15N in symbiotic fungi and plants estimates nitrogen and carbon flux rates in Arctic tundra

When soil nitrogen is in short supply, most terrestrial plants form symbioses with fungi (mycorrhizae): hyphae take up soil nitrogen, transport it into plant roots, and receive plant sugars in return. In ecosystems, the transfers within the pathway fractionate nitrogen isotopes so that the natural abundance of 15N in fungi differs from that in their host plants by as much as 12% per hundred. Here we present a new method to quantify carbon and nitrogen fluxes in the symbiosis based on the fractionation against 15N during transfer of nitrogen from fungi to plant roots. We tested this method, which is based on the mass balance of 15N, with data from arctic Alaska where the nitrogen cycle is well studied. Mycorrhizal fungi provided 61-86% of the nitrogen in plants; plants provided 8-17% of their photosynthetic carbon to the fungi for growth and respiration. This method of analysis avoids the disturbance of the soil-microbe-root relationship caused by collecting samples, mixing the soil, or changing substrate concentrations. This analytical technique also can be applied to other nitrogen-limited ecosystems, such as many temperate and boreal forests, to quantify the importance for terrestrial carbon and nitrogen cycling of nutrient transfers mediated by mycorrhizae at the plant-soil interface.     

Spatial decoupling of facilitation and competition at the origin of gapped vegetation patterns

Spatially periodic vegetation patterns, forming gaps, bands, labyrinths, or spots, are characteristic of arid and semiarid landscapes. Self-organization models can explain this variety of structures within a unified conceptual framework. All these models are based on the interplay of positive and negative effects of plants on soil water, but they can be divided according to whether they assume the interactions to be mediated by water redistribution through runoff/diffusion or by plants' organs. We carried out a multi-proxy approach of the processes operating in a gapped pattern in southwest Niger dominated by a shrub species. Soil moisture within the root layer was monitored in time and space over one month of the rainy season. Soil water recharge displayed no spatial variation with respect to vegetation cover, but the stock half-life under cover was twice that of bare areas. A kernel of facilitation by the aboveground parts of shrubs was parameterized, and soil water half-life was significantly correlated to the cumulated facilitative effects of shrubs. The kernel range was found to be smaller than the canopy radius (81%). This effect of plants on soil water dynamics, probably through a reduction of evaporation by shading, is shown to be a better explanatory variable than potentially relevant soil and topography parameters. The root systems of five individuals of Combretum micranthum G. Don were excavated. Root density data were used as a proxy to parameterize a kernel function of interplant competition. The range of this kernel was larger than the canopy radius (125%). The facilitation-to-competition range ratio, reflecting the above-to-belowground ratio of plant lateral extent, was smaller than 1 (0.64), a result supporting models assuming that patterning may emerge from an adaptation of plant morphology to aridity and shallow soils by means of an extended lateral root system. Moreover, observed soil water gradients had directions opposite to those assumed by alternative mathematical models based on underground water diffusion. This study contributes to the growing awareness that combined facilitative and competitive plant interactions can induce landscape-scale patterns and shape the two-way feedback loops between environment and vegetation.     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

大气CO2升高对土壤碳循环影响的研究进展

土壤有机碳是陆地碳库的重要组成部分,其积累和分解的变化直接影响全球的碳平衡。据估计,全球土壤（表层1m）有机碳积累总量相当于大气中碳总量的2～3倍。土壤是温室气体的源或汇,土壤碳库的变化将影响大气C02的浓度,因此,土壤碳库对人类活动的响应也是全球碳循环和全球变化研究的热点。在全球变化的大背景下,大气CO2升高导致植被生态系统碳平衡的改变进而对土壤碳循环产生影响。总结了陆地生态系统碳循环对大气C02浓度升高响应的主要生物学机制及过程,简述了大气C02浓度升高对影响土壤碳输入和输出的各因素的研究进展,并指出未来研究的主要方向。在大气C02浓度升高条件下,陆地生态系统碳循环的变化主要反映在以下几个方面：1）不同类型植物群落的净初级生产力（NPP）显著增加,但湿地植物的净初级生产力也有可能降低;2）光合产物向根系分配的数量增加,地上／地下生物量降低,根系形态发生变化,根系周转速率和根系分泌等过程的碳流量提高;3）植物含氮量降低,C／N提高,次生代谢产物增加,微生物生长受到抑制,植物残体分解速率降低;4）土壤呼吸速率显著增加,提高幅度受植物类型与土壤状况的影响;5）进入土壤的植物残体及分泌物的数量和性质影响土壤酶的活性,脱氢酶和转化酶活性增加,酚氧化酶和纤维素酶受植物类型与环境条件的影响;6）土壤中真菌的数量的增加幅度要高于细菌;7）CH4释放量增加,在植物的生长期表现更为明显。由于陆地生态系统碳循环的复杂性,研究结果仍有很大的不确定性。大气C02浓度升高与全球变化的其它表现间的交互作用将是今后研究的重点,同时由于土壤碳循环是一个由微生物介导的生物地球化学循环过程,因此,加强陆地生态系统碳循环的微生物机制研究也将为全面理解碳循环的过程提供更加准确的研究理论基础。     

