Nitrogen fixation and the mass balances of carbon and nitrogen in ecosystems

Ecosystems with high rates of nitrogen fixation often have high loss rates through leaching or possibly denitrification. However, there is no formal theoretical context to examine why this should be the case nor of how nitrogen accumulates in such open systems. Here, we propose a simple model coupling nitrogen inputs and losses to carbon inputs and losses. The nitrogen balance of this model system depends on plant (nitrogen fixer) growth rate, its carrying capacity, N fixed/C fixed, residence time of nitrogen and carbon in biomass, litter decay rate, litter N/C, and fractional loss rate of mineralized nitrogen. The model predicts the requirements for equilibrium in a nitrogen-fixing system, and the conditions on nitrogen fixation and losses in order for the system to accumulate nitrogen and carbon. In particular, the accumulation of nitrogen and carbon in a nitrogen-fixing system depend on an interaction between residence time in vegetation and litter decay rate in soil. To reflect a possible increased uptake of soil nitrogen and decreased respiratory cost of symbiotic nitrogen fixers, the model was then modified so that fixation rate decreased and growth rate increased as nitrogen capital accumulated. These modifications had only small effects on carbon and nitrogen accumulation. This suggests that switching from uptake of atmospheric nitrogen to mineral soil nitrogen as nitrogen capital accumulates simply results in a trade-off between energetic limitations and soil nitrogen limitations to carbon and nitrogen accumulation. Experimental tests of the model are suggested.     

Nitrogen use efficiency. 1. Uptake of nitrogen from the soil

The nitrogen use efficiency (NUE) of crop plants can be expressed very simply as the yield of nitrogen per unit of available nitrogen in the soil. This NUE can be divided into two processes: uptake efficiency, the ability of the plant to remove N from the soil normally present as nitrate or ammonium ions, and the utilisation efficiency, the ability of the plant to transfer the N to the grain, predominantly present as protein. In this article, we have highlighted the latest developments in the isolation and characterisation of the genes involved in the uptake of nitrogen from the soil.     

Plant root distributions and nitrogen uptake predicted by a hypothesis of optimal root foraging

CO(2)-enrichment experiments consistently show that rooting depth increases when trees are grown at elevated CO(2) (eCO(2)), leading in some experiments to increased capture of available soil nitrogen (N) from deeper soil. However, the link between N uptake and root distributions remains poorly represented in forest ecosystem and global land-surface models. Here, this link is modeled and analyzed using a new optimization hypothesis (MaxNup) for root foraging in relation to the spatial variability of soil N, according to which a given total root mass is distributed vertically in order to maximize annual N uptake. MaxNup leads to analytical predictions for the optimal vertical profile of root biomass, maximum rooting depth, and N-uptake fraction (i.e., the proportion of plant-available soil N taken up annually by roots). We use these predictions to gain new insight into the behavior of the N-uptake fraction in trees growing at the Oak Ridge National Laboratory free-air CO(2)-enrichment experiment. We also compare MaxNup with empirical equations previously fitted to root-distribution data from all the world's plant biomes, and find that the empirical equations underestimate the capacity of root systems to take up N.     

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.     

Role of Arbuscular Mycorrhizal Fungi in Uptake of Phosphorus and Nitrogen From Soil

Abstract

Colonization of plant roots by arbuscular mycorrhizal fungi can greatly increase the plant uptake of phosphorus and nitrogen. The most prominent contribution of arbuscular mycorrhizal fungi to plant growth is due to uptake of nutrients by extraradical mycorrhizal hyphae. Quantification of hyphal nutrient uptake has become possible by the use of soil boxes with separated growing zones for roots and hyphae. Many (but not all) tested fungal isolates increased phosphorus and nitrogen uptake of the plant by absorbing phosphate, ammonium, and nitrate from soil. However, compared with the nutrient demand of the plant for growth, the contribution of arbuscular mycorrhizal fungi to plant phosphorus uptake is usually much larger than the contribution to plant nitrogen uptake. The utilization of soil nutrients may depend more on efficient uptake of phosphate, nitrate, and ammonium from the soil solution even at low supply concentrations than on mobilization processes in the hyphosphere. In contrast to ectomycorrhizal fungi, nonsoluble nutrient sources in soil are used only to a limited extent by hyphae of arbuscular mycorrhizal fungi. Side effects of mycorrhizal colonization on, for example, plant health or root activity may also influence plant nutrient uptake.     

