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.     

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 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.     

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.     

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.     

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.     

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.     

氮沉降下西南亚高山针叶林生长的养分限制特征

长期氮(N)沉降及其诱导的N、磷(P)养分平衡性对森林生长与生产力的生态反馈效应已成为当前森林生态学研究的前沿与热点,但目前大多研究主要基于已有文献数据整合分析,而缺乏野外原位系统性研究与试验证据.以西南亚高山两种典型人工针叶林——云杉(Picea asperata)林和华山松(Pinus armandii)林为对象,...     

不同林龄马尾松（Pinus massoniana）人工林碳氮磷分配格局及化学计量特征

为了解长江上游低山丘陵区马尾松（Pinus massoniana）人工林生态系统的C、N、P分配格局及化学计量特征,本文采用时空互代的方法,在宜宾高县来复林区选取三种不同林龄（5年生幼龄林、14年生中龄林、39年生成熟林）,但立地条件相近、样地情况基本一致的马尾松（Pinus massoniana）人工林作为研究对象,对马尾松针叶、凋落物及土壤中的C、N、P含量及 w（C）？w（N）？w（P）化学计量特征进行测定和分析。结果表明,（1）C、N、P 含量均表现为针叶〉凋落物〉土壤,且在三个库之间差异显著;（2）林龄对针叶、凋落物、土壤的 C、N、P 及 w（C）？w（N）、w（C）？w（P）计量比均有显著影响。（3）土壤 C、N、P含量在成熟林中最高;针叶和凋落物的C含量在成熟林中最低,N、P含量则在中龄林中最高。（4）随林龄增加马尾松对N、P的利用效率降低,针叶、凋落物及土壤的w（C）？w（N）与 w（C）？w（P）均表现为下降。（5）马尾松针叶w（N）？w（P）比值在14.37~15.53之间,说明该地区马尾松人工林受N和P的共同限制,但林龄对N、P养分限制的影响不显著。为提高该区马尾松人工林的生产力,建议在人工林的抚育管理中要适当增加N肥和P肥,同时也可在马尾松人工林引入豆科固氮植物以提高地力。该研究将马尾松针叶、凋落物及土壤结合起来探究随林龄增长C、N、P养分元素的分配格局及化学计量特征的变化,有助于全面、系统地揭示马尾松人工林生态系统的养分循环,对指导马尾松人工林生产,调节和改善林木生长环境,提高系统的养分利用效率及林地生产力具有重要意义。     

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。     

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.     

Fate of nitrogen in riparian forest soils and trees: an 15N tracer study simulating salmon decay

We introduced an 15N-NH4+ tracer to the riparian forest of a salmon-bearing stream (Kennedy Creek, Washington, USA) to quantify the cycling and fate of a late-season pulse of salmon N and, ultimately, mechanisms regulating potential links between salmon abundance and tree growth. The 15N tracer simulated deposition of 7.25 kg of salmon (fresh) to four 50-m2 plots. We added NH4+ (the initial product of salmon carcass decay) and other important nutrients provided by carcasses (P, S, K, Mg, Ca) to soils in late October 2003, coincident with local salmon spawning. We followed the 15N tracer through soil and tree pools for one year. Biological uptake of the 15N tracer occurred quickly: 64% of the 15N tracer was bound in soil microbiota within 14 days, and roots of the dominant riparian tree, western red cedar (Thuja plicata), began to take up 15N tracer within seven days. Root uptake continued through the winter. The 15N tracer content of soil organic matter reached a maximum of approximately 52%, five weeks after the application, and a relative equilibrium of approximately 40% within five months. Six months after the addition, in spring 2004, at least 37% of the 15N tracer was found in tree tissues: approximately 23% in foliage, approximately 11% in roots, and approximately 3% in stems. Within the stems, xylem and phloem sap contained approximately 96% of the tracer N, and approximately 4% was in structural xylem N. After one year, at least 28% of the 15N tracer was still found in trees, and loss from the plots was only approximately 20%. The large portion of tracer N taken up in the fall and reallocated to leaves and stems the following spring provides mechanistic evidence for a one-year-lagged tree-growth response to salmon nutrients. Salmon nutrients have been deposited in the Kennedy Creek system each fall for centuries, but the system shows no evidence of nutrient saturation. Rates of N uptake and retention are a function of site history and disturbance and also may be the result of a legacy effect, in which annual salmon nutrient addition may lead to increased efficiency of nutrient uptake and use.     

氮沉降对森林土壤碳收支机制的影响

森林土壤储存着全球陆地生态系统大约45%的碳,在维持全球碳平衡方面具有重要的作用。不断加剧的全球氮沉降对森林生态系统碳循环和碳吸存产生了深刻的影响,进而改变了森林生态系统的生产力和生物量积累。本文以欧洲和北美温带地区开展的有关氮沉降对森林生态系统影响的研究为基础,提炼出最可能决定加氮影响碳输入、输出效应方向和大小的因素：凋落物分解、细根周转、外生菌根真菌、土壤呼吸及可溶性有机碳淋失,并探讨了森林生态系统碳动态对氮沉降响应的不确定性。陆地生态系统碳氮循环密切相关,由于氮循环的复杂性,尽管以往碳循环研究都考虑了氮对碳循环的限制作用,但在碳氮循环耦合机理方面的研究还比较少见。在未来研究中,应通过探寻森林土壤碳氮相互作用特征,及土壤微生物、土壤酶等与土壤碳氮过程的互动机制,来增进氮沉降对森林碳储量和碳通量的理解。     

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.     

