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.     

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.     

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.     

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.     

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.     

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.     

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。     

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.     

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.     

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.     

Nitrogen controls on ecosystem carbon sequestration: a model implementation and application to Saskatchewan,Canada

A plant–soil nitrogen (N) cycling model was developed and incorporated into the Integrated BIosphere Simulator (IBIS) of Foley et al. [Foley, J.A., Prentice, I.C., Ramankutty, N., Levis, S., Pollard, D., Sitch, S., Haxeltine, A., 1996. An integrated biosphere model of land surface process, terrestrial carbon balance and vegetation dynamics. Global Biogeochem. Cycles 10, 603–628]. In the N-model, soil mineral N regulates ecosystem carbon (C) fluxes and ecosystem C:N ratios. Net primary productivity (NPP) is controlled by feedbacks from both leaf C:N and soil mineral N. Leaf C:N determines the foliar and canopy photosynthesis rates, while soil mineral N determines the N availability for plant growth and the efficiency of biomass construction. Nitrogen controls on the decomposition of soil organic matter (SOM) are implemented through N immobilization and mineralization separately. The model allows greater SOM mineralization at lower mineral N, and conversely, allows greater N immobilization at higher mineral N. The model's seasonal and inter-annual behaviours are demonstrated. A regional simulation for Saskatchewan, Canada, was performed for the period 1851–2000 at a 10 km × 10 km resolution. Simulated NPP was compared with high-resolution (1 km × 1 km) NPP estimated from remote sensing data using the boreal ecosystem productivity simulator (BEPS) [Liu, J., Chen, J.M., Cihlar, J., Park, W.M., 1997. A process-based boreal ecosystem productivity simulator using remote sensing inputs. Remote Sens. Environ. 44, 81–87]. The agreement between IBIS and BEPS, particularly in NPP spatial variation, was considerably improved when the N controls were introduced into IBIS.     

闽南山区连续年龄序列桉树人工林土壤养分动态

对闽南山区连续年龄序列(2 a、3 a、4 a、5 a、6 a)的尾巨桉(Eucalyptus urophyllaxE.grandis)人工林土壤有机碳及氮、磷、钾含量进行了分析,以评价土壤养分随林分年龄的变化动态.土壤有机碳、全氮、水解氮、全磷、速效磷均表现出随林分年龄增加而增加,到4 a生时达到最大,随后则呈下降趋势,6 a生时下降到最低值.速效钾含量则随林分年龄增加逐步下降,6 a生林分0～20 cm、20～40 cm和40～60 cm这3个土层速效钾含量分别比2 a生下降了53.2%、44.4%和55.2%;土壤全钾含量也表现出一定的下降趋势.所有年龄林分的土壤有机碳、全氮、水解氮、全磷、速效磷和速效钾含量均随土层深度增加而降低,而土壤伞钾则表现出相反的趋势.因此,在桉树人工林经营过程中,应定期增施钾肥,适时施用有机肥及氮、磷肥,以控制地力衰退.图4表2参24     

干旱区不同种植年限苜蓿人工草地土壤CO2排放的主要影响因素

采用静态箱-气象色谱法对干旱区不同种植年限苜蓿人工草地土壤系统、土壤-植被系统的CO2排放通量进行了测定,并对土壤 CO2排放通量的主要影响因素,包括土壤碳氮质量分数、土壤温度、气温、土壤水分体积分数、生物量进行了测定分析,从而得到了苜蓿生长季内CO2排放通量规律及影响土壤CO2排放的主要因素。结果表明：观测期内各个年限土壤-植被系统与土壤系统的CO2排放趋势基本一致,总体表现为4a〉5a〉8a〉3a〉1a,表现出“夏季高秋季低”的季节变化规律,具有明显的季节变化动态,各样地CO2通量在7月中旬达到最高排放通量;土壤CO2排放通量与0~20 cm土层内土壤有机碳质量分数、活性有机碳质量分数、土壤C：N及土壤全N质量分数都具有正相关关系,并且都达到了显著水平;土壤CO2排放通量与5 cm土壤温度和大气温度都有正相关关系,并且显著水平很高（P＜0.001）;土壤CO2排放通量与0~10 cm土壤水分体积分数之间没有显著的相关性（P〉0.05）;土壤CO2排放通量与地上部分生物量的大小有直接的正相关关系,并且CO2排放通量与地上生物量具有相同的变化趋势。     

