Atmospheric CO2 and O3 alter the flow of 15N in developing forest ecosystems

Anthropogenic O3 and CO2-induced declines in soil N availability could counteract greater plant growth in a CO2-enriched atmosphere, thereby reducing net primary productivity (NPP) and the potential of terrestrial ecosystems to sequester anthropogenic CO2. Presently, it is uncertain how increasing atmospheric CO2 and O3 will alter plant N demand and the acquisition of soil N by plants as well as the microbial supply of N from soil organic matter. To address this uncertainty, we initiated an ecosystem-level 15N tracer experiment at the Rhinelander (Wisconsin, USA) free air CO2-O3 enrichment (FACE) facility to understand how projected increases in atmospheric CO2 and 03 alter the distribution and flow of N in developing northern temperate forests. Tracer amounts of 15NH4+ were applied to the forest floor of developing Populus tremuloides and P. tremuloides-Betula papyrifera communities that have been exposed to factorial CO2 and O3 treatments for seven years. One year after isotope addition, both forest communities exposed to elevated CO2 obtained greater amounts of 15N (29%) and N (40%) from soil, despite no change in soil N availability or plant N-use efficiency. As such, elevated CO2 increased the ability of plants to exploit soil for N, through the development of a larger root system. Conversely, elevated O3 decreased the amount of 15N (-15%) and N (-29%) in both communities, a response resulting from lower rates of photosynthesis, decreases in growth, and smaller root systems that acquired less soil N. Neither CO2 nor 03 altered the amount of N or 15N recovery in the forest floor, microbial biomass, or soil organic matter. Moreover, we observed no interaction between CO2 and 03 on the amount of N or 15N in any ecosystem pool, suggesting that 03 could exert a negative effect regardless of CO2 concentration. In a CO2-enriched atmosphere, greater belowground growth and a more thorough exploitation of soil for growth-limiting N is an important mechanism sustaining the enhancement of NPP in developing forests (0-8 years following establishment). However, as CO2 accumulates in the Earth's atmosphere, future O3 concentrations threaten to diminish the enhancement of plant growth, decrease plant N acquisition, and lessen the storage of anthropogenic C in temperate forests.     

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.     

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.     

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。     

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.     

Changes in microbial community characteristics and soil organic matter with nitrogen additions in two tropical forests

Microbial communities and their associated enzyme activities affect the amount and chemical quality of carbon (C) in soils. Increasing nitrogen (N) deposition, particularly in N-rich tropical forests, is likely to change the composition and behavior of microbial communities and feed back on ecosystem structure and function. This study presents a novel assessment of mechanistic links between microbial responses to N deposition and shifts in soil organic matter (SOM) quality and quantity. We used phospholipid fatty acid (PLFA) analysis and microbial enzyme assays in soils to assess microbial community responses to long-term N additions in two distinct tropical rain forests. We used soil density fractionation and 13C nuclear magnetic resonance (NMR) spectroscopy to measure related changes in SOM pool sizes and chemical quality. Microbial biomass increased in response to N fertilization in both tropical forests and corresponded to declines in pools of low-density SOM. The chemical quality of this soil C pool reflected ecosystem-specific changes in microbial community composition. In the lower-elevation forest, there was an increase in gram-negative bacteria PLFA biomass, and there were significant losses of labile C chemical groups (O-alkyls). In contrast, the upper-elevation tropical forest had an increase in fungal PLFAs with N additions and declines in C groups associated with increased soil C storage (alkyls). The dynamics of microbial enzymatic activities with N addition provided a functional link between changes in microbial community structure and SOM chemistry. Ecosystem-specific changes in microbial community composition are likely to have far-reaching effects on soil carbon storage and cycling. This study indicates that microbial communities in N-rich tropical forests can be sensitive to added N, but we can expect significant variability in how ecosystem structure and function respond to N deposition among tropical forest types.     

大气CO2与植物氮素营养的关系

大气CO2浓度升高对植物吸收氮素,以及对植物和土壤中的氮浓度、C/N比和氮循环都存在着影响。大气CO2浓度与植物氮素营养之间存在着交互作用。大气CO2浓度升高对植物氮素营养影响的结果与氮浓度、氮形态等因素有关。     

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.     

