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.     

Modeling the forest transition: forest scarcity and ecosystem service hypotheses.

An historical generalization about forest cover change in which rapid deforestation gives way over time to forest restoration is called "the forest transition." Prior research on the forest transition leaves three important questions unanswered: (1) How does forest loss influence an individual landowner's incentives to reforest? (2) How does the forest recovery rate affect the likelihood of forest transition? (3) What happens after the forest transition occurs? The purpose of this paper is to develop a minimum model of the forest transition to answer these questions. We assume that deforestation caused by landowners' decisions and forest regeneration initiated by agricultural abandonment have aggregated effects that characterize entire landscapes. These effects include feedback mechanisms called the "forest scarcity" and "ecosystem service" hypotheses. In the forest scarcity hypothesis, forest losses make forest products scarcer, which increases the economic value of forests. In the ecosystem service hypothesis, the environmental degradation that accompanies the loss of forests causes the value of ecosystem services provided by forests to decline. We examined the impact of each mechanism on the likelihood of forest transition through an investigation of the equilibrium and stability of landscape dynamics. We found that the forest transition occurs only when landowners employ a low rate of future discounting. After the forest transition, regenerated forests are protected in a sustainable way if forests regenerate slowly. When forests regenerate rapidly, the forest scarcity hypothesis expects instability in which cycles of large-scale deforestation followed by forest regeneration repeatedly characterize the landscape. In contrast, the ecosystem service hypothesis predicts a catastrophic shift from a forested to an abandoned landscape when the amount of deforestation exceeds the critical level, which can lead to a resource degrading poverty trap. These findings imply that incentives for forest conservation seem stronger in settings where forests regenerate slowly as well as when decision makers value the future.     

CO2 fluxes near a forest edge: a numerical study

In contrast with recent advances on the dynamics of the flow at a forest edge, few studies have considered its role on scalar transport and, in particular, on CO2 transfer. The present study addresses the influence of the abrupt roughness change on forest atmosphere CO2 exchange and contrasts the concentration and flux fields against those of a uniform forested surface. We use an atmospheric boundary layer two-equation closure model that accounts for the flow dynamics and vertical divergence of CO2 sources/sinks within a plant canopy. This paper characterizes the spatial variation of CO2 fluxes as a function of both sources/sinks distribution and the vertical structure of the canopy. Results suggest that the ground source plays a major role in the formation of wave-like vertical CO2 flux behavior downwind of a forest edge, despite the fact that the contribution of foliage sources/sinks changes monotonously. Such a variation is caused by scalar advection in the trunk space and reveals itself as a decrease or increase in vertical fluxes over the forest relative to carbon dioxide exchange of the underlying forest. The effect was more pronounced in model forests where the leaf area is concentrated in the upper part of the canopy. These results can be useful both for interpretation of existing measurements of net ecosystem exchange of CO2 (NEE) from flux towers in limited fetch conditions and in planning future CO2 transport experiments.     

Potentials and limitations of using large-scale forest inventory data for evaluating forest succession models

Forest gap models have been applied widely to examine forest development under natural conditions and to investigate the effect of climate change on forest succession. Due to the complexity and parameter requirements of such models a rigorous evaluation is required to build confidence in the simulation results. However, appropriate data for model assessment are scarce at the large spatial and temporal scales of successional dynamics. In this study, we explore a data source for the evaluation of forest gap models that has been used only little in the past, i.e., large-scale National Forest Inventory data. The key objectives of this study were (a) to examine the potentials and limitations of using large-scale forest inventory data for evaluating the performance of forest gap models and (b) to test two particular models as case studies to derive recommendations for their future improvement.     

Modelling of matter cycle in a mesotrophic bog ecosystem II. Dynamic model and ecological succession

Based upon the the results of static environ analysis of the organic matter cycle through the ecosystem, a number of dynamic models are developed, in this second part, for the matter and nitrogen cycles. Probable outcomes of ecological succession are obtained within the framework of the dynamic model without using the ergodic hypothesis that is implicitly adopted in ecological studies. The dynamics of both stocks in the ecosytem components and flows among them can be observed within the model in order to compare some turnover characteristics of the succession series with those of the ecological one.     