Gastropod herbivory in response to elevated CO2 and N addition impacts plant community composition

In this study, the influence of elevated carbon dioxide (CO2) and nitrogen (N) deposition on gastropod herbivory was investigated for six annual species in a California annual grassland community. These experimentally simulated global changes increased availability of important resources for plant growth, leading to the hypothesis that species with the most positive growth and foliar nutrient responses would experience the greatest increase in herbivory. Counter to the expectations, shifts in tissue N and growth rates caused by N deposition did not predict shifts in herbivore consumption rates. N deposition increased seedling N concentrations and growth rates but did not increase herbivore consumption overall, or for any individual species. Elevated CO2 did not influence growth rates nor have a statistically significant influence on seedling N concentrations. Elevated CO2 at ambient N levels caused a decline in the number of seedlings consumed, but the interaction between CO2 and N addition differed among species. The results of this study indicate that shifting patterns of herbivory will likely influence species composition as environmental conditions change in the future; however, a simple trade-off between shifting growth rates and palatability is not evident.     

岷江干旱河谷灌木幼苗的土壤养分适应性研究

岷江上游干旱河谷地区的植被恢复是目前急需解决的问题,小马鞍羊蹄甲（Bauhinia faberi var. microphylla）是干旱河谷地区的优势乡土灌木,并作为当地植被恢复过程中的重要备选物种,因而进行小马鞍羊蹄甲种群在土壤贫瘠的干旱河谷的养分限制研究十分必要。针对性地进行施肥将影响到小马鞍羊蹄甲幼苗存活、生长和定殖,而如何有效提高该物种的存活和生长速率对于干旱河谷植被恢复将具有重要的意义。采用野外调查与模拟实验相结合的方法,分别研究了干旱河谷地区3个典型群落（干旱河谷的核心区——飞虹,干旱河谷灌丛与亚高山森林的过渡区——北部的石大关、干旱河谷灌丛与温性森林的过渡区——南部的蓝新镇）中小马鞍羊蹄甲幼苗叶片的化学计量特征,和施肥试验[w（有效氮）分别为100、280、460 mg·kg–1、w（有效磷）分别为12、24、48 mg·kg–1、w（氮）分别为40、70、100 mg·kg–1、w（磷）分别为12和24 mg·kg–1]中叶片化学计量特征及幼苗生长参数（叶片数、基径和株高）和各器官（根茎叶）生物量。结果表明：在野外各演替阶段的小马鞍羊蹄甲幼苗生物量和营养元素质量分数都随着磷肥的增加而增加,表明磷素是植物生长的限制因子,同时N∶P比均大于16,也暗示其受到P养分的限制;在室内施肥试验中,施加N肥没有促进小马鞍羊蹄甲生物量的积累,反而抑制了幼苗生长;施加P肥促进了幼苗生物量的积累,表明幼苗缺乏P元素,养分限制类型为P限制。     

Dynamics and physiology of saxitoxin production by the dinoflagellatesAlexandrium spp.

Toxin content (fmol cell^–1) and a suite of elemental and macromolecular variables were measured in batch cultures of the dinoflagellatesAlexandrium fundyense, A. tamarense andAlexandrium sp. from the southern New England region, USA. A different perspective was provided by semicontinuous cultures which revealed sustained, steady-state physiological adaptations by cells to N and P limitation. Two types of variability were investigated. In batch culture, changes in nutrient availability with time caused growth stage variability in toxin content, which often peaked in mid-exponential growth. A second type of variability that could be superimposed on growth stage differences is best exemplified by the high toxin content of cells grown at suboptimal temperatures. Calculations of the net rate of toxin production (R _tox ; fmol cell^–1 d^–1) for these different culture treatments and modes made it possible to separate the dynamics of toxin production from cell division. Over a wide range of growth rates, cells produced toxin at rates approximating those needed to replace losses to daughter cells during division. The exception to this direct proportionality was with P limitation, which was associated with a dramatic increase in the rate of toxin production as cells stopped dividing due to nutrient limitation in batch culture. Growth stage variability in batch culture thus reflects small imbalances (generally within a factor of two) between the specific rates of toxin production and cell division. N limitation and CO₂ depletion both affect pathways involved in toxin synthesis before those needed for cell division; P limitation does the opposite. The patterns of toxin accumulation were the same as for major cellular metabolites or elemental pools. The highest rates of toxin production appear to result from an increased availability of arginine (Arg) within the cell, due to either a lack of competition for this amino acid from pathways involved in cell division or to increased de novo synthesis. There were no significant changes in toxin content with either acclimated growth at elevated salinity, or with short term increases or decreases of salinity. These results demonstrate that toxin production is a complex process which, under some conditions, is closely coupled to growth rate; under other conditions, these processes are completely uncoupled. Explanations for the observed variability probably relate to pool sizes of important metabolites and to the differential response of key biochemical reactions to these pool sizes and to environmental conditions.     

森林土地利用变化及其对碳循环的影响

由于人口剧增，人类活动的影响不断加大，在过去100年全球土地利用／土地覆被发生了巨大的变化。最常见的土地利用变化是由森林转变为农业用地。森林砍伐使森林生态系统地上部生物量大大减少，砍伐后作农业用地，降低了植被生产力，减少了土壤有机质的输入，增强了腐殖质的矿化作用，有机质分解速率增加，有机碳贮量随之降低，从而影响到森林生态系统的碳循环，使大量碳元素释放到大气中，引起温室效应，导致全球变暖。另一个常见的土地利用变化是植树造林和森林恢复，这一过程可以增加森林生态系统的碳储量，从而减缓大气CO2体积分数的上升。     

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.