Plant diversity, CO2, and N influence inorganic and organic N leaching in grasslands

In nitrogen (N)-limited systems, the potential to sequester carbon depends on the balance between N inputs and losses as well as on how efficiently N is used, yet little is known about responses of these processes to changes in plant species richness, atmospheric CO2 concentration ([CO2]), and N deposition. We examined how plant species richness (1 or 16 species), elevated [CO2] (ambient or 560 ppm), and inorganic N addition (0 or 4 g x m(-2) x yr(-1)) affected ecosystem N losses, specifically leaching of dissolved inorganic N (DIN) and organic N (DON) in a grassland field experiment in Minnesota, USA. We observed greater DIN leaching below 60 cm soil depth in the monoculture plots (on average 1.8 and 3.1 g N x m(-2) x yr(-1) for ambient N and N-fertilized plots respectively) than in the 16-species plots (0.2 g N x m(-2) x yr(-1) for both ambient N and N-fertilized plots), particularly when inorganic N was added. Most likely, loss of complementary resource use and reduced biological N demand in the monoculture plots caused the increase in DIN leaching relative to the high-diversity plots. Elevated [CO2] reduced DIN concentrations under conditions when DIN concentrations were high (i.e., in N-fertilized and monoculture plots). Contrary to the results for DIN, DON leaching was greater in the 16-species plots than in the monoculture plots (on average 0.4 g N x m(-2) x yr(-1) in 16-species plots and 0.2 g N x m(-2) x yr(-1) in monoculture plots). In fact, DON dominated N leaching in the 16-species plots (64% of total N leaching as DON), suggesting that, even with high biological demand for N, substantial amounts of N can be lost as DON. We found no significant main effects of elevated [CO2] on DIN or DON leaching; however, elevated [CO2] reduced the positive effect of inorganic N addition on DON leaching, especially during the second year of observation. Our results suggest that plant species richness, elevated [CO2], and N deposition alter DIN loss primarily through changes in biological N demand. DON losses can be as large as DIN loss but are more sensitive to organic matter production and turnover.     

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.     

No evidence that chronic nitrogen additions increase photosynthesis in mature sugar maple forests

Atmospheric nitrogen (N) deposition can increase forest growth. Because N deposition commonly increases foliar N concentrations, it is thought that this increase in forest growth is a consequence of enhanced leaf-level photosynthesis. However, tests of this mechanism have been infrequent, and increases in photosynthesis have not been consistently observed in mature forests subject to chronic N deposition. In four mature northern hardwood forests in the north-central United States, chronic N additions (30 kg N ha(-1) yr(-1) as NaNO3 for 14 years) have increased aboveground growth but have not affected canopy leaf biomass or leaf area index. In order to understand the mechanism behind the increases in growth, we hypothesized that the NO3(-) additions increased foliar N concentrations and leaf-level photosynthesis in the dominant species in these forests (sugar maple, Acer saccharum). The NO3(-) additions significantly increased foliar N. However, there was no significant difference between the ambient and +NO3(-) treatments in two seasons (2006-2007) of instantaneous measurements of photosynthesis from either canopy towers or excised branches. In measurements on excised branches, photosynthetic nitrogen use efficiency (micromol CO2 s(-1) g(-1) N) was significantly decreased (-13%) by NO3(-) additions. Furthermore, we found no consistent NO3(-) effect across all sites in either current foliage or leaf litter collected annually throughout the study (1993-2007) and analyzed for delta 13C and delta 18O, isotopes that can be used together to integrate changes in photosynthesis over time. We observed a small but significant NO3(-) effect on the average area and mass of individual leaves from the excised branches, but these differences varied by site and were countered by changes in leaf number. These photosynthesis and leaf area data together suggest that NO3(-) additions have not stimulated photosynthesis. There is no evidence that nutrient deficiencies have developed at these sites, so unlike other studies of photosynthesis in N-saturated forests, we cannot attribute the lack of a stimulation of photosynthesis to nutrient limitations. Rather than increases in C assimilation, the observed increases in aboveground growth at our study sites are more likely due to shifts in C allocation.     

Nutrient limitation in soils exhibiting differing nitrogen availabilities: what lies beyond nitrogen saturation?

The nature of nutrient limitation in large areas of temperate forest may be changing due to human activities. As N availability in these forests increases, other nutrients could increasingly constrain productivity and other ecosystem processes. To determine the nature of nutrient limitation (N, P, and Ca) in forest soils exhibiting differing N availability, we conducted three field studies in the Fernow Experimental Forest, West Virginia, USA. The first used a ubiquitous herbaceous species, Viola rotundifolia, to compare indices of N availability to the activity of root-associated phosphomonoesterase (PME) activity at two spatial scales. The second study used fertilized, root in-growth cores to assess the extent of N, P, and Ca limitation. Finally, we measured the root-associated PME activity of V. rotundifolia growing in experimental plots that have received various combinations of nutrient additions and harvest treatments. For entire watersheds, stream water nitrate concentrations were positively related to PME activities (R2 = 0.986). For small plots, PME activities were positively associated with soil nitrate availability (R2 = 0.425), and to a lesser extent with the leaf N concentrations (R2 = 0.291). Root growth into microsites fertilized with P was greater than growth into microsites fertilized with either N or Ca, especially in watersheds with high N availability. Experimental additions of N increased the root-associated PME activity of V. rotundifolia, supporting the causality of the relationship between N availability and PME activity. Collectively, our results indicate that, as N availability increases, P becomes increasingly limiting at the sites examined. Understanding how nutrient limitations change during N saturation should improve ecosystem models and better inform our attempts to mitigate any undesired effects.     

