increased seedling vigour and can be effected by a range of nutritional as well as non-nutritional treatments of seedlings that increase seedling dry matter, nutrient content, and nutrient concentration. Impairment of leaf growth and to a lesser extent root growth in the nursery depressed seedling vigour after transplanting. However, rather than increasing stress tolerance, seedling vigour was more beneficial when post transplant growth was not limited by nutrient or water stresses.     

EFFECTS OF ROOT PRUNING AT THE REGREENING STAGE ON DROUGHT TOLERANCE AND WATER USE EFFICIENCY OF WINTER WHEAT IN LATE GROWING STAGE

 通过盆栽试验研究了返青期根修剪对冬小麦(Triticum aestivum)后期耐旱性及水分利用效率的影响。在返青期设置了两个根修剪处理: 1)小剪根, 在植株一侧切去部分侧生根; 2)大剪根, 在主茎四周切去部分侧生根。不剪根者设为对照(CK)。研究结果显示, 两个根修剪处理均显著减少了小麦的根系, 但对根冠比没有显著影响。在花期, 两个根修剪处理的小麦旗叶的叶绿素荧光参数最大光化学效率(The maximum photochemical efficiency of PSⅡ, F_v/F_m)、 PSⅡ潜在活性 (PSⅡ potential activity, F_v/F_o)、实际光化学量子产量(Effective PSⅡ quantum yield, Φ_PSⅡ)、表观光合电子传递速率(Apparent rate of photosynthetic electron transport, ETR)、光化学淬灭系数( Coefficient of photochemical quenching, q_P)和非光化学淬灭系数(Coefficient of non-photochemical quenching, NPQ)值, 在停止供水7 d后, 均显著高于对照, 这表明根修剪小麦的耐旱性强于对照, 因此在干旱胁迫下有较高的光化学活性。小剪根处理在高水条件下对小麦产量无显著影响, 而在中度干旱条件下显著提高了小麦的产量, 因此, 小剪根处理显著提高了小麦的抗旱系数; 小剪根处理在高水分处理(土壤水分含量为田间持水量的85%)和中度干旱胁迫处理(土壤水分含量为田间持水量的55%)条件下, 均显著提高了小麦的水分利用效率。但大剪根处理由于严重影响了群体数量和产量, 水分利用效率和抗旱系数均没有提高。可见, 适当地减少根系有助于小麦的耐旱性和水分利用效率的提高。     

根修剪对黄土旱塬冬小麦(Triticum aestivum)根系分布、根系效率及产量形成的影响

通过田间试验研究了不同时期根修剪处理对冬小麦（Triticum aestivum）根系大小与分布、根系效率、水分利用效率及产量形成的影响。设置4个根修剪处理：越冬期小剪根（WS）、越冬期大剪根（WB）,返青期小剪根（GS）、返青期大剪根（GB）,未剪根小麦作为对照（CK）。结果表明,到花期时,各根修剪处理小麦的在0～120cm总根量均显著小于对照。与对照相比各根修剪处理主要是显著地减少了上层土壤中的根量。但WS和GS两小剪根处理和对照相比在中层土壤中有较大的根量;花后各处理小麦旗叶的气孔导度和蒸腾速率均显著大于对照。这说明根修剪处理减少了小麦表层的根量,从而削弱了表土干旱信号对作物与外界气体交换的抑制作用。花期时各根修剪小麦的净光合速率均显著高于对照,而单位面积上的根呼吸速率均显著小于对照,根修剪处理提高了小麦的根系效率,使更多的光合产物用于籽粒生产,从而提高了小麦的收获指数。根修剪还提高了小麦的水分利用效率,其中WS、WB、GS处理的水分利用效率显著高于对照。但是GB处理的水分利用效率却没有显著提高。因此,本研究进一步证明了由不同年代品种得到的推测,认为在旱地农业中,通过遗传育种或采用适当农艺措施优化根系分布,既可以减少生长前期作物对水分的过度消耗,又能够削弱花后表土过度干旱对作物生长抑制作用,同时降低根系对同化产物的消耗,对作物产量及水分利用效率的提高具有积极的作用。     

Genes controlling legume nodule numbers affect phenotypic plasticity responses to nitrogen in the presence and absence of rhizobia

