A highly controversial issue in global biogeochemistry is the regulation of terrestrial carbon (C) sequestration by soil nitrogen (N) availability. This controversy translates into great uncertainty in predicting future global terrestrial C sequestration. We propose a new framework that centers on the concept of progressive N limitation (PNL) for studying the interactions between C and N in terrestrial ecosystems. In PNL, available soil N becomes increasingly limiting as C and N are sequestered in long-lived plant biomass and soil organic matter. Our analysis focuses on the role of PNL in regulating ecosystem responses to rising atmospheric carbon dioxide concentration, but the concept applies to any perturbation that initially causes C and N to accumulate in organic forms. This article examines conditions under which PNL may or may not constrain net primary production and C sequestration in terrestrial ecosystems. While the PNL-centered framework has the potential to explain diverse experimental results and to help researchers integrate models and data, direct tests of the PNL hypothesis remain a great challenge to the research community.

Soil carbon is a large component of the global carbon cycle and its management can significantly affect the atmospheric CO2 concentration. An important management issue is the extent of soil carbon (C) release when forest is converted to agricultural land. We reviewed the literature to assess changes in soil C upon conversion of forests to agricultural land. Analyses are confounded by changes in soil bulk density upon land‐use change, with agricultural soils on average having 13% higher bulk density. Consistent with earlier reviews, we found that conversion of forest to cultivated land led to an average loss of approximately 30% of soil C. When we restricted our analysis to studies that had used appropriate corrections for changes in bulk density, soil C loss was 22%. When, from all the studies compiled, we considered only studies reporting both soil C and nitrogen (N), average losses of C and N were 24% and 15%, respectively, hence showing a decrease in the average C : N ratio. The magnitude of these changes in the C : N ratio did not correlate with either C or N changes. When considering the transition from forest to pasture, there was no significant change in either soil C or N, even though reported changes in soil C ranged from −50% to +160%. Among studies that reported changes in soil N as well as soil C, C : N ratios both increased and decreased, with trends depending on changes in system N. Systems with increasing soil N generally had decreased C : N ratios, whereas systems with decreasing soil N had increased C : N ratios. Our survey confirmed earlier findings that conversion of forest to cropland generally leads to a loss of soil carbon, although the magnitude of change might have been inflated in many studies by the confounding influence of bulk‐density changes. In contrast, conversion of forest to uncultivated grazing land did not, on average, lead to loss of soil carbon, although individual sites may lose or gain soil C, depending on specific circumstances, such as application of fertiliser or retention or removal of plant residues.

Stimulation of terrestrial plant production by rising CO 2 concentration is projected to reduce the airborne fraction of anthropogenic CO 2 emissions. Coupled climate-carbon cycle models are sensitive to this negative feedback on atmospheric CO 2 , but model projections are uncertain because of the expectation that feedbacks through the nitrogen (N) cycle will reduce this so-called CO 2 fertilization effect. We assessed whether N limitation caused a reduced stimulation of net primary productivity (NPP) by elevated atmospheric CO 2 concentration over 11 y in a free-air CO 2 enrichment (FACE) experiment in a deciduous Liquidambar styraciflua (sweetgum) forest stand in Tennessee. During the first 6 y of the experiment, NPP was significantly enhanced in forest plots exposed to 550 ppm CO 2 compared with NPP in plots in current ambient CO 2 , and this was a consistent and sustained response. However, the enhancement of NPP under elevated CO 2 declined from 24% in 2001-2003 to 9% in 2008. Global analyses that assume a sustained CO 2 fertilization effect are no longer supported by this FACE experiment. N budget analysis supports the premise that N availability was limiting to tree growth and declining over time -an expected consequence of stand development, which was exacerbated by elevated CO 2 . Leaf-and stand-level observations provide mechanistic evidence that declining N availability constrained the tree response to elevated CO 2 ; these observations are consistent with stand-level model projections. This FACE experiment provides strong rationale and process understanding for incorporating N limitation and N feedback effects in ecosystem and global models used in climate change assessments.CO 2 fertilization | free air CO 2 enrichment | global carbon cycle | sweetgum | coupled climate-carbon cycle models P olicy decisions to mitigate climate change require dependable predictions of the forcings and feedbacks between the terrestrial biosphere and the climate (1). Currently, climate models that are coupled to terrestrial and oceanic carbon (C)-cycle models simulate a positive feedback to climate change such that the airborne fraction of anthropogenic CO 2 emissions increases with, and amplifies, climatic warming (1). However, the uncertainty in these projections is high, largely because of uncertainty in the offsetting negative feedback that may occur if stimulation of terrestrial plant production by rising CO 2 concentration increases land C storage and thereby reduces the airborne fraction of anthropogenic CO 2 emissions. Coupled climate-C-cycle models, including those used in the Intergovernmental Panel on Climate Change Fourth Assessment Report (AR4) (2), are sensitive to this negative feedback on atmospheric CO 2 (3). For example, dynamic global vegetation models (4) simulate an increased terrestrial C sink resulting from the physiological responses of plants to elevated atmospheric CO 2 concentration (eCO 2 ), and when coupled to climate models, inclusion of the CO 2 fertilization effect slows the increase in...