二氧化碳浓度升高对植物入侵的影响

从入侵植物和入侵植物群落两个方面,综述了大气二氧化碳浓度升高对植物入侵的影响。二氧化碳浓度升高,可以增加C3植物的入侵性,提高入侵植物的生物量、资源利用率以及繁殖能力,直接影响植物入侵;还可以通过改变土壤水分、氮循环、干扰体系等其它环境因子间接地影响植物入侵。此外,二氧化碳浓度升高,对入侵群落的初级生产量、组成与结构以及群落动态产生重要影响,改变群落的可入侵性。今后应当着重从群落水平,结合其它全球变化因子的共同作用研究二氧化碳浓度升高对植物入侵的影响,同时深入探讨其作用机制以及不同植物类群对二氧化碳的响应,为入侵种的预防和控制提供理论指导。     

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.     

大气CO_2浓度升高对稻田土壤Ca、Mg含量的影响

为了研究稻田生态系统中土壤钙、镁元素生物地球化学循环对大气CO2浓度升高的响应。利用中国稻/麦轮作FACE（Free Air Carbon-dioxide Enrichment）试验平台,研究大气CO2浓度升高（比周围大气高200μmol mol-1）对2007年稻季各生育期不同深度土壤溶液ρ（Ca）、ρ（Mg）的影响。结果表明,大气CO2浓度升高降低了5 cm处土壤溶液ρ（Ca）、ρ（Mg）,有增加稻季30、60和90 cm处土壤溶液ρ（Ca）的趋势,其增幅分别为18.3%、12.4%和15.3%;大气CO2浓度升高会增加稻季Ca淋溶损失风险;稻季不同深度土壤溶液ρ（Ca）、ρ（Mg）对大气CO2浓度升高的响应有所不同。稻田生态系统不同深度土壤Ca、Mg循环对大气CO2浓度升高的响应值得深入研究。     

Soil responses to management, increased precipitation, and added nitrogen in ponderosa pine forests.

Forest management, climatic change, and atmospheric N deposition can affect soil biogeochemistry, but their combined effects are not well understood. We examined the effects of water and N amendments and forest thinning and burning on soil N pools and fluxes in ponderosa pine forests near Flagstaff, Arizona (USA). Using a 15N-depleted fertilizer, we also documented the distribution of added N into soil N pools. Because thinning and burning can increase soil water content and N availability, we hypothesized that these changes would alleviate water and N limitation of soil processes, causing smaller responses to added N and water in the restored stand. We found little support for this hypothesis. Responses of fine root biomass, potential net N mineralization, and the soil microbial N to water and N amendments were mostly unaffected by stand management. Most of the soil processes we examined were limited by N and water, and the increased N and soil water availability caused by forest restoration was insufficient to alleviate these limitations. For example, N addition caused a larger increase in potential net nitrification in the restored stand, and at a given level of soil N availability, N addition had a larger effect on soil microbial N in the restored stand. Possibly, forest restoration increased the availability of some other limiting resource, amplifying responses to added N and water. Tracer N recoveries in roots and in the forest floor were lower in the restored stand. Natural abundance delta15N of labile soil N pools were higher in the restored stand, consistent with a more open N cycle. We conclude that thinning and burning open up the N cycle, at least in the short-term, and that these changes are amplified by enhanced precipitation and N additions. Our results suggest that thinning and burning in ponderosa pine forests will not increase their resistance to changes in soil N dynamics resulting from increased atmospheric N deposition or increased precipitation due to climatic change. Restoration plans should consider the potential impact on long-term forest productivity of greater N losses from a more open N cycle, especially during the period immediately after thinning and burning.     

菌根真菌对大气CO2浓度升高的响应研究进展

大气CO2浓度升高对植物的光合作用、呼吸作用等产生直接影响，进而影响到运送到根系中碳的量，菌根真菌也随之受到影响．本文对全球CO2浓度升高对菌根真菌的影响、菌根真菌在植物对大气CO2增加响应中的作用、菌根真菌在大气CO2浓度增加条件下对整个生态系统的作用等进行了综述，同时对当前存在的问题和未来的发展做了探讨．图1参37     

Atmospheric deposition may affect northern hardwood forest composition by altering soil nutrient supply.