Nitrogen Use Efficiency of Crop Plants: Physiological Constraints upon Nitrogen Absorption

Current global nitrogen fertilizer use has reached approximately one hundred billion kg per annum. In many agricultural systems, a very substantial portion of this applied nitrogen fertilizer is lost from soil to groundwaters, rivers and oceans. While soil physicochemical properties play a significant part in these losses, there are several characteristic features of plant nitrogen transporter function that facilitate N losses. Nitrate and ammonium efflux from roots result in a reduction of net nitrogen uptake. As external nitrate and ammonium concentrations, respectively, are increased, particularly into the range of concentrations that are typical of agricultural soils, elevated rates of nitrate and ammonium efflux result. The rapid down-regulation of high-affinity influx as plants become nitrogen replete further reduces the root's capacity to acquire external nitrogen; only nitrogen-starved roots absorb with both high capacity and high affinity. The results of studies using molecular biology methods demonstrate that genes encoding nitrate and ammonium transporters are rapidly down-regulated when nitrogen is resupplied to nitrogen-starved plants. Provision of ammonium to roots of plants actively absorbing nitrate imposes a block on nitrate uptake, the extent of which depends on the ammonium concentration, thus further reducing the efficient utilization of soil nitrate. During the daily variation of incoming light and during periods of low incident irradiation (i.e. heavy cloud cover) the expression levels of genes encoding nitrate and ammonium transporters, and rates of nitrate and ammonium uptake, are substantially reduced. Low temperatures reduce growth and nitrogen demand, and appear to discriminate against high-affinity nitrogen influx. In sum, these several factors conspire to limit rates of plant nitrogen uptake to values that are well below capacity. These characteristics of the plant's nitrogen uptake systems facilitate nitrogen losses from soils.     

不同有机氮效率的甜菜基因型筛选及差异分析

通过对不同基因型甜菜土壤有机氮利用及吸收效率的筛选和差异分析,为土壤有机氮高效基因型甜菜的栽培及品种选育提供理论依据。2014-2015年选取100份不同基因型的甜菜材料通过室内及田间试验在甜菜的不同发育阶段比较并分析土壤有机氮效率,筛选出对有机氮利用及吸收效率均显著差异的高效和低效基因型甜菜材料。结果表明,初步筛选得到的有机氮高效基因型甜菜材料KWS8138、HI0466和有机氮低效基因型甜菜材料BETA176、T230苗期全株及根部有机氮利用效率（Organic Nitrogen Use Efficiency,ONUE）差异显著;之后通过田间试验对有机氮吸收效率（Organic Nitrogen Assimilation Efficiency,ONAE）做了进一步筛选,发现KWS8138不但对ONUE有显著优势,还具有较高的有机氮素吸收能力,包括苗期之后对土壤有机氮素的运转量较高,合理的根冠比等。有机氮低效基因型甜菜材料BETA176的有机氮素吸收利用能力很弱、氮素转运能力过低等限制了植株对有机氮素的合理利用,不利于有机氮效率的提高。因此确定KWS8138为有机氮高效基因型材料,BETA176为有机氮低效基因型材料,均可作为进一步试验的材料。有机氮高效基因型甜菜较高的土壤有机氮转运量及合理的根冠比促进了其对有机氮素的吸收,是有机氮高效的基础。较高的干物质生产效率反应了甜菜对有机氮素的高效利用,是有机氮高效的关键。     

土壤易矿化有机态氮和微生物态氮作为土壤氮素生物有效性指标的评价

通过盆栽试验研究了土壤易矿化有机态氮和土壤微生物态氮与土壤净矿化氮及植物吸氮量之间的关系。结果表明,种植前土壤易矿化有机态氮和土壤微生物态氮以及种植前后土壤易矿化有机态氮的变化量均与土壤氮素净矿化量和植物吸氮量之间存在显著的相关性。在盆栽试验条件下,土壤易矿化有机态氮和种植前土壤微生物态氮能够较好地反映土壤氮素的矿化和供应能力,可以作为土壤氮素的生物有效性指标。     

Effect of Nitrogen Supply on the Nitrogen Use Efficiency of an Annual Herb, Helianthus annuus L.

