Nitrogen uptake, distribution, turnover, and efficiency of use in a CO2-enriched sweetgum forest

The Progressive Nitrogen Limitation (PNL) hypothesis suggests that ecosystems in a CO2-enriched atmosphere will sequester C and N in long-lived biomass and soil organic pools, thereby limiting available N and constraining the continued response of net primary productivity to elevated [CO2]. Here, we present a six-year record of N dynamics of a sweetgum (Liquidambar styraciflua) stand exposed to elevated [CO2] in the free-air CO2 enrichment (FACE) experiment at Oak Ridge, Tennessee, USA. We also evaluate the concept of PNL for this ecosystem from the perspective of N uptake, content, distribution, and turnover, and N-use efficiency. Leaf N content was 11% lower on a leaf mass basis (NM) and 7% lower on a leaf area basis (NA) in CO2-enriched trees. However, there was no effect of [CO2] on total canopy N content. Resorption of N during senescence was not altered by [CO2], so NM of litter, but not total N content, was reduced. The NM of fine roots was not affected, but the total amount of N required for fine-root production increased significantly, reflecting the large stimulation of fine-root production in this stand. Hence, total N requirement of the trees was higher in elevated [CO2], and the increased requirement was met through an increase in N uptake rather than increased retranslocation of stored reserves. Increased N uptake was correlated with increased net primary productivity (NPP). N-use efficiency, however, did not change with CO2 enrichment because increased N productivity was offset by lower mean residence time of N in the trees. None of the measured responses of plant N dynamics in this ecosystem indicated the occurrence of PNL, and the stimulation of NPP by elevated [CO2] was sustained for the first six years of the experiment. Although there are some indications of developing changes in the N economy, the N supply in the soil at this site may be sufficient to meet an increasing demand for available N, especially as the roots of CO2-enriched trees explore deeper in the soil profile.     

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.     

Litterfall 15N abundance indicates declining soil nitrogen availability in a free-air CO2 enrichment experiment

Forest productivity increases in response to carbon dioxide (CO2) enrichment of the atmosphere. However, in nitrogen-limited ecosystems, increased productivity may cause a decline in soil nitrogen (N) availability and induce a negative feedback on further enhancement of forest production. In a free-air CO2 enrichment (FACE) experiment, the response of sweetgum (Liquidambar styraciflua L.) productivity to elevated CO2 concentrations [CO2] has declined over time, but documenting an associated change in soil N availability has been difficult. Here we assess the time history of soil N availability through analysis of natural 15N abundance in archived samples of freshly fallen leaf litterfall. Litterfall delta15N declined from 1998 to 2005, and the rate of decline was significantly faster in elevated [CO2]. Declining leaf litterfall delta15N is indicative of a tighter ecosystem N cycle and more limited soil N availability. By integrating N availability over time and throughout the soil profile, temporal dynamics in leaf litterfall delta15N provide a powerful tool for documenting changes in N availability and the critical feedbacks between C and N cycles that will control forest response to elevated atmospheric CO2 concentrations.     

Nitrogen limitation of net primary productivity in terrestrial ecosystems is globally distributed

Our meta-analysis of 126 nitrogen addition experiments evaluated nitrogen (N) limitation of net primary production (NPP) in terrestrial ecosystems. We tested the hypothesis that N limitation is widespread among biomes and influenced by geography and climate. We used the response ratio (R approximately equal ANPP(N)/ANPP(ctrl)) of aboveground plant growth in fertilized to control plots and found that most ecosystems are nitrogen limited with an average 29% growth response to nitrogen (i.e., R = 1.29). The response ratio was significant within temperate forests (R = 1.19), tropical forests (R = 1.60), temperate grasslands (R = 1.53), tropical grasslands (R = 1.26), wetlands (R = 1.16), and tundra (R = 1.35), but not deserts. Eight tropical forest studies had been conducted on very young volcanic soils in Hawaii, and this subgroup was strongly N limited (R = 2.13), which resulted in a negative correlation between forest R and latitude. The degree of N limitation in the remainder of the tropical forest studies (R = 1.20) was comparable to that of temperate forests, and when the young Hawaiian subgroup was excluded, forest R did not vary with latitude. Grassland response increased with latitude, but was independent of temperature and precipitation. These results suggest that the global N and C cycles interact strongly and that geography can mediate ecosystem response to N within certain biome types.     

