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.     

Root: shoot ratio as a balance between nitrogen productivity and photosynthesis

Abstract It is shown how root : shoot ratios can be derived from two plant characteristics, (i) the nitrogen productivity (amount of biomass produced per unit of nitrogen in the biomass and per unit of time) and (ii) the relationship between photo-synthetic rate and internal nitrogen concentration. Derived root: shoot ratios are compared with experimental data for seedlings of seven tree species grown with stable internal nitrogen concentrations. Agreement is good but more exact knowledge of the relationship between nutrient status and photosynthesis is needed. It is shown that a formula derived for the root: shoot ratio can also be applied under non-steady-state conditions.     

Combined effects of elevated CO2 and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia

Impacts of either elevated CO₂ or drought stress on plant growth have been studied extensively, but interactive effects of these on plant carbon and nitrogen allocation is inadequately understood yet. In this study the response of the dominant desert shrub, Caragana intermedia Kuanget H.c.Fu, to the interaction of elevated CO₂ (700 ± 20 μmol mol⁻¹) and soil drought were determined in two large environmental growth chambers (18 m²). Elevated CO₂ increased the allocation of biomass and carbon into roots and the ratio of carbon to nitrogen (C:N) as well as the leaf soluble sugar content, but decreased the allocation of biomass and carbon into leaves, leaf nitrogen and leaf soluble protein concentrations. Elevated CO₂ significantly decreased the partitioning of nitrogen into leaves, but increased that into roots, especially under soil drought. Elevated CO₂ significantly decreased the carbon isotope discrimination (Δ) in leaves, but increased them in roots, and the ratio of Δ values between root and leaf, indicating an increased allocation into below-ground parts. It is concluded that stimulation of plant growth by CO₂ enrichment may be negated under soil drought, and under the future environment, elevated CO₂ may partially offset the negative effects of enhanced drought by regulating the partitioning of carbon and nitrogen.     

Nitrogen dynamics in the intact grasses Poa trivialis and Panicum maximum receiving contrasting supplies of nitrogen

The C(3) grass Poa trivialis and the C(4) grass Panicum maximum were grown in sand culture and received a complete nutrient solution with nitrogen supplied as 1.5 mol m(-3) NH(4)NO(3). (15)N tracer techniques were used to quantify the relative use of root uptake and mobilization in supplying nitrogen to growing leaves in intact plants which either continued to receive nitrogen or which received the complete nutrient solution without nitrogen. The allocation of both (15)N-labelled nitrogen uptake and unlabelled mobilized nitrogen indicated that, under their conditions of growth, the sink strength of growing leaves was relatively greater in P. maximum than P. trivialis. The supply of nitrogen by mobilization to side tillers of P. trivialis was completely stopped as the external nitrogen supply was reduced, whilst in P. maximum some allocation of mobilized nitrogen to side tillers, roots and growing leaves was maintained. In both plant species receiving an uninterrupted supply of nitrogen the allocation pattern of mobilized nitrogen differed from that of nitrogen derived from root uptake. Differences exist in the degree to which P. trivialis and P. maximum utilized uptake and mobilization to supply nitrogen to the growing leaves. In P. trivialis roots were always a net sink of mobilized nitrogen, irrespective of the external nitrogen supply. In P. maximum, roots were a net sink of mobilized nitrogen when external nitrogen was withdrawn, but exhibited both source and sink behaviour when nitrogen supply was continued.     

Resource capture, biomass allocation and growth in herbaceous plants

Plant species differ widely in their rate of biomass production, even when grown under optimal conditions. A key question concerns the extent to which these growth rates correlate with the uptake of carbon and nitrogen and with the biomass allocation between leaves and roots. Recent data show that the answer to this question differs for mono- and dicotyledons, and that more than biomass allocation, it is the ratio between the activities of leaves and roots that correlates with the growth rate of a plant.     

Allocation strategies of carbon,nitrogen and phosphorus following a gradient of wildfire severities