Nitrogen use efficiency (NUE) is the product of nitrogen productivity (NP) and the mean residence time of nitrogen (MRT). Theory suggests that there should be a trade-off between both components,but direct experimental evidence is still scarce. To test this hypothesis, we analyzed the effect of varying nitrogen supply levels on NUEand its two components (NP, MRT) in Helianthus annuus L., an annual herb.The plants investigated were subjected to six nitrogen levels (0, 2, 4, 8, 16, and 32 g N/m2). Total plant production increased substantially with increasing nitrogen supply. Nitrogen uptake and loss also in creased with nitrogen supply. Nitrogen influx (rin) and outflux (rout) were defined as the rates of nitrogen uptake and loss per unit aboveground nitrogen, respectively. Both rin and rout increased with increasing nitrogen supply. In addition, rin was far higher than rout. Consequently, the relative rate of nitrogen incre ment (rin- rout) also increased with nitrogen supply. There were marked differences between treatments with respect to parameters related to the stress resistance syndrome: nitrogen pool size, leaf nitrogen concentration,and net aboveground productivity increased with nitrogen supply. Plants at high nitrogen levels showed a higher NP (the growth rate per unit aboveground nitrogen) and a shorter MRT (the inverse of rout), whereas plants at low nitrogen levels displayed the reverse pattern. Shorter MRT for plants at high nitrogen levels was caused by the abscission of leaves that contained relatively large fractions of total plant nitrogen. We found a negative relationship between NP and MRT, the components of NUE, along the gradient of nitrogen availability, suggesting that there was a trade-off between NP and MRT. The NUE increased with increasing nitrogen availability, up to a certain level, and then decreased. These results offer support for the hypoth esis that adaptation to infertile habitats involves a low nitrogen loss (long MRT in the plant) rather than a high NUE per se. The higher NUE at the plant level was a result, in part, of greater nitrogen resorption during senescence. We suggest that a long MRT (an index of nitrogen conservation) is a potentially successful strategy in nitrogen-poor environments.     

向日葵种群中植株个体大小对其氮素利用策略的影响

我们利用Berendse和Aerts提出的氮素利用效率(NUE)概念及原理研究了高密度一年生草本植物向日葵(Helianthus annuus L.)种群中植株个体大小对其氮素吸收利用的影响,并对种内竞争进行了分析.结果表明,植株对氮素的吸收与其个体大小不成线性关系,说明种群内不同植株个体对土壤氮素的竞争属于非对称竞争.植株的氮素损失随着个体大小的增加而增加.个体较大的植株具有较高的氮素输入率和较低的氮素输出率,因而具有较高的氮素净增加值.植株的氮素生产力(NP)和氮素平均滞留时间(MRT)均与植株个体大小呈正相关.较大的植物个体具有较高的NP和较长的MRT,由于NUE为NP和MRT二者的乘积,因而较大个体植株的NUE高于个体较小的植株.同种植物的不同个体的NP和MRT之间不存在协衡关系.氮素回收效率(NRE)与植株个体大小密切相关.在个体水平上,较大的植株个体具有较高的NUE与其较高的NRE有关.种群内植株个体对土壤氮素的非对称竞争主要由于植株对氮素的吸收和利用效率不同所致.因此,Berendse和Aerts提出的氮素利用效率概念不仅适用于研究种间的养分利用策略,对于种内不同植株的养分策略研究也同样适用.     