Calcium constrains plant control over forest ecosystem nitrogen cycling

Forest ecosystem nitrogen (N) cycling is a critical controller of the ability of forests to prevent the movement of reactive N to receiving waters and the atmosphere and to sequester elevated levels of atmospheric carbon dioxide (CO2). Here we show that calcium (Ca) constrains the ability of northern hardwood forest trees to control the availability and loss of nitrogen. We evaluated soil N-cycling response to Ca additions in the presence and absence of plants and observed that when plants were present, Ca additions "tightened" the ecosystem N cycle, with decreases in inorganic N levels, potential net N mineralization rates, microbial biomass N content, and denitrification potential. In the absence of plants, Ca additions induced marked increases in nitrification (the key process controlling ecosystem N losses) and inorganic N levels. The observed "tightening" of the N cycle when Ca was added in the presence of plants suggests that the capacity of forests to absorb elevated levels of atmospheric N and CO2 is fundamentally constrained by base cations, which have been depleted in many areas of the globe by acid rain and forest harvesting.     

Temperature influences carbon accumulation in moist tropical forests

Evergreen broad-leaved tropical forests can have high rates of productivity and large accumulations of carbon in plant biomass and soils. They can therefore play an important role in the global carbon cycle, influencing atmospheric CO2 concentrations if climate warms. We applied meta-analyses to published data to evaluate the apparent effects of temperature on carbon fluxes and storages in mature, moist tropical evergreen forest ecosystems. Among forests, litter production, tree growth, and belowground carbon allocation all increased significantly with site mean annual temperature (MAT); total net primary productivity (NPP) increased by an estimated 0.2-0.7 Mg C x ha(-1) x yr(-1) x degrees C(-1). Temperature had no discernible effect on the turnover rate of aboveground forest biomass, which averaged 0.014 yr(-1) among sites. Consistent with these findings, forest biomass increased with site MAT at a rate of 5-13 Mg C x ha(-1) x degrees C(-1). Despite greater productivity in warmer forests, soil organic matter accumulations decreased with site MAT, with a slope of -8 Mg C x ha(-1) x degrees C(-1), indicating that decomposition rates of soil organic matter increased with MAT faster than did rates of NPP. Turnover rates of surface litter also increased with temperature among forests. We found no detectable effect of temperature on total carbon storage among moist-tropical evergreen forests, but rather a shift in ecosystem structure, from low-biomass forests with relatively large accumulations of detritus in cooler sites, to large-biomass forests with relatively smaller detrital stocks in warmer locations. These results imply that, in a warmer climate, conservation of forest biomass will be critical to the maintenance of carbon stocks in moist tropical forests.     

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。     

Improved estimates of net primary productivity from modis satellite data at regional and local scales.

We compared estimates of net primary production (NPP) from the MODIS satellite with estimates from a forest ecosystem process model (PnET-CN) and forest inventory and analysis (FIA) data for forest types of the mid-Atlantic region of the United States. The regional means were similar for the three methods and for the dominant oak-hickory forests in the region. However, MODIS underestimated NPP for less-dominant northern hardwood forests and overestimated NPP for coniferous forests. Causes of inaccurate estimates of NPP by MODIS were (1) an aggregated classification and parameterization of diverse deciduous forests in different climatic environments into a single class that averages different radiation conversion efficiencies; and (2) lack of soil water constraints on NPP for forests or areas that occur on thin or sandy, coarse-grained soil. We developed the "available soil water index" for adjusting the MODIS NPP estimates, which significantly improved NPP estimates for coniferous forests. The MODIS NPP estimates have many advantages such as globally continuous monitoring and remarkable accuracy for large scales. However, at regional or local scales, our study indicates that it is necessary to adjust estimates to specific vegetation types and soil water conditions.     