Rapid ecosystem shifts in peatlands: linking plant physiology and succession

Stratigraphic records from peatlands suggest that the shift from a rich fen (calcareous fen) to an ombrotrophic bog can occur rapidly. This shift constitutes a switch from a species-rich ecosystem to a species-poor one with greater carbon storage. In this process, the invasion and expansion of acidifying bog species of Sphagnum (peat mosses) play a key role. To test under what conditions an acidifying bog species could invade a rich fen, we conducted three experiments, contrasting the bog species S. fucsum with the rich-fen species S. warnstorfii and S. teres. We first tested the effect of calcareous water by growing the three species at different constant height above the water table (HWT; 2, 7, and 14 cm) in a rich-fen pool and measured maximum photosynthetic rate and production and difference in length growth as an indicator of competition. In none of the species was the photosynthetic capacity negatively affected when placed at low HWT, but S. fuscum was a weaker competitor at low HWT. In our second experiment we transplanted the three species into microhabitats with different and naturally varying HWT in a rich fen. Here, S. fuscum nearly ceased to photosynthesize when transplanted to low HWT (brown moss carpet), while it performed similarly to the two rich-fen species at the intermediate level (S. warnstorfii hummock level). In contrast to S. fuscum, the rich-fen sphagna performed equally well in both habitats. The brown moss carpet was seasonally flooded, and in our third experiment we found that S. fuscum, but not S. teres, was severely damaged when submerged in rich-fen water. Our results suggest two thresholds in HWT affecting the ecosystem switch: one level that reduces the risk of submergence and a higher one that makes bog sphagna competitive against the rich-fen species.     

Simulating forest succession after blowdown events: The crucial role of space for a realistic management

Ecological patterns vary in space and time. Therefore, when using dynamic models in ecology, the spatial aspect should not be neglected prematurely since it could possibly change the model outcomes to a considerable extent. In view of this problem, we describe here a method how to construct a non-spatial version from a spatially explicit simulation model. The principle idea is to suppress the spatial correlations of cells in a grid in time by continuously re-assigning a random neighbourhood for each cell on the grid. Since this procedure actually eliminates the spatial dimensions, it allows to quantify the unadulterated impact of spatial processes on the model results. To illustrate an important application of this approach in the context of forest management we use a grid-based model that simulates succession of Norway spruce (Picea abies (L.) Karst.) at mountainous sites after blowdown events. The output of this model is compared with the results of the deduced non-spatial version of this model regarding the predicted amount of re-growing trees. The non-spatial version dramatically overestimates the number of spruce trees on different microsites. Thus, the uncritical use of the non-spatial model might give reason to wrong management decisions that are based on too optimistic predictions. In practice, this may lead to dangerous situations, especially in mountain forests serving as protection against avalanches and landslides. This example demonstrates the successful applicability of our approach. Our method can be interpreted as a contribution to an extended sensitivity analysis: it analyses the sensitivity of the results due to structural changes of the model. This sensitivity allows one to estimate the redundancy or the necessity of spatially explicit processes in a model with regard to the parsimony principle of modelling. Since our approach is not dependent on special features of the simulation model used here, it is assumed to be applicable for other spatial models, too, and can thus be considered of general interest for a diligent model analysis.     

不同土地覆被下岩溶表层系统CO2体积分数研究

对重庆金佛山林地、裸地表层岩溶生态系统CO2体积分数进行了野外监测,揭示了CO2体积分数变化规律,这种变化与土壤温度有密切的关系。林地与裸地各个层次土壤的CO2体积分数与土温呈一致性变化,随着土温的升高或降低而相应的增加或减少。文章进一步揭示了林地植被平抑这种动态效应,而裸地则响应于温度变化;这种不同植被系统下的动态差异在解释岩溶沉积记录和讨论岩溶作用与碳循环的关系时值得充分注意。     

Establishment, persistence, and introgression of entomopathogenic nematodes in a forest ecosystem.