陆地生态系统植物的氮源及氮素吸收

氮是植物生长发育所必需的营养元素,也是其主要的限制因子之一.陆地生态系统植物所需氮的来源及植物对氮素的吸收利用均受控于其种类和生长环境.环境条件的改变,一方面可能改变植物生长区原有氮的形态、浓度、赋存方式等,从而改变氮对植物的供给状况;另一方面可能引起植物生长区土壤质量、水分利用状况、光照等的改变,从而产生耦合现象,直接影响植物的生理生态特性,使植物对氮素的吸收利用发生改变,导致植物生长区的种群类型及物种多样性发生改变,并直接影响到生态系统的功能及演替.本文主要对陆地生态系统中高等植物生长发育所需氮素的来源及植物对氮素吸收利用过程中的影响因素进行了综述和讨论,并结合国内外在该领域的研究现状对其研究前景进行了展望.     

植物氮形态利用策略及对外来植物入侵性的影响

氮是影响外来植物入侵性的重要因素之一, 但相关研究多关注土壤氮水平的效应, 较少考虑氮形态的作用。为从土壤氮形态利用的角度阐释外来植物的入侵机制, 本文在植物氮形态利用策略分析的基础上, 综述了外来植物氮形态利用的偏好性及其对入侵性的影响。植物的氮形态利用策略有偏好性和可塑性两种, 这可能与植物对土壤氮形态特性的长期适应有关; 植物不仅可以对土壤氮形态做出响应, 而且还能改造土壤氮形态, 并对改变后的土壤氮形态做出反馈响应。很多外来植物入侵硝态氮占优势的干扰生境, 偏好硝态氮的外来植物与本地植物竞争硝态氮; 而偏好铵态氮的外来植物通过抑制土壤硝化作用, 营造铵态氮环境, 促进自身生长, 同时抑制偏好硝态氮的本地植物生长。然而, 植物氮形态利用策略不是一成不变的, 而是受多种生物和非生物因素共同作用影响的复杂过程, 今后应加强多因素交互作用对外来入侵植物氮形态利用策略的影响及机制研究, 更好地揭示氮形态利用策略, 尤其是氮形态利用的可塑性与外来植物入侵性的关系。     

硝态氮（NO3^—）对水稻侧根生长及其氮吸收的影响

侧根是植物吸收利用土壤养分的重要器官 ,其生长发育受内部遗传因子和外部环境矿质养分的影响。通过琼脂分层培养发现 :局部供应NO-3 可以诱导水稻 (OryzasativaL .)主根或不定根上侧根的生长。为研究旱种条件下NO-3 对水稻侧根发育及其N吸收的影响 ,设置了 3个蛭石培养实验 :分根处理、全株缺N、全株供N处理。分根处理 (一半根系供应 3mmol/LKNO3,另一半根系供应 3mmol/LKCl)结果表明 :局部供应NO-3 能够促进水稻侧根生长。而在全株处理下 ,N饥饿诱导了侧根的伸长。水稻根系对NO-3 的这两种反应都存在着显著的基因型差异。同时对地上部N浓度、可溶性总糖含量及N含量分析表明 ,这些生理指标在分根处理与全株加N处理中的差异均不显著 ,表明分根处理也能基本满足植株正常生长对N的需求。在分根处理中 ,水稻的N含量与分根处理中供N一侧的平均侧根长度存在显著正相关 ,这表明在养分不均一的介质中 ,侧根长度对水稻N素吸收具有十分重要的作用。而在N素充足的条件下 ,两者之间的相关性并不显著 ,这暗示在养分充足的环境下 ,侧根长度可能并不是决定根系吸收N素的主要因素     

Nitrogen use efficiency. 2. Amino acid metabolism

In a previous article, we highlighted the latest developments in the isolation and characterisation of genes involved in the uptake of nitrogen from the soil, which might be used to improve the nitrogen use efficiency (NUE) of crop plants. In this article, we have concentrated on the genes controlling the enzymes of amino acid metabolism that may be involved in transferring nitrogen to the protein in the grain. Evidence is now accumulating from the use of knockout mutants, of the role of individual isoenzymes involved in amino acid metabolism, which are encoded by specific genes that are often members of a multigene family. In addition, a significant number of overexpressing plant lines have been obtained, which have increased activities of cytosol located, glutamine synthetase, asparagine synthetase and alanine aminotransferase that appear to have improved NUE.     