We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.

Stimulation of terrestrial productivity by rising CO~2~ concentration is projected to reduce the airborne fraction of anthropogenic CO~2~ emissions; coupled climate-carbon (C) cycle models, including those used in the IPCC Fourth Assessment Report (AR4), are sensitive to this negative feedback on atmospheric CO~2~^1^. The representation of the so-called CO~2~ fertilization effect in the 11 models used in AR4 and subsequent models^2,3^ was broadly consistent with experimental evidence from four free-air CO~2~ enrichment (FACE) experiments, which indicated that net primary productivity (NPP) of forests was increased by 23 +/- 2% in response to atmospheric CO~2~ enrichment to 550 ppm^4^. Substantial uncertainty remains, however, because of the expectation that feedbacks through the nitrogen (N) cycle will reduce the CO~2~ stimulation of NPP^5,6^; these feedbacks were not included in the AR4 models and heretofore have not been confirmed by experiments in forests^7^. Here, we provide new evidence from a FACE experiment in a deciduous Liquidambar styraciflua (sweetgum) forest stand in Tennessee, USA, that N limitation has significantly reduced the stimulation of NPP by elevated atmospheric CO~2~ concentration (eCO~2~). Isotopic evidence and N budget analysis support the premise that N availability in this forest ecosystem has been declining over time, and declining faster in eCO~2~. Model analyses and evidence from leaf- and stand-level observations provide mechanistic evidence that declining N availability constrained the tree response to eCO2. These results provide a strong rationale and process understanding for incorporating N limitation and N feedback effects in ecosystem and global models used in climate change assessments.

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Established process-based models of forest biomass production in relation to atmospheric CO"2 concentration (McMurtrie 1991) and soil carbon/nutrient dynamics (Parton et al. 1987) are integrated to derive the @'Generic Decomposition and Yield@' model (G'DAY). The model is used to describe how photosynthesis and nutritional factors interact to determine the productivity of forests growing under nitrogen-limited conditions. A simulated instantaneous doubling of atmospheric CO"2 concentration leads to a growth response that is initially large (27% above productivity at current CO"2) but declines to <10% elevation within 5 yr. The decline occurs because increases in photosynthetic carbon gain at elevated CO"2 are not matched by increases in nutrient supply. Lower foliar N concentrations at elevated CO"2 have two countervailing effects on forest production: decreased rates of N cycling between vegetation and soils (with negative consequences for productivity), and reduced rates of N loss through gaseous emission, fire, and leaching. Theoretical analysis reveals that there is an enduring response to CO"2 enrichment, but that the magnitude of the long-term equilibrium response is extremely sensitive to the assumed rate of gaseous emission resulting from mineralization of nitrogen. Theory developed to analyze G'DAY is applicable to other published production-decomposition models describing the partitioning of soil carbon among compartments with widely differing decay-time constants.

In a forest ecosystem at steady state, net carbon (C) assimilation by plants and C loss through soil and litter decomposition by heterotrophic organisms are balanced. However, a perturbation to the system, such as increased mean soil temperature, will lead to faster decay, enhancing CO 2 release from decomposers, and thus upsetting the balance. Recent in situ experiments have indicated that the stimulation of soil respiration following a step increase in annual average soil temperature declines over time. One possible explanation for this decline may be changes in substrate availability. This hypothesis is examined by using the ecosystem model G'DAY, which simulates C and nitrogen (N) dynamics in plants and soil.We applied the model to observations from a soil-warming experiment in a Norway spruce (Picea abies (L.) Karst.) stand by simulating a step increase of soil temperature. The model provided a good qualitative reproduction of the observed reduction of heterotrophic respiration (R h ) under sustained warming. The simulations showed how the combined effects of faster turnover and reduced substrate availability lead to a transient increase of R h . The simulated annual increase in R h from soil was 60% in the first year after perturbation but decreased to 30% after a decade.One conclusion from the analysis of the simulations is that R h can decrease even though the temperature response function for decomposition remains unchanged. G'DAY suggests that acclimation of R h to soil warming is partly an effect of substrate depletion of labile C pools during the first decade of warming as a result of accelerated rates of mineralization. The response is attributed mainly to changing levels of C in pools with short time constants, reflecting the importance of high-quality soil C fractions. Changes of the structure or physiology of the decomposer community were not invoked. Therefore, it becomes a question of definition whether the simulated dynamics of the declining response of CO 2 release to the warming should be named acclimation or seen as a natural part of the system dynamics.