Entomopathogenic nematodes (EPN) are currently marketed worldwide for use in inundative biological control, where the applied natural enemy population (rather than its offspring) is expected to reduce insect numbers. Unlike classical biological control, in inundative control natural enemy establishment is not crucial in order to achieve pest suppression. Field trials in Irish forestry provided the opportunity to test predictions regarding the establishment of two exotic (Steinernema carpocapsae and Heterorhabditis megidis) and two indigenous (Steinernema feltiae and Heterorhabditis downesi) species. Nematodes were inundatively applied to pine stumps to control populations of pine weevil, Hylobius abietis, on three clearcut sites, and their persistence and spread monitored for up to five years. All species were recovered three years after application but only S. feltiae was recovered in years 4 and 5. Limited horizontal dispersal to 20 cm (but not 100 cm) was observed, but the majority of nematodes were recovered close to the area of application. Steinernema feltiae was also recovered from nearby stumps to which it had not been applied, indicating possible phoretic dispersal by weevils or other stump-associated fauna. EPN were not recovered from stumps outside the treated area, suggesting that such dispersal is quite localized. Two strains of S. feltiae (Irish and exotic) were applied. Amplified fragment length polymorphism (AFLP) analysis on 11 populations isolated from soil four years later showed that all had a much closer affinity to the applied Irish strain, suggesting persistence of this genotype and extinction of the exotic one. Some strains were clustered close together, and this is interpreted in the light of possible population genetic scenarios. The findings from the field study confirm predictions based on background knowledge of the species and demonstrate the importance of medium-term studies, as a 3-year study would have overestimated the risk of establishment of exotic species. Short-term persistence and spread of S. carpocapsae, S. feltiae, and H. downesi was also studied in pine forest mesocosms. Similar trends to field results, such as limited horizontal dispersal, even vertical distribution, and more abundant recovery of S. feltiae than of other species, point to the utility of mesocosm studies as a predictive tool.     

Net emissions of CH4 and CO2 in Alaska: implications for the region's greenhouse gas budget.

We used a biogeochemistry model, the Terrestrial Ecosystem Model (TEM), to study the net methane (CH4) fluxes between Alaskan ecosystems and the atmosphere. We estimated that the current net emissions of CH4 (emissions minus consumption) from Alaskan soils are approximately 3 Tg CH4/yr. Wet tundra ecosystems are responsible for 75% of the region's net emissions, while dry tundra and upland boreal forests are responsible for 50% and 45% of total consumption over the region, respectively. In response to climate change over the 21st century, our simulations indicated that CH4 emissions from wet soils would be enhanced more than consumption by dry soils of tundra and boreal forests. As a consequence, we projected that net CH4 emissions will almost double by the end of the century in response to high-latitude warming and associated climate changes. When we placed these CH4 emissions in the context of the projected carbon budget (carbon dioxide [CO2] and CH4) for Alaska at the end of the 21st century, we estimated that Alaska will be a net source of greenhouse gases to the atmosphere of 69 Tg CO2 equivalents/yr, that is, a balance between net methane emissions of 131 Tg CO2 equivalents/yr and carbon sequestration of 17 Tg C/yr (62 Tg CO2 equivalents/yr).     

开放式空气CO2浓度升高对水稻根系形态的影响

在FACE（free-air carbon dioxide enrichment）技术平台上，采用水培的研究方法，观测了大气CO2浓度升高和两种氮水平下水稻根系形态的变化。结果表明，在水稻各生育期，CO2浓度升高都极显著增加了根干质量，且主要增加于根粗为2．0～2．5mm／n的部位。根系形态的各项指标均对高CO2浓度有积极的响应，在抽穗期尤为明显；N处理的差异很明显，低氮条件下根系表现为根长、根尖数和根表面积增加，常氮条件下根粗和发根数增加。各生育期的根冠比在高CO2浓度下极显著增加，尤其在LN处理下。水稻从分蘖期到抽穗期，因地上部分的增幅大，根冠比表现为逐渐降低的趋势。     

Nocturnal accumulation of CO2 underneath a tropical forest canopy along a topographical gradient