根际微生物促进水稻氮利用的机制

根际微生物影响植物的生长及环境适应性。不同种属、不同种群的植物影响其环境微生物群落;反之,根际微生物也影响宿主植物生长发育与生态适应性。植物与根际微生物的互作现象及其机制,是生命科学研究关注的热点,也是农业微生物利用的关键问题。近期,中国科学家在该领域取得了突破性进展。通过对不同籼稻(Oryza sativa subsp. indica)和粳稻(O. sativa subsp. japonica)品种的根际微生物组进行研究,发现籼稻根际比粳稻根际富集更多参与氮代谢的微生物群落,且该现象与硝酸盐转运蛋白基因NRT1.1B在籼粳之间的自然变异相关联。通过对籼稻接种籼稻根际特异富集的微生物群体可以提高前者对有机氮的利用,促进其生长。该研究揭示了籼稻和粳稻根际微生物分化的分子基础,展示了利用根际微生物提高水稻营养高效吸收的应用前景。     

Nitrogen Relations in a Forest Plantation--Soil Organic Matter Ecosystem Model

Two previously published models, after minor modification, areamalgamated to give a model that describes the major carbonand nitrogen pools and fluxes in a plantation forest soil system.The first model is a transport-resistance model of forest growthand dry-matter partitioning. The second is a soil organic mattermodel that was constructed for temperate grasslands. The combinedmodel is used to examine the relations between plantation growth,soil organic matter content, nitrogen deposition rate from theatmosphere, mineralization flux, nitrogen uptake by the plantation,dry matter partitioning between foliage and root, litter productionand the timing and quantity of fertilizer application. The highdemand for N by even-aged plantations during the period of canopybuilding is highlighted. The marked ontogenetic shifts in thegrowth pattern during plantation development is emphasized,indicating several phases of forest development. The resultsindicate that the potential growth of even-aged plantationsmay be greater than that realized in poor soils with commonlevels of atmospheric N deposition and normal fertilizer regimes.The simulations show how the concentrations of soil mineralN change during the development of a plantation, and point towardsthe importance of atmospheric N deposition. They also show thatfertilizer application must be accurately matched to growthstage if fertilizer is to be used efficiently. The nitrogencycle (N-uptake by plant     

Modeling long-term crop response to fertilizer and soil nitrogen

A simple nitrogen balance model to calculate long-term changes in soil organic nitrogen, nitrogen uptake by the crop and recovery of applied nitrogen, is presented. It functions with time intervals of one year or one growing season. In the model a labile and a stable pool of soil organic nitrogen are distinguished. Transfer coefficients for the various inputs of nitrogen are established that specify the fractions taken up by the crop, lost from the system, and incorporated in soil organic nitrogen. It is shown how input data, model parameters and initial pool sizes can be derived and how the model can be used for calculating long-term changes in total soil organic nitrogen and uptake by the crop. For nitrogen applied annually as fertilizer or organic material the time course of nitrogen uptake and recovery of applied nitrogen is calculated. To test the sensitivity of the model, calculations have been performed for different environmental conditions with higher or lower risks for losses. The model has also been applied to establish fertilizer recommendations for a certain target nitrogen uptake by the crop. Finally, for agricultural systems where periods of cropping alternate with peroids of green fallow the time course of nitrogen uptake by the crop is calculated.     