We investigated the role of three autoregulation of nodulation (AON) genes in regulating of root and shoot phenotypes when responding to changing nitrogen availability in the model legume, Medicago truncatula. These genes, RDN1‐1 (ROOT DETERMINED NODULATION1‐1), SUNN (SUPER NUMERIC NODULES), and LSS (LIKE SUNN SUPERNODULAOR), act in a systemic signalling pathway that limits nodule numbers. This pathway is also influenced by nitrogen availability, but it is not well known if AON genes control root and shoot phenotypes other than nodule numbers in response to nitrogen. We conducted a controlled glasshouse experiment to compare root and shoot phenotypes of mutants and wild type plants treated with four nitrate concentrations. All AON mutants showed altered rhizobia‐independent phenotypes, including biomass allocation, lateral root length, lateral root density, and root length ratio. In response to nitrogen, uninoculated AON mutants were less plastic than the wild type in controlling root mass ratio, root length ratio, and lateral root length. This suggests that AON genes control nodulation‐independent root architecture phenotypes in response to nitrogen. The phenotypic differences between wild type and AON mutants were exacerbated by the presence of nodules, pointing to resource competition as an additional mechanism affecting root and shoot responses to nitrogen.     

Influence of nitrogen application rate on the magnitude of root and shoot growth flushes of Viburnum odoratissimum Ker-Gawl

Most trees and shrubs have cyclic, rather than continuous shoot and root growth. Growth cycles have important implications with regards to transplant and fertilization timing. Although growth cycles of several woody species have been characterized, no information is available on sweet viburnum (Viburnum odoratissimum). A series of experiments was conducted to study shoot and root growth flushes of sweet viburnum and the influence of nitrogen fertilization on flushes. Plants were grown in observation tubes and kept under greenhouse conditions. Sweet viburnum exhibited alternating periods of root and shoot elongation, and the root elongation peak preceded shoot elongation peak by 6–18 days. Increased nitrogen fertilization rate negatively impacted the magnitude and number of root growth flushes. Further research is needed to determine when maximum nitrogen uptake is occurring, relative to root and shoot growth cycles. Section Editor: P. J. Gregory     

Transplanting shock in white spruce; effect of cold-storage and root pruning on water relations and stomatal conditioning

White spruce [ Piceae glauca (Moench) Voss] seedlings were pre-conditioned by cold storage and root pruning and a study was made of the effects of conditioning on root regeneration and stomatal adaptation following transplanting.
Root pruning (0 to 75% of the initial root area) of dormant seedlings immediately prior to transplanting did not confer any detectable physiological advantage in either cold-stored or freshly-lifted seedlings. Transpiration rates and stomatal resistance were not greatly affected by root pruning in the 6 week period after planting and root pruning did not greatly alter the response to drought.
Cold-stored seedlings appear better able to avoid transplanting shock and an early drought despite delayed root growth as evidenced by a) a delay in flushing of new shoot growth and b) pre-conditioning of the stomatal apparatus which reduced water loss after planting and when moisture was limiting. Seedlings freshly-lifted in the spring did not show these adaptive responses and their stomata were poorly adapted to reduce water loss in the post-planting period after a drought.     

Nutrition and growth of coniferous seedlings at varied relative nitrogen addition rate

The growth of two provenances of Pinus sylvestris L. were compared with two provenances of Picea abies (L.) Karst. and with Pinus contorta Dougl. when grown in solution cultures with low nutrient concentrations. Nitrogen was added at different exponentially increasing rates, and the other nutrients were added at a rate high enough to ensure free access of them to the seedlings. During an initial period of the culture (a lag phase), when the internal nutrient status was changing from optimum to the level of the treatment, deficiency symptoms appeared. The needles yellowed and the root/shoot ratio increased. The initial phase was followed by a period of exponential growth and steady-state nutrition. The needles turned green again, and the root/shoot ratio stabilized at a level characteristic of the treatment. These patterns were the same as previously reported for other tree species. The relative growth rate during exponential growth was numerically closely equal to the relative nitrogen addition rate. The maximum relative growth rates were about 6 to 7.5% dry weight increase day^-1. This is a much lower maximum than for broad-leaved species (about 20 to 30% day^-1) under similar growth conditions. The internal nitrogen concentrations of the seedlings and the relative growth rates were stable during the exponential period. Close linear relationships were found between these parameters and the relative addition rate up to maximum growth. During steady state the relative growth rates of the different plant parts were equal. However, there were large differences between genotypes in absolute root growth rate at the same seedling size because of differences in root/shoot ratio. Lodgepole pine had the highest root growth rate, whereas that of Norway spruce, especially the southern provenance, was remarkably low. Yet, Norway spruce had a high ability to utilize available nutrients. In treatments with free nutrient access, growth allocation to the shoot had a high priority in all genotypes, but there was still a marked tendency for luxury uptake of nutrients. Nitrogen productivity (growth rate per unit of nitrogen) was lower than in broadleaved species and highest in lodgepole pine. The relevance of the dynamic factors, i.e. maximum relative growth rate, nutrient uptake rate, nitrogen productivity, growth allocation and root growth rate, are discussed with regard to conifer characteristics and selection value.     