From single fine roots to a black alder forest ecosystem: How system behaviour emerges from single component activities

Based on empirical findings in a natural black alder ecosystem in Northern Germany we developed an individual based model that integrates components of a black alder ecosystem interacting on different levels of organisation. The factors determining seasonal fine root biomass development of forest ecosystems are not yet fully understood.We used an object oriented model approach to investigate this complex matter for black alder trees. Processes like growth, storage, respiration, transport, nutrient mineralisation and uptake as well as interactions among these factors are described on the level of functionally differentiated plant organs (fine roots, coarse roots, stem, branches, leaves) and soil units. The object structure of the model is determined by spatial relations between plant modules as well as between plant modules and their local environment modules.As results of model application we found that (i) on the organ level, spatio-temporal plasticity of (root) growth allocation is related to spatio-temporal variation of resource availability, (ii) on the plant level, balanced root:shoot growth appears in response to variation of available resources light and nutrients, (iii) on the population level, tree stand development (population structure, self-thinning) resulted from coexistence and competition between plant individuals.For the understanding of the root compartment it seems relevant that the model implementation of local scale fine root dynamics is consistent with a self-organised large scale spatial heterogeneity of fine root activity pattern. On the other hand, fine-root dynamics cannot be explained as a result of autonomous dynamics. A reference to above-ground processes is a necessary condition and the overall plant seems to act as an integrator providing boundary conditions for local activity pattern. At the same time fine-root characteristics are of some importance for properties on hierarchically higher levels, e.g. co-existence in a tree population or element cycling in the ecosystem.As a conclusion, modelling of the spatio-temporal dynamics of tree root systems appears as a paradigmatic example of scale and organisation level integrating processes.     

Grassland establishment under varying resource availability: a test of positive and negative feedback

The traditional logic of carbon (C) and nitrogen (N) interactions in ecosystems predicts further increases or decreases in productivity (positive feedback) in response to high and low fertility in the soil, respectively; but the potential for development of feedback in ecosystems recovering from disturbance is less well understood. Furthermore, this logic has been challenged in grassland ecosystems where frequent fires or grazing may reduce the contribution of aboveground litter inputs to soil organic matter pools and nutrient supply for plant growth, relative to forest ecosystems. Further, if increases in plant productivity increase soil C content more than soil N content, negative feedback may result from increased microbial demand for N making less available for plant growth. We used a field experiment to test for feedback in an establishing grassland by comparing aboveground net primary productivity (ANPP) and belowground pools and fluxes of C and N in soil with enriched, ambient, and reduced N availability. For eight years annual N enrichment increased ANPP, root N, and root tissue quality, but root C:N ratios remained well above the threshold for net mineralization of N. There was no evidence that N enrichment increased root biomass, soil C or N accrual rates, or storage of C in total, microbial, or mineralizable pools within this time frame. However, the net nitrogen mineralization potential (NMP) rate was greater following eight years of N enrichment, and we attributed this to N saturation of the microbial biomass. Grassland developing under experimentally imposed N limitation through C addition to the soil exhibited ANPP, root biomass and quality, and net NMP rate similar to the ambient soil. Similarity in productivity and roots in the reduced and ambient N treatments was attributed to the potentially high nitrogen-use efficiency (NUE) of the dominant C4 grasses, and increasing cover of legumes over time in the C-amended soil. Thus, in a developing ecosystem, positive feedback between soil N supply and plant productivity may promote enhanced long-term N availability and override progressive N limitation as C accrues in plant and soil pools. However, experimentally imposed reduction in N availability did not feed back to reduce ANPP, possibly due to shifts in NUE and functional group composition.     