植物碳、氮、磷在不同火烧强度下的分配策略 森林野火是影响北方针叶林演替过程中养分分配规律的重要因素。然而,植物叶片和细根之间 的碳(C)、氮(N)、磷(P)分配策略在不同强度森林野火后的研究尚不充分。本研究旨在探讨不同野火强度下叶片和细根间C、N、P的分配策略。运用化学计量学理论和异速生长方程,选取中国东北大兴安岭地区的4个不同火烧强度(未火烧、低、中、高)恢复10年后的火烧迹地为研究样地,比较不同火烧强 度下各物种叶片和细根的C、N、P含量。研究结果表明,与未受到火烧的样地相比,轻度火烧迹地的植物叶片和细根C浓度增加,重度火烧迹地植物叶片N浓度最高,但是细根N浓度最低。N:P比值的平均值大于16的结果表示植物养分利用策略在高火烧强度下趋于P限制。更重要的是,随着火烧严重程度的增加,细根与叶片间的C、N、P分配规律出现由异速生长向等速生长的转变,即随着火烧强度的增加,元素分配表现为对叶片的分配多于细根。这些结果表明,植物叶片和细根之间的元素分配策略在受到不同强度的野火干扰以后发生了失衡。本研究加深了我们对火后森林生态系统演替过程中植物与土壤养分动态的认识。     

Increased forest carbon storage with increased atmospheric CO2 despite nitrogen limitation: a game‐theoretic allocation model for trees in competition for nitrogen and light

Changes in resource availability often cause competitively driven changes in tree allocation to foliage, wood, and fine roots, either via plastic changes within individuals or through turnover of individuals with differing strategies. Here, we investigate how optimally competitive tree allocation should change in response to elevated atmospheric CO₂ along a gradient of nitrogen and light availability, together with how those changes should affect carbon storage in living biomass. We present a physiologically‐based forest model that includes the primary functions of wood and nitrogen. From a tree's perspective, wood is an offensive and defensive weapon used against neighbors in competition for light. From a biogeochemical perspective, wood is the primary living reservoir of stored carbon. Nitrogen constitutes a tree's photosynthetic machinery and the support systems for that machinery, and its limited availability thus reduces a tree's ability to fix carbon. This model has been previously successful in predicting allocation to foliage, wood, and fine roots along natural productivity gradients. Using game theory, we solve the model for competitively optimal foliage, wood, and fine root allocation strategies for trees in competition for nitrogen and light as a function of CO₂ and nitrogen mineralization rate. Instead of down‐regulating under nitrogen limitation, carbon storage under elevated CO₂ relative to carbon storage at ambient CO₂ is approximately independent of the nitrogen mineralization rate. This surprising prediction is a consequence of both increased competition for nitrogen driving increased fine root biomass and increased competition for light driving increased allocation to wood under elevated CO₂.     

不同供氮水平对水曲柳苗木生物量、氮分配及其季节变化的影响

在温室内以水曲柳苗木为材料进行砂培试验,探讨了4种不同氮素浓度处理(1、4、8和16mmol·L^-1)下的水曲柳苗木根系和叶片内氮分配以及对苗木生物量的影响．结果表明,氮素供给浓度显著影响苗木根系氮浓度和叶片氮浓度．随着供氮水平提高,苗木体内根系和叶片氮浓度明显提高．在生长初期(6月)和中期(7、8月),叶片中氮浓度分别高于根系9．40、9．55和4．21mg·g^-1,而在生长末期(9月)叶片中氮浓度低于根系;随着水曲柳幼苗生长发展,体内氮贮量呈明显上升趋势．9月份全株氮贮量比6月份平均增加了4倍．不同氮处理下水曲柳体内氮贮量明显不同．高氮处理下氮贮量平均为N1处理下4倍．氮贮量分配在不同部位有很大不同．分配到叶片中的比例在6月份最高,平均为43％．分配到根系中氮贮量比例随生长而增加,9月份根系氮贮量相对值最高,为81％．如果不考虑氮浓度和季节的作用,根系中分配的氮最多,其次为叶片,茎中最少．     

Resource allocation in different parts of juvenile mountain birch plants: effect of nitrogen supply on seedling phenolics and growth

The composition and concentrations of phenolic compounds were studied in the first true leaves, cotyledons, stems and roots of 2.5-week-old seedlings of mountain birch ( Betula pubescens ssp. czerepanovii ). The differences in secondary compounds among these plant parts were both qualitative and quantitative. In all parts, condensed tannins accounted for more than 50% of the phenolics. In the first true leaves and cotyledons, chlorogenic acid was the most abundant of the HPLC phenolics. The main components in stems were (+)-catechins and rhododendrins whereas in roots, the main components were ellagitannins. The seedlings were grown at three levels of nitrogen supply (very low-N, low-N, moderate-N), and the effect of nitrogen on concentrations of phenolic compounds was studied in all plant parts. The dry weight of all plant parts, except the roots, increased with increased nitrogen. In all parts, the concentration of condensed tannins was higher at lower levels of nitrogen than at moderate-N. The concentrations of total HPLC phenolics and also those of the compound groups of HPLC phenolics were, however, affected only in the first true leaves and roots. The concentrations in the first true leaves were generally higher in seedlings grown at very low-N and low-N than in seedlings grown at moderate-N. The concentrations in roots were highest at low-N. Not all compounds responded to nitrogen supply in the same manner. The changes in concentrations cannot be exclusively interpreted as changes in the accumulation of phenolic compounds, due to dilution caused by the increase in biomass in better nitrogen availability. There were differences in carbon allocation between condensed tannins and HPLC phenolics in seedlings grown at different nitrogen levels.     