Progressive N limitation in forests: review and implications for long-term responses to elevated CO2

Field studies have shown that elevated CO2 can cause increased forest growth over the short term (<6 years) even in the face of N limitation. This is facilitated to some degree by greater biomass production per unit N uptake (lower tissue N concentrations), but more often than not, N uptake is increased with elevated CO2 as well. Some studies also show that N sequestration in the forest floor is increased with elevated CO2. These findings raise the questions of where the "extra" N comes from and how long such growth increases can continue without being truncated by progressive N limitation (PNL). This paper reviews some of the early nutrient cycling literature that describes PNL during forest stand development and attempts to use this information, along with recent developments in soil N research, to put the issue of PNL with elevated CO2 into perspective. Some of the early studies indicated that trees can effectively "mine" N from soils over the long term, and more recent developments in soil N cycling research suggest mechanisms by which this might have occurred. However, both the early nutrient cycling literature and more recent simulation modeling suggest that PNL will at some point truncate the observed increases in growth and nutrient uptake with elevated CO2, unless external inputs of N are increased by either N fixation or atmospheric deposition.     

Progressive nitrogen limitation of ecosystem processes under elevated CO2 in a warm-temperate forest

A hypothesis for progressive nitrogen limitation (PNL) proposes that net primary production (NPP) will decline through time in ecosystems subjected to a step-function increase in atmospheric CO2. The primary mechanism driving this response is a rapid rate of N immobilization by plants and microbes under elevated CO2 that depletes soils of N, causing slower rates of N mineralization. Under this hypothesis, there is little long-term stimulation of NPP by elevated CO2 in the absence of exogenous inputs of N. We tested this hypothesis using data on the pools and fluxes of C and N in tree biomass, microbes, and soils from 1997 through 2002 collected at the Duke Forest free-air CO2 enrichment (FACE) experiment. Elevated CO2 stimulated NPP by 18-24% during the first six years of this experiment. Consistent with the hypothesis for PNL, significantly more N was immobilized in tree biomass and in the O horizon under elevated CO2. In contrast to the PNL hypothesis, microbial-N immobilization did not increase under elevated CO2, and although the rate of net N mineralization declined through time, the decline was not significantly more rapid under elevated CO2. Ecosystem C-to-N ratios widened more rapidly under elevated CO2 than ambient CO2 indicating a more rapid rate of C fixation per unit of N, a processes that could delay PNL in this ecosystem. Mass balance calculations demonstrated a large accrual of ecosystem N capital. Is PNL occurring in this ecosystem and will NPP decline to levels under ambient CO2? The answer depends on the relative strength of tree biomass and O-horizon N immobilization vs. widening C-to-N ratios and ecosystem-N accrual as processes that drive and delay PNL, respectively. Only direct observations through time will definitively answer this question.     

Elevated CO2 stimulates net accumulations of carbon and nitrogen in land ecosystems: a meta-analysis

The capability of terrestrial ecosystems to sequester carbon (C) plays a critical role in regulating future climatic change yet depends on nitrogen (N) availability. To predict long-term ecosystem C storage, it is essential to examine whether soil N becomes progressively limiting as C and N are sequestered in long-lived plant biomass and soil organic matter. A critical parameter to indicate the long-term progressive N limitation (PNL) is net change in ecosystem N content in association with C accumulation in plant and soil pools under elevated CO2. We compiled data from 104 published papers that study C and N dynamics at ambient and elevated CO2. The compiled database contains C contents, N contents, and C:N ratio in various plant and soil pools, and root:shoot ratio. Averaged C and N pool sizes in plant and soil all significantly increase at elevated CO2 in comparison to those at ambient CO2, ranging from a 5% increase in shoot N content to a 32% increase in root C content. The C and N contents in litter pools are consistently higher in elevated than ambient CO2 among all the surveyed studies whereas C and N contents in the other pools increase in some studies and decrease in other studies. The high variability in CO2-induced changes in C and N pool sizes results from diverse responses of various C and N processes to elevated CO2. Averaged C:N ratios are higher by 3% in litter and soil pools and 11% in root and shoot pools at elevated relative to ambient CO2. Elevated CO2 slightly increases root:shoot ratio. The net N accumulation in plant and soil pools at least helps prevent complete down-regulation of, and likely supports, long-term CO2 stimulation of C sequestration. The concomitant C and N accumulations in response to rising atmospheric CO2 may reflect intrinsic nature of ecosystem development as revealed before by studies of succession over hundreds to millions of years.     