硝态氮(NO-3)对水稻侧根生长及其氮吸收的影响

侧根是植物吸收利用土壤养分的重要器官,其生长发育受内部遗传因子和外部环境矿质养分的影响.通过琼脂分层培养发现:局部供应NO-3可以诱导水稻( Oryza sativa L.)主根或不定根上侧根的生长.为研究旱种条件下NO-3对水稻侧根发育及其N吸收的影响,设置了3个蛭石培养实验:分根处理、全株缺N、全株供N处理.分根处理(一半根系供应3 mmol/L KNO3,另一半根系供应3 mmol/L KCl)结果表明:局部供应NO-3 能够促进水稻侧根生长.而在全株处理下,N饥饿诱导了侧根的伸长.水稻根系对NO-3的这两种反应都存在着显著的基因型差异.同时对地上部N浓度、可溶性总糖含量及N含量分析表明,这些生理指标在分根处理与全株加N处理中的差异均不显著,表明分根处理也能基本满足植株正常生长对N的需求.在分根处理中,水稻的N含量与分根处理中供N一侧的平均侧根长度存在显著正相关,这表明在养分不均一的介质中,侧根长度对水稻N素吸收具有十分重要的作用.而在N素充足的条件下,两者之间的相关性并不显著,这暗示在养分充足的环境下,侧根长度可能并不是决定根系吸收N素的主要因素.     

The Pitcher Plant Sarracenia purpurea Can Directly Acquire Organic Nitrogen and Short-Circuit the Inorganic Nitrogen Cycle

Background

Despite the large stocks of organic nitrogen in soil, nitrogen availability limits plant growth in many terrestrial ecosystems because most plants take up only inorganic nitrogen, not organic nitrogen. Although some vascular plants can assimilate organic nitrogen directly, only recently has organic nitrogen been found to contribute significantly to the nutrient budget of any plant. Carnivorous plants grow in extremely nutrient-poor environments and carnivory has evolved in these plants as an alternative pathway for obtaining nutrients. We tested if the carnivorous pitcher plant Sarracenia purpurea could directly take up intact amino acids in the field and compared uptake of organic and inorganic forms of nitrogen across a gradient of nitrogen deposition. We hypothesized that the contribution of organic nitrogen to the nitrogen budget of the pitcher plant would decline with increasing nitrogen deposition.

Methodology and Principal Findings

At sites in Canada (low nitrogen deposition) and the United States (high nitrogen deposition), individual pitchers were fed two amino acids, glycine and phenylalanine, and inorganic nitrogen (as ammonium nitrate), individually and in mixture. Plants took up intact amino acids. Acquisition of each form of nitrogen provided in isolation exceeded uptake of the same form in mixture. At the high deposition site, uptake of organic nitrogen was higher than uptake of inorganic nitrogen. At the low deposition site, uptake of all three forms of nitrogen was similar. Completeness of the associated detritus-based food web that inhabits pitcher-plant leaves and breaks down captured prey had no effect on nitrogen uptake.

Conclusions and Significance

By taking up intact amino acids, Sarracenia purpurea can short-circuit the inorganic nitrogen cycle, thus minimizing potential bottlenecks in nitrogen availability that result from the plant''s reliance for nitrogen mineralization on a seasonally reconstructed food web operating on infrequent and irregular prey capture.     

向日葵种群中植株个体大小对其氮素利用策略的影响

我们利用Berendse和Aerts提出的氮素利用效率(NUE)概念及原理研究了高密度一年生草本植物向日葵(Helianthus annuus L.)种群中植株个体大小对其氮素吸收利用的影响,并对种内竞争进行了分析。结果表明,植株对氮素的吸收与其个体大小不成线性关系,说明种群内不同植株个体对土壤氮素的竞争属于非对称竞争。植株的氮素损失随着个体大小的增加而增加。个体较大的植株具有较高的氮素输入率和较低的氮素输出率,因而具有较高的氮素净增加值。植株的氮素生产力(NP)和氮素平均滞留时间(MRT)均与植株个体大小呈正相关。较大的植物个体具有较高的NP和较长的MRT,由于NUE为NP和MRT二者的乘积,因而较大个体植株的NUE高于个体较小的植株。同种植物的不同个体的NP和MRT之间不存在协衡关系。氮素回收效率(NRE)与植株个体大小密切相关。在个体水平上,较大的植株个体具有较高的NUE与其较高的NRE有关。种群内植株个体对土壤氮素的非对称竞争主要由于植株对氮素的吸收和利用效率不同所致。因此,Berendse和Aerts提出的氮素利用效率概念不仅适用于研究种间的养分利用策略,对于种内不同植株的养分策略研究也同样适用。