The effect of diesel fuel on common vetch (Vicia sativa L.) plants

When petroleum hydrocarbons contaminate soil, the carbon:nitrogen (C:N) ratio of the soil is altered. The added carbon stimulates microbial numbers but causes an imbalance in the C:N ratio which may result in immobilization of soil nitrogen by the microbial biomass, leaving none available for plant growth. As members of Leguminosae fix atmospheric nitrogen to produce their own nitrogen for growth, they may prove more successful at growing on petroleum hydrocarbon contaminated sites. During a wider study on phytoremediation of diesel fuel contaminated soil, particular attention was given to the performance of legumes versus other plant species. During harvesting of pot experiments containing leguminous plants, a recurring difference in the number and formation of root nodules present on control and contaminated Common vetch (Vicia sativa L.) plants was observed. The total number of nodules per plant was significantly reduced in contaminated plants compared to control plants but nodules on contaminated plants were more developed than corresponding nodules on control plants. Plant performance of Common vetch and Westerwold's ryegrass (Lolium multiflorum L.) was compared to illustrate any difference between the ability of legumes and grasses to grow on diesel fuel contaminated soil. Common vetch was less affected by diesel fuel and performed better in low levels of diesel fuel contaminated soil than Westerwolds ryegrass. The total amount of diesel fuel remaining after 4 months in Common vetch planted soil was slightly less than in Westerwolds ryegrass planted soil.     

Nitrogen regulation of the climate-carbon feedback: evidence from a long-term global change experiment

Modeling studies have shown that nitrogen (N) strongly regulates ecosystem responses and feedback to climate warming. However, it remains unclear what mechanisms underlie N regulation of ecosystem-climate interactions. To examine N regulation of ecosystem feedback to climate change, we have conducted a warming and clipping experiment since November 1999 in a tallgrass prairie of the Great Plains, USA. Infrared heaters were used to elevate soil temperature by an average of 1.96 degrees C at a depth of 2.5 cm from 2000 to 2008. Yearly biomass clipping mimicked hay or biofuel feedstock harvest. We measured carbon (C) and N concentrations, estimated their content and C:N ratio in plant, root, litter, and soil pools. Warming significantly stimulated C storage in aboveground plant, root, and litter pools by 17%, 38%, and 29%, respectively, averaged over the nine years (all P < 0.05) but did not change soil C content or N content in any pool. Plant C:N ratio and nitrogen use efficiency increased in the warmed plots compared to the control plots, resulting primarily from increased dominance of C4 plants in the community. Clipping significantly decreased C and N storage in plant and litter pools (all P < 0.05) but did not have interactive effects with warming on either C or N pools over the nine years. Our results suggest that increased ecosystem nitrogen use efficiency via a shift in species composition toward C4 dominance rather than plant N uptake is a key mechanism underlying warming stimulation of plant biomass growth.     

气候变暖背景下森林土壤碳循环研究进展

由人类活动引起的温室效应以及由此造成的气候变暖对森林牛态系统的影响已引起人们的普遍关注.森林土壤碳循环作为全球碳循环的重要组成部分,是决定未来陆地牛物嘲表现为碳源/碳汇的关键环节,揭示这一作用对于准确理解全球变化背景下陆地生态系统碳循环过程具有重要的指导意义.本文主要通过论述影响土壤碳循环过程的5个方面(土壤呼吸、土壤微生物、土壤酶活性、凋落物输入与分解、土壤碳库),综述了近10 a来全球气候变暖对土壤碳循环过程的影响.近年来,尽管已开展了大量有关土壤碳循环对气候变暖的响应及反馈机制的研究,并取得了一定的成果,但研究结果仍然存在很大的不确定性.整合各种密切关联的全球变化现象,完善研究方法和实验手段,加强根际微生态系统碳循环过程与机理研究将是下一步研究的方向和重点.参70     

不同浓度植物根系分泌物微生态效应研究

探讨夹竹桃(Nerium oleander)根系分泌物对土壤微生物量碳、呼吸强度等微生物学特征及土壤微生物群落功能多样性的影响,深入揭示夹竹桃根系分泌物的微生态效应,通过向土壤中添加植物根系分泌物溶液的方法,研究了不同浓度(CK,15 mL去离子水作为对照;LC,5 mL分泌物+10 mL去离子水;MC,10 mL分泌...     