Long-term CO2 enrichment of a forest ecosystem: implications for forest regeneration and succession.

The composition and successional status of a forest affect carbon storage and net ecosystem productivity, yet it remains unclear whether elevated atmospheric carbon dioxide (CO2) will impact rates and trajectories of forest succession. We examined how CO2 enrichment (+200 microL CO2/L air differential) affects forest succession through growth and survivorship of tree seedlings, as part of the Duke Forest free-air CO2 enrichment (FACE) experiment in North Carolina, USA. We planted 2352 seedlings of 14 species in the low light forest understory and determined effects of elevated CO2 on individual plant growth, survival, and total sample biomass accumulation, an integrator of plant growth and survivorship over time, for six years. We used a hierarchical Bayes framework to accommodate the uncertainty associated with the availability of light and the variability in growth among individual plants. We found that most species did not exhibit strong responses to CO2. Ulmus alata (+21%), Quercus alba (+9.5%), and nitrogen-fixing Robinia pseudoacacia (+230%) exhibited greater mean annual relative growth rates under elevated CO2 than under ambient conditions. The effects of CO2 were small relative to variability within populations; however, some species grew better under low light conditions when exposed to elevated CO2 than they did under ambient conditions. These species include shade-intolerant Liriodendron tulipifera and Liquidambar styraciflua, intermediate-tolerant Quercus velutina, and shade-tolerant Acer barbatum, A. rubrum, Prunus serotina, Ulmus alata, and Cercis canadensis. Contrary to our expectation, shade-intolerant trees did not survive better with CO2 enrichment, and population-scale responses to CO2 were influenced by survival probabilities in low light. CO2 enrichment did not increase rates of sample biomass accumulation for most species, but it did stimulate biomass growth of shade-tolerant taxa, particularly Acer barbatum and Ulmus alata. Our data suggest a small CO2 fertilization effect on tree productivity, and the possibility of reduced carbon accumulation rates relative to today's forests due to changes in species composition.     

Interactive effects of plant species diversity and elevated CO2 on soil biota and nutrient cycling

Terrestrial ecosystems consist of mutually dependent producer and decomposer subsystems, but not much is known on how their interactions are modified by plant diversity and elevated atmospheric CO2 concentrations. Factorially manipulating grassland plant species diversity and atmospheric CO2 concentrations for five years, we tested whether high diversity or elevated CO2 sustain larger or more active soil communities, affect soil aggregation, water dynamics, or nutrient cycling, and whether plant diversity and elevated CO2 interact. Nitrogen (N) and phosphorus (P) pools, symbiotic N2 fixation, plant litter quality, soil moisture, soil physical structure, soil nematode, collembola and acari communities, soil microbial biomass and microflora community structure (phospholipid fatty acid [PLFA] profiles), soil enzyme activities, and rates of C fluxes to soils were measured. No increases in soil C fluxes or the biomass, number, or activity of soil organisms were detected at high plant diversity; soil H2O and aggregation remained unaltered. Elevated CO2 affected the ecosystem primarily by improving plant and soil water status by reducing leaf conductance, whereas changes in C cycling appeared to be of subordinate importance. Slowed-down soil drying cycles resulted in lower soil aggregation under elevated CO2. Collembola benefited from extra soil moisture under elevated CO2, whereas other faunal groups did not respond. Diversity effects and interactions with elevated CO2 may have been absent because soil responses were mainly driven by community-level processes such as rates of organic C input and water use; these drivers were not changed by plant diversity manipulations, possibly because our species diversity gradient did not extend below five species and because functional type composition remained unaltered. Our findings demonstrate that global change can affect soil aggregation, and we advocate that soil aggregation should be considered as a dynamic property that may respond to environmental changes and feed back on other ecosystem functions.     