Robust biological nitrogen fixation in a model grass–bacterial association

Nitrogen‐fixing rhizobacteria can promote plant growth; however, it is controversial whether biological nitrogen fixation (BNF) from associative interaction contributes to growth promotion. The roots of Setaria viridis, a model C₄ grass, were effectively colonized by bacterial inoculants resulting in a significant enhancement of growth. Nitrogen‐13 tracer studies provided direct evidence for tracer uptake by the host plant and incorporation into protein. Indeed, plants showed robust growth under nitrogen‐limiting conditions when inoculated with an ammonium‐excreting strain of Azospirillum brasilense. ¹¹C‐labeling experiments showed that patterns in central carbon metabolism and resource allocation exhibited by nitrogen‐starved plants were largely reversed by bacterial inoculation, such that they resembled plants grown under nitrogen‐sufficient conditions. Adoption of S. viridis as a model should promote research into the mechanisms of associative nitrogen fixation with the ultimate goal of greater adoption of BNF for sustainable crop production.     

不同管理模式对茶树碳氮磷含量及其生态化学计量比的影响

植物不同器官的碳(C)、氮(N)、磷(P)含量及其生态化学计量特征能够反映植物内部的养分分配与平衡关系。该研究以福建安溪3种不同管理模式的铁观音茶园为研究对象, 设置了常规管理模式下的茶园(M1)、间作套种模式下的茶园(M2)和现代技术管理模式下的茶园(M3) 3种样地, 分析茶树根、茎、叶器官的C、N、P含量及其化学计量学特征, 养分的变异特征与异速生长关系。结果表明: M2和M3管理模式下茶树根、茎、叶N、P含量均显著高于M1管理模式, C含量差异不明显; 茶树根、茎、叶C:N、C:P、N:P均表现为M1 > M2 > M3。茶树不同器官C、N、P含量差异较大, 根据变异来源分析, 管理模式因素对C、N、P含量变异的影响均达到显著水平。根茎叶N-P的异速生长关系表明茶树不同器官的养分需求存在相似性; 土壤pH和容重是影响C:N、C:P、N:P的重要因素, 而土壤含水量和盐度对茶树根和叶C含量的影响较大。总体来讲, 间作套种以及现代化滴灌、水肥等管理模式可以改善茶树对养分的吸收效率, 对解决土壤养分不均衡问题具有正面效应。     

Carbon and nitrogen content of congeneric annual and perennial grass species: relationships with growth

Seven annual-perennial pairs of grass species (six congeneric and one pair taken at random) were grown under productive conditions in the laboratory in order to investigate which plant characters were responsible for the higher relative growth rate (RGR) of annuals as compared to perennials under these conditions. The nitrogen and carbon concentrations of shoot organs and of the whole plant were higher in annuals than in perennials. This was also the case for the specific absorption rate for nitrate and nitrogen productivity (on whole plant and leaf basis). The range of RGR displayed by the 14 species was large enough (0.15–0.33d⁻¹) to examine the general relationships between RGR and the various parameters measured in the present study. RGR was positively related to plant, leaf blade and sheath nitrogen concentrations, but there was no relationship between RGR and any of the carbon concentrations. RGR also strongly correlated with specific absorption rate for nitrate and with nitrogen productivity. A new factorization of this latter parameter led to the definition of the ‘leaf nitrogen productivity’ (N_LP), which is likely to depend on photosynthetic nitrogen use efficiency. RGR was shown to be strongly correlated with N_LP, but not with the second term of the factorization, namely the proportion of plant nitrogen allocated to the leaves.     

Changes in the acquisition and partitioning of carbon and nitrogen in the gibberellin-deficient mutants A70 and W335 of tomato (Solanum lycopersicum L.)