温带阔叶红松林表层土壤活性碳、氮库的季节动态

以长白山温带阔叶红松林为观测对象,在植被生长季节对森林表层土壤进行连续性采样分析,以研究森林土壤活性碳、氮库的季节性变化特征及其影响因素。结果表明,不同季节土壤水溶性有机碳和硝态氮含量从高到低依次为春季、夏季和秋季,其变化范围分别为48～80和0.68～2.30μg.g-1。不同季节土壤水浸提溶液的特征吸光系数ASUVA,254和土壤铵态氮含量均表现为夏季高于春季和秋季,其变化范围分别为3.19～3.94 L.mg-1.m-1和6.1～12.0μg.g-1。在小时间尺度上,土壤活性碳、氮库对土壤含水量变化具有敏感性。水分条件和植物根系活动是影响表层土壤活性碳、氮库动态变化的主要因素,土壤微生物活动起调节作用。     

CERES — A model of forest stand biomass dynamics for predicting trace contaminant,nutrient, and water effects. I. Model description

CERES is a forest stand growth model which incorporates sugar transport in order to predict both short-term effects and long-term accumulation of trace contaminants and/or nutrients when coupled with the soil chemistry model (SCHEM), and models of solute uptake (DIFMAS and DRYADS) of the Unified Transport Model, UTM. An important feature of CERES is its ability to interface with the soil-plant-atmosphere water model (PROSPER) as a means of both predicting and studying the effects of plant water status on growth and solute transport.CERES considers the biomass dynamics of plants, standing dead and litter with plants divided into leaves, stems, roots, and fruits. The plant parts are divided further into sugar substrate, storage, and in the case of stems and roots, heartwood components.Each ecosystem component is described by a mass balance equation written as a first-order ordinary differential equation.     

土壤多环芳烃污染根际修复研究进展

多环芳烃(polycyclicaromatichydrocarbons,PAHs)是环境中普遍存在的具有代表性的一类重要持久性有机污染物,具“三致性”、难降解性,在土壤环境中不断积累,严重危害着土壤的生产和生态功能、农产品质量和人类健康。修复土壤多环芳烃污染已成为研究的焦点。根际修复是利用植物-微生物和根际环境降解有机污染物的复合生物修复技术,是目前最具潜力的土壤生物修复技术之一。对国内外学者近年来在土壤多环芳烃污染根际修复的效果、根际修复机理和根际修复的影响因素方面的研究进展作了较系统的综述,并分别分析了单作体系、混作体系、多进程根际修复系统和接种植物生长促进菌根际修复系统对土壤多环芳烃的修复效果。指出根际环境对PAHs的修复主要有3种机制:根系直接吸收和代谢PAHs;植物根系释放酶和分泌物去除PAHs,增加根际微生物数量,提高其活性,强化微生物群体降解PAHs。并讨论了影响根际修复PAHs的环境因素如植物、土壤类型、PAHs理化性质、菌根真菌以及表面活性剂等。植物-表面活性剂结合的根际修复技术、PAHs胁迫下根际的动态调节过程、运用分子生物学技术并结合植物根分泌物的特异性筛选高效修复植物以及植物富集的PAHs代谢产物进行跟踪与风险评价将成为未来研究的主流。     

Herbivores enable plant survival under nutrient limited conditions in a model grazing system

We make a theoretical study of nitrogen cycling in a model of a grazing system with five compartments. The rates of uptake of nutrient by plants and herbivores are allowed nonlinear forms which involve no a priori assumptions about whether the system is subject to top-down or bottom-up control. We derive a method of piecewise linear approximation which allows analytical study of the system. We then use this method to investigate the properties of the equilibrium states of the system, and in particular whether the system favours donor- or recipient-control, the grazing optimization problem, and the potential benefits of herbivory to plant growth. We are able to generalise our results to all uptake functions of the same qualitative class as those considered, and to show that in general the system will tend to a stable equilibrium state of donor-controlled herbivory. In this model, the presence of the ‘right’ class of herbivore is not only beneficial to plant growth in certain circumstances, but can be essential to their survival, allowing plants to co-exist with herbivores under conditions in which they would be unable to survive alone.     