大气CO2体积分数升高对植物N素吸收的影响

从影响植物N素吸收的因素来看，大气CO2体积分数升高条件下植物净光合作用增强，碳同化产物增多，利于改善N素吸收的能量和物质基础：植物根系生长增强，生物量增多且空间分布加大，有利于N素吸收；但土壤有效N供应能力的变化存在增强和减弱两种观点。从植物N素吸收的实际情况来看，大气CO2体积分数升高条件下植物N吸收总量并末增加，植物体内N质量分数普遍降低，某些种类植物N吸收形态也发生了改变。因此要阐明大气CO2体积分数升高对植物N素吸收的影响机制，必须探明土壤有效N供应能力的变化：CO2体积分数升高条件下N矿化作用是否增强，微生物和植物间是否存在对有效N的竞争，此外，CO2体积分数升高条件下植物根系形态特征变化和N素吸收(包括主动和被动吸收)的生理机制及其与环境因素的关系也值得进一步研究。     

Influences of global environmental changes on arbuscular mycorrhizal fungal communities: a review北大核心CSCD

丛枝菌根(arbuscular mycorrhizal, AM)真菌是生态系统地上地下部的重要连接体,对其群落结构特征的研究有助于菌种资源的发掘和生态系统的可持续发展.人类生产生活活动对全球环境带来了一系列的改变,如二氧化碳和臭氧浓度升高、氮沉降、增温及降水减少/增多等,全球环境变化对AM真菌群落结构的影响也引起了广泛关注.针对二氧化碳和臭氧浓度升高、增温、氮沉降和降水减少/增多等全球环境变化因子,总结其对AM真菌群落结构影响的国内外研究进展,探讨全球环境变化对AM真菌群落的可能作用途径.已有模拟全球环境变化实验研究主要集中于北半球的草原、农田和森林系统.大多研究发现二氧化碳和臭氧浓度升高未对AM真菌多样性产生不利影响,但使AM真菌群落结构显著分异.氮沉降和增温对AM真菌多样性的影响表现为降低、无显著影响和增加等多种情况,对AM真菌群落结构的影响也表现为未显著和显著分异,主要与模拟实验处理方式、增加幅度、土壤养分水平和生态系统类型等因素有关.降水减少未显著影响AM真菌群落结构和多样性,而降水增加使AM真菌群落结构发生显著分异.这些研究主要注重AM真菌群落结构和多样性如何改变等生态现象而潜在机理探索以及热带和南半球不同生态系统下的研究尚不足.另外,鉴于全球变化因子间的关联性,复合因子对AM真菌群落结构的影响值得重视.(图1表4参113)     

CO2 enrichment inhibits shoot nitrate assimilation in C3 but not C4 plants and slows growth under nitrate in C3 plants

The CO2 concentration in Earth's atmosphere may double during this century. Plant responses to such an increase depend strongly on their nitrogen status, but the reasons have been uncertain. Here, we assessed shoot nitrate assimilation into amino acids via the shift in shoot CO2 and O2 fluxes when plants received nitrate instead of ammonium as a nitrogen source (deltaAQ). Shoot nitrate assimilation became negligible with increasing CO2 in a taxonomically diverse group of eight C3 plant species, was relatively insensitive to CO2 in three C4 species, and showed an intermediate sensitivity in two C3-C4 intermediate species. We then examined the influence of CO2 level and ammonium vs. nitrate nutrition on growth, assessed in terms of changes in fresh mass, of several C3 species and a Crassulacean acid metabolism (CAM) species. Elevated CO2 (720 micromol CO2/mol of all gases present) stimulated growth or had no effect in the five C3 species tested when they received ammonium as a nitrogen source but inhibited growth or had no effect if they received nitrate. Under nitrate, two C3 species grew faster at sub-ambient (approximately 310 micromol/mol) than elevated CO2. A CAM species grew faster at ambient than elevated or sub-ambient CO2 under either ammonium or nitrate nutrition. This study establishes that CO2 enrichment inhibits shoot nitrate assimilation in a wide variety of C3 plants and that this phenomenon can have a profound effect on their growth. This indicates that shoot nitrate assimilation provides an important contribution to the nitrate assimilation of an entire C3 plant. Thus, rising CO2 and its effects on shoot nitrate assimilation may influence the distribution of C3 plant species.     