In the northeastern United States, the input of reactive nitrogen (N) via atmospheric deposition has increased rapidly since the onset of the industrial revolution. During the same period of time, acid precipitation and forest harvest have removed substantial quantities of base cations from soil. Because of the dominance of base-poor soils and the low rates of atmospheric base cation deposition, soils throughout the northeastern United States may be increasingly rich in N but poor in calcium (Ca). We studied the consequences of a change in soil N and Ca availability on forest composition by transplanting seedlings of four tree species into replicate plots in the understory and in canopy gaps amended with N and Ca in factorial combination. In this paper, we report on the growth and survivorship of seedlings over a four-year period. Relative to control plots, fertilization with N increased red maple growth by an average of 39% whereas fertilization with Ca decreased survivorship in the understory by 41%. In sugar maple, fertilization with Ca increased growth by 232% and 46% in the forest understory and in canopy gaps, respectively, and significantly increased high light survivorship. Fertilization with N decreased white pine survivorship by 69% in the understory whereas high Ca availability significantly increased survivorship. Fertilization with N or Ca alone reduced red oak growth but had no effect on survivorship. The results of this study suggest that historical losses of soil Ca and the continuing effects of atmospheric-N deposition on N availability are likely to alter the composition of northeastern North American forests because of the positive effects of N enrichment on the growth of red maple and the negative effects of Ca loss on the growth and survivorship of sugar maple and white pine.     

Potential nitrogen constraints on soil carbon sequestration under low and elevated atmospheric CO2

The interaction between nitrogen cycling and carbon sequestration is critical in predicting the consequences of anthropogenic increases in atmospheric CO2 (hereafter, Ca). The progressive N limitation (PNL) theory predicts that carbon sequestration in plants and soils with rising Ca may be constrained by the availability of nitrogen in many ecosystems. Here we report on the interaction between C and N dynamics during a four-year field experiment in which an intact C3/C4 grassland was exposed to a gradient in Ca from 200 to 560 micromol/mol. There were strong species effects on decomposition dynamics, with C loss positively correlated and N mineralization negatively correlated with Ca for litter of the C3 forb Solanum dimidiatum, whereas decomposition of litter from the C4 grass Bothriochloa ischaemum was unresponsive to Ca. Both soil microbial biomass and soil respiration rates exhibited a nonlinear response to Ca, reaching a maximum at approximately 440 micromol/mol Ca. We found a general movement of N out of soil organic matter and into aboveground plant biomass with increased Ca. Within soils we found evidence of C loss from recalcitrant soil C fractions with narrow C:N ratios to more labile soil fractions with broader C:N ratios, potentially due to decreases in N availability. The observed reallocation of N from soil to plants over the last three years of the experiment supports the PNL theory that reductions in N availability with rising Ca could initially be overcome by a transfer of N from low C:N ratio fractions to those with higher C:N ratios. Although the transfer of N allowed plant production to increase with increasing Ca, there was no net soil C sequestration at elevated Ca, presumably because relatively stable C is being decomposed to meet microbial and plant N requirements. Ultimately, if the C gained by increased plant production is rapidly lost through decomposition, the shift in N from older soil organic matter to rapidly decomposing plant tissue may limit net C sequestration with increased plant production.     

Dynamics of nitrogen and dissolved organic carbon at the Hubbard brook experimental forest

The factors controlling spatial and temporal patterns in soil solution and streamwater chemistry are highly uncertain in northern hardwood forest ecosystems in the northeastern United States, where concentrations of reactive nitrogen (Nr) in streams have surprisingly declined over recent decades in the face of persistent high rates of atmospheric Nr deposition and aging forests. Reactive nitrogen includes inorganic species (e.g., ammonium [NH4+], nitrate [NO3-]) and some organic forms (e.g., amino acids) available to support the growth of plants and microbes. The objective of this study was to examine controls on the spatial and temporal patterns in the concentrations and fluxes of nitrogen (N) species and dissolved organic carbon (DOC) in a 12-year record of soil solutions and streamwater along an elevational gradient (540-800 m) of a forested watershed at the Hubbard Brook Experimental Forest (HBEF) in the White Mountains of New Hampshire, USA. Dissolved organic N and DOC concentrations were elevated in the high-elevation spruce-fir-white birch (SFB) zone of the watershed, while NO3- was the dominant N species in the lower elevation hardwood portion of the watershed. Within the soil profile, N retention was centered in the mineral horizon, and significant amounts of N were retained between the lower mineral soil and the stream, supporting the idea that near- and in-stream processes are significant sinks for N at the HBEF. Temporal analysis suggested that hydrologic flow paths can override both abiotic and biotic retention mechanisms (i.e., during the non-growing season when most hydrologic export occurs, or during years with high rainfall), there appears to be direct flushing of N from the organic horizons into the stream via horizontal flow. Significant correlations between soil NO3- concentrations, nitrification rates and streamwater NO3- exports show the importance of biological production as a regulator of inorganic N export. The lack of internal production response (e.g., mineralization, nitrification) to a severe ice storm in 1998 reinforces the idea that plant uptake is the dominant regulator of export response to disturbance.