菌根类型对森林树木净初级生产力的影响

菌根是土壤真菌与植物根系形成的共生体,存在于绝大多数植物（90%）的根系和生境中。菌根共有7种类型,在生态系统的过程和功能方面都扮演着十分重要的角色。为了增强对菌根在森林生态系统中重要功能的理解,文章基于全球森林数据库,在全球尺度上研究了不同菌根类型对森林树木净初级生产力（NPP）的影响。结果表明,森林树木NPP随菌根类型的不同而不同,AM类型菌根森林的NPP[679.49 g.m-2.a-1（以C计）]要显著高于含ECM类型菌根的森林[479.00 g.m-2.a-1（以C计）];菌根类型的不同对森林树木地上和地下及其各组分NPP的影响和贡献也存在着显著的不同,AM类型菌根对地下NPP的贡献要高于ECM菌根,而ECM菌根对地上NPP的贡献则较大。菌根类型对地上、地下NPP组分的影响分析则表明,AM类型的菌根对树叶和细根NPP的贡献较大,而ECM类型菌根则对树木主干和枝NPP的贡献较大。可见,森林树木总体NPP及其各组分NPP都随着菌根类型的不同而存在显著的差异。     

The role of canopy structural complexity in wood net primary production of a maturing northern deciduous forest

The even-aged northern hardwood forests of the Upper Great Lakes Region are undergoing an ecological transition during which structural and biotic complexity is increasing. Early-successional aspen (Populus spp.) and birch (Betula papyrifera) are senescing at an accelerating rate and are being replaced by middle-successional species including northern red oak (Quercus rubra), red maple (Acer rubrum), and white pine (Pinus strobus). Canopy structural complexity may increase due to forest age, canopy disturbances, and changing species diversity. More structurally complex canopies may enhance carbon (C) sequestration in old forests. We hypothesize that these biotic and structural alterations will result in increased structural complexity of the maturing canopy with implications for forest C uptake. At the University of Michigan Biological Station (UMBS), we combined a decade of observations of net primary productivity (NPP), leaf area index (LAI), site index, canopy tree-species diversity, and stand age with canopy structure measurements made with portable canopy lidar (PCL) in 30 forested plots. We then evaluated the relative impact of stand characteristics on productivity through succession using data collected over a nine-year period. We found that effects of canopy structural complexity on wood NPP (NPPw) were similar in magnitude to the effects of total leaf area and site quality. Furthermore, our results suggest that the effect of stand age on NPPw is mediated primarily through its effect on canopy structural complexity. Stand-level diversity of canopy-tree species was not significantly related to either canopy structure or NPPw. We conclude that increasing canopy structural complexity provides a mechanism for the potential maintenance of productivity in aging forests.     

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.     

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.     

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。     

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.     

CO_2倍增和气候变化对北京山区栓皮栎林NPP影响研究

应用生物地球化学过程模型BIOME-BGC估算了1977—1992年北京妙峰山栓皮栎（Quercus variabilis）林的净第一性生产力（NPP）,并分析气候对NPP年际变化的影响以及未来气候变化情景下对NPP的影响。结果表明：1977—1992年15年间栓皮栎的NPP（以C计）平均值为340.17g·m-2·a-1,NPP（以C计）变化在143.56～431.56g·m-2·a-1之间,并无明显的整体变化趋势,但表现出明显的年际变化,年际变动率达18%。在这段时间内降水量成为控制栓皮栎林NPP年际变化的主要气候因子。通过设置18种不同未来气候方案进行栓皮栎林NPP模拟表明,CO2浓度加倍会降低栓皮栎林的NPP但降低幅度较小。在CO2浓度不变的情况下,温度升高2.0℃和降水的协同增加以及单个因子的增加都有利于NPP的积累,但协同增加不如单个因子的增加对NPP的积累效应明显;在CO2和气候同时改变的情况下,CO2浓度加倍、温度升高2.0℃和降水的协同增加有利于NPP的积累且协同增加比单个因子的增加对NPP的积累效应明显,但各因子之间交互作用较弱。