N fertilization effects on denitrification and N cycling in an aggrading forest.

We investigated N cycling and denitrification rates following five years of N and dolomite amendments to whole-tree harvested forest plots at the long-term soil productivity experiment in the Fernow Experimental Forest in West Virginia, USA. We hypothesized that changes in soil chemistry and nutrient cycling induced by N fertilization would increase denitrification rates and the N2O:N2 ratio. Soils from the fertilized plots had a lower pH (2.96) than control plots (3.22) and plots that received fertilizer and dolomite (3.41). There were no significant differences in soil %C or %N between treatments. Chloroform-labile microbial biomass carbon was lower in fertilized plots compared to control plots, though this trend was not significant. Extractable soil NO3- was elevated in fertilized plots on each sample date. Soil-extractable NH4+, NO3-, pH, microbial biomass carbon, and %C varied significantly by sample date suggesting important seasonal patterns in soil chemistry and N cycling. In particular, the steep decline in extractable NH4+ during the growing season is consistent with the high N demands of a regenerating forest. Net N mineralization and nitrification also varied by date but were not affected by the fertilization and dolomite treatments. In a laboratory experiment, denitrification was stimulated by NO3- additions in soils collected from all field plots, but this effect was stronger in soils from the unfertilized control plots, suggesting that chronic N fertilization has partially alleviated a NO3- limitation on denitrification rates. Dextrose stimulated denitrification only in the whole-tree-harvest soils. Denitrification enzyme activity varied by sample date and was elevated in fertilized plots for soil collected in July 2000 and June 2001. There were no detectable treatment effects on N2O or N2 flux from soils under anaerobic conditions, though there was strong temporal variation. These results suggest that whole-tree harvesting has altered the N status of these soils so they are less prone to N saturation than more mature forests. It is likely that N losses associated with the initial harvest and high N demand by aggrading vegetation is minimizing, at least temporarily, the amount of inorganic N available for nitrification and denitrification, even in the fertilized plots in this experiment.     

施肥与不施肥措施下小麦田的CO2、CH4、N2O排放日变化特征

二氧化碳（CO2）、甲烷（CH4）、氧化亚氮（N2O）是对全球气候变化影响最大的温室气体。由于土壤与大气之间的水热交换需要一定的传导平衡时间,因此土壤温室气体与温湿度之间的关系存在不同的表现形式。目前,有关温室气体研究多集中于季节性排放特征,而关于CO2、CH4、N2O的日变化研究却少见报道。以北京小麦（Triticum aestivuml）农田土壤为研究对象,对施肥和不施肥条件下CO2、CH4、N2O交换通量和气温、土壤温度进行连续观测,来探讨3种温室气体的日变化特征。采用人工静态暗箱法对小麦田土壤进行连续48 h原位观测,每2 h测定1次,每次盖箱时间为30 min。气体样品中的CO2、CH4、N2O用气相色谱仪（Agilent 6890A,FID/ECD）测定。结果表明：施肥与不施肥条件下小麦生育后期麦田土壤CO2、CH4、N2O交换通量具有明显的日变化特征。土壤表现为CH4的吸收汇、CO2和N2O的排放源。CH4的吸收通量、CO2和N2O的排放通量均表现为施肥区＞对照区。CO2、CH4的交换通量的70%以上出现在白天,而施肥区和对照区的N2O白天排放通量分别达到全天的81.8%、91.1%。另外,相关性分析表明,CO2、N2O交换通量的日变化与气温和5 cm地温呈极显著（P＜0.01）或显著（P＜0.05）的正相关关系,且N2O交换通量日变化与10 cm地温呈现极显著的正相关关系,说明温度是影响CO2、N2O交换通量日变化的重要因素;而气温、5 cm地温、10 cm地温对CH4交换通量日变化不存在显著性影响。     