Even though the growth‐promoting effects of gibberellins (GAs) in plants are well established, little is known about GA action on carbon metabolism and the available reports seem contradictory. We studied the effects of GA deficiency in mutants of tomato (Solanum lycopersicum L.) on rates of carbon acquisition and the allocation of acquired carbon to growth and respiration of leaves, stems and roots. Carbon budgets were calculated from 24 h measurements of photosynthesis and respiration. The partitioning of nitrogen compounds to leaves, stems and roots, which strongly influences carbon budgets, was also studied. The GA‐deficient mutants acquired less carbon per unit plant mass per day than did the wild type and used a larger fraction of it for root growth and root respiration. To find out to what extent these changes were just consequences of restriction of growth, the experiment was repeated at a low exponential nitrate addition rate, which forced all genotypes to grow at the same rate. Under these conditions, the low‐GA mutants still photosynthesized and respired faster and partitioned more carbon to root growth than the wild type did. The reasons for the observed differences in carbon economies between the wild type and the low‐GA mutants are discussed.     

不同施氮量对乌龙茶生长和生理的影响

以武夷肉桂为研究对象,研究不同施氮量对乌龙茶幼龄茶树生长和生理的影响。结果表明,幼龄茶树对氮肥的需求不强烈,其新梢生物量、根生物量和总生物量以及茶叶产量随施氮量的增加而下降。茶树新梢全氮、叶绿素、游离氨基酸、茶多酚和咖啡碱的含量随施氮量的增加而增加,而茶树碳氮比随着施氮量增加而下降;但施氮并没有影响茶树总碳含量。老叶叶绿素含量、根全氮和硝态氮含量、新梢总糖含量与施氮量呈二次曲线回归关系,适度施氮促进根对氮的吸收、老叶叶绿素合成和新梢总糖代谢,过度施氮则相反。新梢生物量与其硝态氮含量和游离氨基酸总量显著负相关;根生物量与根碳氮比和新梢咖啡碱含量显著负相关;茎叶生物量和总生物量与根含氮量显著正相关,但与新梢硝态氮和氨基酸含量显著负相关。过度施氮造成茶树生产力下降的主要原因是因为过度施氮极显著提高了茶树氨基酸代谢水平,使用于茶树生长的碳代谢产物（如总糖）减少,进而影响茶树的生长。     

Flooding affects uptake and distribution of carbon and nitrogen in citrus seedlings

Soil flooding has been widely reported to affect large areas of the world. In this work, we investigated the effect of waterlogging on citrus carbon and nitrogen pools and partitioning. Influence on their uptake and translocation was also studied through 1?N and 13C labeling to provide insight into the physiological mechanisms underlying the responses. The data indicated that flooding severely reduced photosynthetic activity and affected growth and biomass partitioning. Total nitrogen content and concentration in the plant also progressively decreased throughout the course of the experiment. After 36 days of treatment, nitrogen content of flooded plants had decreased more than 2.3-fold compared to control seedlings, and reductions in nitrogen concentration ranged from 21 to 55% (in roots and leaves, respectively). Specific absorption rate and transport were also affected, leading to important changes in the distribution of this element inside the plant. Additionally, experiments involving labeled nitrogen revealed that 1?N uptake rate and accumulation were drastically decreased at the end of the experiment (93% and 54%, respectively). 13CO? assimilation into the plant was strongly reduced by flooding, with δ13C reductions ranging from 22 to 37% in leaves and roots, respectively. After 36 days, the relative distribution of absorbed 13C was also altered. Thus, 13C recovery in flooded leaves increased compared to controls, whereas roots exhibited the opposite pattern. Interestingly, when carbohydrate partitioning was examined, the data revealed that sucrose concentration was augmented significantly in roots (37-56%), whereas starch was reduced. In leaves, a marked increase in sucrose was detected from the first sampling onwards (36-66%), and the same patter was observed for starch. Taken together, these results indicate that flooding altered carbon and nitrogen pools and partitioning in citrus. On one hand, reduced nitrogen concentration appears to be a consequence of impaired uptake and transport. On the other hand, the observed changes in carbohydrate distribution suggest that translocation from leaves to roots was reduced, leading to significant starch accumulation in leaves and further decreases in roots.     