A mechanistic model of tree competition and facilitation for Mediterranean forests: Scaling from leaf physiology to stand dynamics

Mechanistic theories of plant competition developed to explain changes in community structure and dynamics along resource availability gradients have been mostly applied to temperate forests and grasslands where light and nutrients are the two main limiting resources. In contrast, the mechanisms underlying the structure and dynamics of water-limited plant communities have been little explored. Also previous mechanistic models rely either on complex simulators, which are difficult to interpret or on simple conceptual models, which ignore too many critical details. In this study, we develop a model of stand dynamics for light and water-limited forests of intermediate complexity and we provide an analytical framework for its analyses. The model is an individual-based simulator that describes the feedback between transpiration, stomatal function and soil water dynamics with asymmetrical competition for light and water. Trees allocate carbon to three main compartments: shoot, stem and roots. We use the model to explore general patterns that may emerge across levels of biological organization from the leaf to the stand. Model predictions are consistent with a number of features of Mediterranean forests structure and dynamics. At the plant-level the leaf-based tradeoff between carbon gain and water loss expresses as a tradeoff between mortality and growth. This tradeoff explains plant morphological changes in above-ground biomass and root to shoot allocation along a water availability gradient. At the community-level, tradeoffs among carbon acquisition and water loss govern the sign of plant interactions along the gradient. Coexistence among morphological types was not observed for the range of parameters and environmental conditions explored. Overall the model provides an unifying explanation for the observed changes in the sign of plant to plant interactions along environmental gradients as well as a process-based formulation that can be linked to empirical studies.     

林木根系衰老研究方法与机制

根系衰老研究方法主要有根窗直接观测和挖根取样间接观测。在整株水平上,树木同化碳的能力,碳在不同器官间的分配,尤其是在地上部分和地下部分间的分配比例,对根的萌生和衰老起着重要作用;地上部分的生长过程和健康状况也对根系的生长和寿命有很大影响;当树木的生长环境受到某种胁迫时,树木的抵抗力下降,容易招致病原菌的危害,造成根系衰老和死亡。在生态系统水平上,干旱洪涝、干扰等胁迫和树木一土壤间的养分循环都会引起树木生长环境的改变,对根系的衰老过程发生作用;季节变化使树木的地上部分和地下部分的生理活动处于不同的旺盛期,从而使碳的分配方式有些改变,影响到根系的生长;土壤中养分的存在形式,某些离子的浓度也直接影响到根的寿命;病原菌对根系的危害与土壤中养分含量变化有关。所以,衰老过程受环境条件的影响,伴随着代谢,RNA和蛋白质合成速率的下降和(或)膜与细胞器结构的改变。     

Contrasting effects of engineered carbon nanotubes on plants: a review

Rapid surge of interest for carbon nanotube (CNT) in the last decade has made it an imperative member of nanomaterial family. Because of the distinctive physicochemical properties, CNTs are widely used in a number of scientific applications including plant sciences. This review mainly describes the role of CNT in plant sciences. Contradictory effects of CNT on plants physiology are reported. CNT can act as plant growth inducer causing enhanced plant dry biomass and root/shoot lengths. At the same time, CNT can cause negative effects on plants by forming reactive oxygen species in plant tissues, consequently leading to cell death. Enhanced seed germination with CNT is related to the water uptake process. CNT can be positioned as micro-tubes inside the plant body to enhance the water uptake efficiency. Due to its ability to act as a slow-release fertilizer and plant growth promoter, CNT is transpiring as a novel nano-carbon fertilizer in the field of agricultural sciences. On the other hand, accumulation of CNT in soil can cause deleterious effects on soil microbial diversity, composition and population. It can further modify the balance between plant-toxic metals in soil, thereby enhancing the translocation of heavy metal(loids) into the plant system. The research gaps that need careful attention have been identified in this review.     