Sinks for nitrogen inputs in terrestrial ecosystems: a meta-analysis of 15N tracer field studies

Effects of anthropogenic nitrogen (N) deposition and the ability of terrestrial ecosystems to store carbon (C) depend in part on the amount of N retained in the system and its partitioning among plant and soil pools. We conducted a meta-analysis of studies at 48 sites across four continents that used enriched 15N isotope tracers in order to synthesize information about total ecosystem N retention (i.e., total ecosystem 15N recovery in plant and soil pools) across natural systems and N partitioning among ecosystem pools. The greatest recoveries of ecosystem 15N tracer occurred in shrublands (mean, 89.5%) and wetlands (84.8%) followed by forests (74.9%) and grasslands (51.8%). In the short term (< 1 week after 15N tracer application), total ecosystem 15N recovery was negatively correlated with fine-root and soil 15N natural abundance, and organic soil C and N concentration but was positively correlated with mean annual temperature and mineral soil C:N. In the longer term (3-18 months after 15N tracer application), total ecosystem 15N retention was negatively correlated with foliar natural-abundance 15N but was positively correlated with mineral soil C and N concentration and C:N, showing that plant and soil natural-abundance 15N and soil C:N are good indicators of total ecosystem N retention. Foliar N concentration was not significantly related to ecosystem 15N tracer recovery, suggesting that plant N status is not a good predictor of total ecosystem N retention. Because the largest ecosystem sinks for 15N tracer were below ground in forests, shrublands, and grasslands, we conclude that growth enhancement and potential for increased C storage in aboveground biomass from atmospheric N deposition is likely to be modest in these ecosystems. Total ecosystem 15N recovery decreased with N fertilization, with an apparent threshold fertilization rate of 46 kg N x ha(-1) x yr(-1) above which most ecosystems showed net losses of applied 15N tracer in response to N fertilizer addition.     

CO2-enrichment and nutrient availability alter ectomycorrhizal fungal communities

Ectomycorrhizal fungi (EMF), a phylogenetically and physiologically diverse guild, form symbiotic associations with many trees and greatly enhance their uptake of nutrients and water. Elevated CO2, which increases plant carbon supply and demand for mineral nutrients, may change the composition of the EMF community, possibly altering nutrient uptake and ultimately forest productivity. To assess CO2 effects on EMF communities, we sampled mycorrhizae from the FACTS-I (Forest-Atmosphere Carbon Transfer and Storage) research site in Duke Forest, Orange County, North Carolina, USA, where Pinus taeda forest plots are maintained at either ambient or elevated CO2 (200 ppm above ambient) concentrations. Mycorrhizae were identified by DNA sequence similarity of the internal transcribed spacer ribosomal RNA gene region. EMF richness was very high; 72 distinct phylotypes were detected from 411 mycorrhizal samples. Overall EMF richness and diversity were not affected by elevated CO2, but increased CO2 concentrations altered the relative abundances of particular EMF taxa colonizing fine roots, increased prevalence of unique EMF species, and led to greater EMF community dissimilarity among individual study plots. Natural variation among plots in mean potential net nitrogen (N) mineralization rates was a key determinant of EMF community structure; increasing net N mineralization rate was negatively correlated with EMF richness and had differential effects on the abundance of particular EMF taxa. Our results predict that, at CO2 concentrations comparable to that predicted for the year 2050, EMF community composition and structure will change, but diversity will be maintained. In contrast, high soil N concentrations can negatively affect EMF diversity; this underscores the importance of considering CO2 effects on forest ecosystems in the context of background soil chemical parameters and other environmental perturbations such as acid deposition or fertilizer runoff.