Flux measurements of carbon dioxide and water vapor above tropical rain forests are often difficult to interpret because the terrain is usually complex. This complexity induces heterogeneity in the surface but also affects lateral movement of carbon dioxide (CO2) not readily detected by the eddy covariance systems. This study describes such variability using measurements of CO2 along vertical profiles and along a toposequence in a tropical rain forest near Manaus, Brazil. Seasonal and diurnal variation was recorded, with atmospheric CO2 concentration maxima around dawn, generally higher CO2 build-up in the dry season and stronger daytime CO2 drawdown in the wet season. This variation was reflected all along the toposequence, but the slope and valley bottom accumulated clearly more CO2 than the plateaus, depending on atmospheric stability. Particularly during stable nights, accumulation was along lines of equal altitude, suggesting that large amounts of CO2 are stored in the valleys of the landscape. Flushing of this store only occurs during mid-morning, when stored CO2 may well be partly transported back to the plateaus. It is clear that, for proper interpretation of tower fluxes in such complex and actively respiring terrain, the horizontal variability of storage needs to be taken into account not only during the night but also during the mornings.     

A simulation model for organic phosphorus transformation in a forest soil ecosystem

A numerical model which simulates the decomposition of litter and mineralization and immobilization of P in the humus layer of a temperate forest (beech site of Solling) is described. The model takes into account the effect of moisture, temperature and C/N ratio. The simulated concentration of P in the effluent of the humus layer agrees well with the measured values. The model predicts an increase in the C/P ratio of the unde-composed litter with time and that there is no direct mineralization of P from litter without passing through a microbial body. The net rate of mineralization is, however, always positive with its highest peak in July. Maximum immobilization of P from solution occurs in June and the minimum in January.The model is stable against changes in the litter input, its C/P ratio and other initial conditions, but it is very sensitive to changes in the efficiency factor which represents the fraction of decomposed C incorporated into microbial tissue. This is a site-specific model but can be used for grassland or agricultural systems with changes in certain parameters.     

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.     

Microbial stress-response physiology and its implications for ecosystem function

Microorganisms have a variety of evolutionary adaptations and physiological acclimation mechanisms that allow them to survive and remain active in the face of environmental stress. Physiological responses to stress have costs at the organismal level that can result in altered ecosystem-level C, energy, and nutrient flows. These large-scale impacts result from direct effects on active microbes' physiology and by controlling the composition of the active microbial community. We first consider some general aspects of how microbes experience environmental stresses and how they respond to them. We then discuss the impacts of two important ecosystem-level stressors, drought and freezing, on microbial physiology and community composition. Even when microbial community response to stress is limited, the physiological costs imposed on soil microbes are large enough that they may cause large shifts in the allocation and fate of C and N. For example, for microbes to synthesize the osmolytes they need to survive a single drought episode they may consume up to 5% of total annual net primary production in grassland ecosystems, while acclimating to freezing conditions switches Arctic tundra soils from immobilizing N during the growing season to mineralizing it during the winter. We suggest that more effectively integrating microbial ecology into ecosystem ecology will require a more complete integration of microbial physiological ecology, population biology, and process ecology.     

Advection and resulting CO2 exchange uncertainty in a tall forest in central Germany