Growth rate, IAA and cytokinin content of wheat seedling after root pruning

Root pruning of wheat seedlings resulted in 2–10 foldincrease in the concentration of IAA in roots ascompared to the control level, which might beresponsible for the observed initiation of lateralroot growth. Cytokinin concentration in xylem sap wasdecreased initially by 60% by pruning in accordancewith the reduction in the hormone-producing organ.Nevertheless cytokinin content in the shoots remainedhigh, which might be due to a decrease in cytokinindecay registered in vitro. A subsequent increasein the export of cytokinins from roots up to thecontrol level demonstrated an elevated ability of thepruned organ to synthesise the hormone. The highcytokinin content in the shoots correlated with theability of the plants to maintain their transpirationand growth at the level of intact plants. Both IAA andcytokinins seem to be important in the restoration ofthe shoot/root balance disturbed by root pruning.     

Oryza sativa Lysine‐Histidine‐type Transporter 1 functions in root uptake and root‐to‐shoot allocation of amino acids in rice

In agricultural soils, amino acids can represent vital nitrogen (N) sources for crop growth and yield. However, the molecular mechanisms underlying amino acid uptake and allocation are poorly understood in crop plants. This study shows that rice (Oryza sativa L.) roots can acquire aspartate at soil concentration, and that japonica subspecies take up this acidic amino acid 1.5‐fold more efficiently than indica subspecies. Genetic association analyses with 68 representative japonica or indica germplasms identified rice Lysine‐Histidine‐type Transporter 1 (OsLHT1) as a candidate gene associated with the aspartate uptake trait. When expressed in yeast, OsLHT1 supported cell growth on a broad spectrum of amino acids, and effectively transported aspartate, asparagine and glutamate. OsLHT1 is localized throughout the rice root, including root hairs, epidermis, cortex and stele, and to the leaf vasculature. Knockout of OsLHT1 in japonica resulted in reduced root uptake of amino acids. Furthermore, in ¹⁵N‐amino acid‐fed mutants versus wild‐type, a higher percentage of ¹⁵N remained in roots instead of being allocated to the shoot. ¹⁵N‐ammonium uptake and subsequently the delivery of root‐synthesized amino acids to Oslht1 shoots were also significantly decreased, which was accompanied by reduced shoot growth. These results together provide evidence that OsLHT1 functions in both root uptake and root to shoot allocation of a broad spectrum of amino acids in rice.     

Signaling mechanisms integrating root and shoot responses to changes in the nitrogen supply

During their life cycle, plants must be able to adapt to wide variations in the supply of soil nitrogen (N). Changes in N availability, and in the relative concentrations of NO₃ ⁻and NH₄ ⁺, are known to have profound regulatory effects on the N uptake systems in the root, on C and N metabolism throughout the plant, and on root and shoot morphology. Optimising the plant’s responses to fluctuations in the N supply requires co-ordination of the pathways of C and N assimilation, as well as establishment of the appropriate allocation of resources between root and shoot growth. Achieving this integration of responses at the whole plant level implies long-distance signaling mechanisms that can communicate information about the current availability of N from root-to-shoot, and information about the C/N status of the shoot in the reverse direction. In this review we will discuss recent advances which have contributed to our understanding of these long-range signaling pathways.     