Nutrient subsidies to belowground microbes impact aboveground food web interactions

Historically, terrestrial food web theory has been compartmentalized into interactions among aboveground or belowground communities. In this study we took a more synthetic approach to understanding food web interactions by simultaneously examining four trophic levels and investigating how nutrient (nitrogen and carbon) and detrital subsidies impact the ability of the belowground microbial community to alter the abundance of aboveground arthropods (herbivores and predators) associated with the intertidal cord grass Spartina alterniflora. We manipulated carbon, nitrogen, and detrital resources in a field experiment and measured decomposition rate, soil nitrogen pools, plant biomass and quality, herbivore density, and arthropod predator abundance. Because carbon subsidies impact plant growth only indirectly (microbial pathways), whereas nitrogen additions both directly (plant uptake) and indirectly (microbial pathways) impact plant primary productivity, we were able to assess the effect of both belowground soil microbes and nutrient availability on aboveground herbivores and their predators. Herbivore density in the field was suppressed by carbon supplements. Carbon addition altered soil microbial dynamics (net potential ammonification, litter decomposition rate, DON [dissolved organic N] concentration), which limited inorganic soil nitrogen availability and reduced plant size as well as predator abundance. Nitrogen addition enhanced herbivore density by increasing plant size and quality directly by increasing inorganic soil nitrogen pools, and indirectly by enhancing microbial nitrification. Detritus adversely affected aboveground herbivores mainly by promoting predator aggregation. To date, the effects of carbon and nitrogen subsidies on salt marshes have been examined as isolated effects on either the aboveground or the belowground community. Our results emphasize the importance of directly addressing the soil microbial community as a factor that influences aboveground food web structure by affecting plant size and aboveground plant nitrogen.     

蒙古栎林生态系统的水分循环与养分循环关系

本文从群体水平上研究了蒙古栎林生态系统中的水分循环与养分循环的关系。结果表明,降雨是系统养分的重要输入。降雨对林冠的淋洗淋溶作用有助于养分的归还及缩短养分循环的途径。蒙古栎树干茎流量大,且其中含有的养分元素较多,从而使蒙古栎的根系能获得更多的水分与养分,有助于适应干旱、瘠薄的立地环境。径流是系统的水分输出,也是养分输出。蒙古栎林的下渗流约占整个径流的90％,且养分元素含量较低,从而有助于养分的积累。降雨在蒙古栎林内的分配格局不一,导致系统养分积累的明显差异。本文还探讨了干扰对养分循环与水分循环的影响。结果表明,采伐森林导致水分分配的变化,也使系统的养分发生亏损。只有把养分循环与水分循环结合起来,才能较完整地评价森林的生态功能。     

Nitrogen cycling during seven years of atmospheric CO2 enrichment in a scrub oak woodland

Experimentally increasing atmospheric CO2 often stimulates plant growth and ecosystem carbon (C) uptake. Biogeochemical theory predicts that these initial responses will immobilize nitrogen (N) in plant biomass and soil organic matter, causing N availability to plants to decline, and reducing the long-term CO2-stimulation of C storage in N limited ecosystems. While many experiments have examined changes in N cycling in response to elevated CO2, empirical tests of this theoretical prediction are scarce. During seven years of postfire recovery in a scrub oak ecosystem, elevated CO2 initially increased plant N accumulation and plant uptake of tracer 15N, peaking after four years of CO2 enrichment. Between years four and seven, these responses to CO2 declined. Elevated CO2 also increased N and tracer 15N accumulation in the O horizon, and reduced 15N recovery in underlying mineral soil. These responses are consistent with progressive N limitation: the initial CO2 stimulation of plant growth immobilized N in plant biomass and in the O horizon, progressively reducing N availability to plants. Litterfall production (one measure of aboveground primary productivity) increased initially in response to elevated CO2, but the CO2 stimulation declined during years five through seven, concurrent with the accumulation of N in the O horizon and the apparent restriction of plant N availability. Yet, at the level of aboveground plant biomass (estimated by allometry), progressive N limitation was less apparent, initially because of increased N acquisition from soil and later because of reduced N concentration in biomass as N availability declined. Over this seven-year period, elevated CO2 caused a redistribution of N within the ecosystem, from mineral soils, to plants, to surface organic matter. In N limited ecosystems, such changes in N cycling are likely to reduce the response of plant production to elevated CO2.