Potential losses by advection were estimated at Hainich Forest, Thuringia, Germany, where the tower is located at a gentle slope. Three approaches were used: (1) comparing nighttime eddy covariance fluxes to an independent value of total ecosystem respiration by bottom-up modeling of the underlying processes, (2) direct measurements of a horizontal CO2 gradient and horizontal wind speed at 2 m height in order to calculate horizontal advection, and (3) direct measurements of a vertical CO2 gradient and a three-dimensional wind profile in order to calculate vertical advection. In the first approach, nighttime eddy covariance measurements were compared to independent values of total ecosystem respiration by means of bottom-up modeling of the underlying biological processes. Turbulent fluxes and storage term were normalized to the fluxes calculated by the bottom-up model. Below a u(*) threshold of 0.6 m/s the normalized turbulent fluxes decreased with decreasing u(*), but the flux to the storage increased only up to values less than 20% of the modeled flux at low turbulence. Horizontal advection was measured by a horizontal CO2 gradient over a distance of 130 m combined with horizontal wind speed measurements. Horizontal advection occurred at most of the evenings independently of friction velocity above the canopy. Nevertheless, horizontal advection was higher when u(*) was low. The peaks of horizontal advection correlated with changes in temperature. A full mass balance including turbulent fluxes, storage, and horizontal and vertical advection resulted in an increase of spikes and scatter but seemed to generally improve the results from the flux measurements. The comparison of flux data with independent bottom-up modeling results as well as the direct measurements resulted in strong indications that katabatic flows along the hill slope during evening and night reduces the measured apparent ecosystem respiration rate. In addition, anabatic flows may occur during the morning. We conclude that direct measurements of horizontal and vertical advection are highly necessary at sites located even on gentle hill slopes.     

An analysis of joint costs in a managed forest ecosystem

This paper analyzes the tenability of line item budgeting and independently determined output costs for a managed forest ecosystem. The general problem of cost allocation in a joint production system is discussed as one of choosing paths of integration. A production structure is hypothesized for describing a managed forest ecosystem and its characteristics of “jointness” are discussed. Finally, an empirical case example is presented which indicates that cost estimates and associated means of production which result from single output costing procedures and from joint costing procedures may be significantly different in a managed forest ecosystem.     

Modeling patterns of nonlinearity in ecosystem responses to temperature, CO2, and precipitation changes.

It is commonly acknowledged that ecosystem responses to global climate change are nonlinear. However, patterns of the nonlinearity have not been well characterized on ecosystem carbon and water processes. We used a terrestrial ecosystem (TECO) model to examine nonlinear patterns of ecosystem responses to changes in temperature, CO2, and precipitation individually or in combination. The TECO model was calibrated against experimental data obtained from a grassland ecosystem in the central United States and ran for 100 years with gradual change at 252 different scenarios. We primarily used the 100th-year results to explore nonlinearity of ecosystem responses. Variables examined in this study are net primary production (NPP), heterotrophic respiration (R(h)), net ecosystem carbon exchange (NEE), runoff, and evapotranspiration (ET). Our modeling results show that nonlinear patterns were parabolic, asymptotic, and threshold-like in response to temperature, CO2, and precipitation anomalies, respectively, for NPP, NEE, and R(h). Runoff and ET exhibited threshold-like pattern in response to both temperature and precipitation anomalies but were less sensitive to CO2 changes. Ecosystem responses to combined temperature, CO2, and precipitation anomalies differed considerably from the responses to individual factors in terms of response patterns and/or critical points of nonlinearity. Our results suggest that nonlinear patterns in response to multiple global-change factors were diverse and were considerably affected by combined climate anomalies on ecosystem carbon and water processes. The diverse response patterns in nonlinearity have profound implications for both experimental design and theoretical development.     

Development of Lake Ladoga ecosystem models: modeling of the phytoplankton succession in the eutrophication process. I

New models of Lake Ladoga ecosystem and the results of modeling are presented. In the first part the model of phytoplankton succession in the process of anthropogenic eutrophication of the lake is considered under the evolution of the phosphorus loading. The still continued anthropogenic eutrophication of the lake started in 1962 when the phosphorus load began to increase. Since 1962 during the evolution of the lake’s state from oligotrophic to developed mezotrophic one, the structure of phytoplankton community dominating species was significantly changed as well as its total productivity. The system state in the model is described by 14 parameters: nine phytoplankton complexes, zooplankton, dissolved organic matter, detritus, dissolved mineral phosphorus and dissolved oxygen. The number of parameters of this model is noticeably larger than that of previous models created by the authors. The relative dynamics of phytoplankton complexes in the lake’s ecosystem evolution was simulated by the new model. It is shown that the modeling results are adequately corresponding to the observation data. The results of phytoplankton structure modeling allow to estimate the impact of phytoplankton on the water quality as well as give the prediction of the lake’s ecosystem evolution with the changes of the phosphorus loading.