A Model Allocating Growth Among Leaf Proteins, Shoot Structure, and Root Biomass to Produce Balanced Activity

A model is developed that considers the allocation of carbonand nitrogen substrates to a protein compartment in the shoots,shoot structural components, and root biomass. Inclusion ofa shoot-protein compartment allows variation in shoot-specificactivity to be modelled as a function of leaf nitrogen concentration.Allocation to the biomass compartments is controlled by twopartitioning variables that are defined by explicitly usingthe balanced activity hypothesis. The model produces balancedactivity where the shoot-specific activity, as well as rootand shoot biomass, vary in response to the above-ground (lightand CO₂) and below-ground (nitrogen) environments. The predictedpatterns of both root: shoot ratio and leaf nitrogen concentrationin response to environmental resource availability are qualitativelyconsistent with general trends observed in plants. Biomass allocation, plant growth, modelling, leaf nitrogen, root: shoot ratio, balanced activity     

Size and morphology of root systems of perennial grasses from contrasting habitats as affected by nitrogen supply

This paper discusses interspecific differences and phenotypic responses to nitrogen supply in various root parameters of five perennial grasses from contrasting habitats. The following root parameters were studied: root:shoot ratio, specific root length, specific root area, mean root diameter, frequency of fine roots, and the length and density of root hairs. Significant between-species variation was found in all of these features. Species from fertile sites had higher root:shoot ratios at high nitrogen supply than species from infertile habitats. All species growing at low nitrogen supply showed a significant increase in root:shoot ratio. Specific root length, specific root area, mean root diameter and frequency of fine roots were not affected significantly by nitrogen supply. Species from infertile sites responded to low nitrogen supply by a significant increase in root hair length and root hair density.     

施氮对三个南方珍稀树种幼苗生长和生物量分配的影响

为了探讨珍稀树种对短期氮素添加的响应,该文研究了氮素添加(0、0.1、0.2、0.4和0.6g·kg~(-1)土)对观光木、棱角山矾和半枫荷幼苗生长和生物量分配的影响。结果表明:3个树种幼苗对外源氮素添加的反应不同,施氮显著促进观光木幼苗株高、基径、冠幅以及全株生物量和各部分生物量的增加,中低氮促进半枫荷幼苗的生长,但高氮抑制其生长;少量施氮对棱角山矾幼苗的形态和生物量参数没有产生显著影响,中量施氮抑制其生长。氮素营养的改变显著影响3种植物幼苗的生物量分配,观光木幼苗的根生物量比和根冠比均随施氮量的增加而显著降低;除高氮处理外,半枫荷幼苗的根生物量比和根冠比均随供氮量的增加而显著升高;棱角山矾的根生物量比和根冠比均随供氮量的增加而显著升高,可能与施氮抑制其茎叶的生长有关。总的来看,观光木幼苗更能耐受高氮条件,半枫荷幼苗次之,而棱角山矾幼苗不耐高氮;但到当年生长季末,各氮处理半枫荷幼苗的株高、基径和总相对生长速率均显著大于其它两种植物。     

The Effect of Nitrogen Supply to Urtica dioica L. Plants on the Distribution of Assimilate Between Shoot and Roots

In this study the influence of nitrogen nutrition on the patterns of carbon distribution was investigated with Urtica dioica. The nettles were grown in sand culture at 3 levels of NO^?₃, namely 3 (low), 15 (medium) and 22 (high) mM. These levels encompassed a range within which nitrogen did not affect total biomass production. The ratio of root: shoot biomass of the low nitrogen plants was, however, significantly higher than that of the nettles grown at medium and high N supply. Carbon allocation from one leaf of each pair of leaves was examined after a ¹⁴CO₂-pulse and a subsequent ¹⁴C distribution period of one night. Only the youngest two leaf pairs did not export assimilates. Carbon (¹⁴C) export to the shoot apex and to the roots, as measured at the individual nodes responded to the nitrogen status: At medium and high nitrogen supply the 3rd, 4th and 5th leaf pairs exported to the shoot apex, while lower leaves exported to the root. At low nitrogen supply only the 3rd leaf exported towards the shoot apex. The results illustrate the plastic response of carbon distribution patterns to the nitrogen supply, even when net photosynthesis, carbon export from the source leaves and biomass production were not affected by the nitrogen supply to the plant.     

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