植物叶片氮分配及其影响因子研究进展

氮是植物生长的基本限制性因子,它的有效利用可以增加植物的适应性。叶片氮分配是指氮在植物叶片细胞各细胞结构以及游离化合物中所分配的比例。叶片氮的分配方式决定了叶片光合作用的强弱,影响叶片的坚韧程度以及化学防御强度,因此研究氮在植物叶片内的分配方式具有重要意义。阐述了叶片氮分配的方式,分析了影响叶片氮分配的生物和非生物因子(CO2,光,土壤养分),介绍了常用的叶片氮分配的研究方法,并对未来的研究进行了展望。     

云杉碳氮化学计量比对土壤水分和氮有效性的响应

以西南亚高山针叶林优势种——粗枝云杉(Picea asperata)为研究对象,探究不同土壤水分状况和氮添加下云杉碳氮化学计量比的变化及其响应过程。采用两因素(水分×氮素)随机区组实验,设置5个土壤水分梯度和3个氮添加浓度,其中土壤水分梯度分别是土壤田间持水量的40%(W1)、50%(W2)、60%(W3)、80%(W4)和100%(W5),氮添加浓度分别为0(N0)、20(N1)、40(N2)gNm^-2 a^-1。结果表明:(1)土壤水分和氮添加显著影响了云杉碳氮化学计量比(P<0.05),具体为:云杉植株和器官碳氮比在N0W4处理下最大值,随土壤水分有效性的降低而减小,随氮添加浓度的增加而降低。(2)随土壤水分有效性的降低,根和叶的碳含量显著升高(P<0.05),茎和叶的碳含量随着氮添加浓度的增加而降低。此外,土壤水分有效性的降低显著提高了根和茎的氮含量(P<0.05),各器官的氮含量随着氮添加浓度的增加而增加。在相同水分和氮添加浓度处理下表现为碳含量:叶>茎>根,氮含量:叶>根>茎。(3)云杉净光...     

土壤碳氮比对平邑甜茶幼苗生长和碳氮分配的影响

为探讨土壤碳氮比(C:N)对苹果(Malus pumila)植株生长和碳氮分配特性的影响, 采用碳氮双标记示踪技术, 以二年生平邑甜茶(Malus hupehensis)幼苗为试验材料, 研究了6个不同土壤C:N处理(T1-T6分别为4.70、9.78、14.70、19.96、25.60和28.83)下平邑甜茶的生长状况和氮素吸收、利用分配以及碳水化合物的运转特性。结果表明, 随着土壤C:N的逐渐增大, 平邑甜茶幼苗根系干重逐渐增加, 而株高、茎粗、地上部干重和植株总干重呈先增加后降低的趋势, 以T4处理最大。土壤C:N显著影响了平邑甜茶幼苗的 ¹⁵N利用率, 从T1到T4处理, 植株的 ¹⁵N利用率逐渐升高, T4处理(18.46%)是T1处理(10.65%)的1.73倍; 随着土壤C:N的进一步增加, 植株的 ¹⁵N利用率逐渐降低, T5和T6处理分别比T4处理降低了1.59%和2.58%。土壤C:N较低的T1和T2处理, 平邑甜茶幼苗各器官从肥料中吸收分配到的 ¹⁵N量对该器官全氮量的贡献率(Ndff)大小顺序为根>叶>茎, 随着土壤C:N的进一步增大, 叶片的Ndff均为最大, 其次是根, 茎最少。随着土壤C:N的增大, 叶片 ¹⁵N分配率逐渐升高, ¹³C分配率逐渐降低; 而根系 ¹⁵N分配率逐渐降低, ¹³C分配率逐渐升高。综合考虑植株生长和氮素利用状况, 本试验条件下适宜平邑甜茶生长的土壤C:N为21-23。     

Glutamate synthase activities and protein changes in relation to nitrogen nutrition in barley: the dependence on different plastidic glucose-6P dehydrogenase isoforms

In barley (Hordeum vulgare L. var. Nure), glutamate synthesis and the production of reducing power by the oxidative pentose phosphate pathway (OPPP) are strictly correlated biochemical processes. NADH-GOGAT was the major root isoform, whose activity increased on a medium supplied with NH4+ or NO3-; by contrast, no noticeable variations could be observed in the leaves of plants supplied with nitrogen. In the leaves, the major isoform is Fd-GOGAT, whose activity increased under nitrogen feeding. G6PDH activity increased in the roots supplied with nitrogen; no variations were observed in the leaves. Moreover, an increase of the P2 isoform in the roots was measured, giving 13.6% G6PDH activity localized in the plastids under ammonium, and 25.2% under nitrate feeding conditions. Western blots confirmed that P2-G6PDH protein was induced in the roots by nitrogen. P1-G6PDH protein was absent in the roots and increased in the leaves by nitrogen supply to the plants. The changes measured in cytosolic G6PDH seem correlated to more general cell growth processes, and do not appear to be directly involved in glutamate synthesis. The effects of light on Fd-GOGAT is discussed, together with the possibility for P2-G6PDH to sustain nitrogen assimilation upon illumination.