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.

Experimental evidence indicates that the stomatal conductance and nitrogen concentration ([N]) of foliage decline under CO2 enrichment, and that the percentage growth response to elevated CO2 is amplified under water limitation, but reduced under nitrogen limitation. We advance simple explanations for these responses based on an optimisation hypothesis applied to a simple model of the annual carbon–nitrogen–water economy of trees growing at a CO2-enrichment experiment at Oak Ridge, Tennessee, USA. The model is shown to have an optimum for leaf [N], stomatal conductance and leaf area index (LAI), where annual plant productivity is maximised. The optimisation is represented in terms of a trade-off between LAI and stomatal conductance, constrained by water supply, and between LAI and leaf [N], constrained by N supply. At elevated CO2 the optimum shifts to reduced stomatal conductance and leaf [N] and enhanced LAI. The model is applied to years with contrasting rainfall and N uptake. The predicted growth response to elevated CO2 is greatest in a dry, high-N year and is reduced in a wet, low-N year. The underlying physiological explanation for this contrast in the effects of water versus nitrogen limitation is that leaf photosynthesis is more sensitive to CO2 concentration ([CO2]) at lower stomatal conductance and is less sensitive to [CO2] at lower leaf [N].

Stable isotope natural abundance measurements integrate across several biogeochemical processes in ecosystem N and C dynamics. Here, we report trends in natural isotope abundance (d 13 C and d 15 N in plant and soil) along a climosequence of 33 Nothofagus forest stands located within Patagonia, Southern Argentina. We measured 28 different abiotic variables (both climatic variables and soil properties) to characterize environmental conditions at each of the 33 sites. Foliar d 13 C values ranged from À35.4& to À27.7&, and correlated positively with foliar d 15 N values, ranging from À3.7& to 5.2&. Soil d 13 C and d 15 N values reflected the isotopic trends of the foliar tissues and ranged from À29.8& to À25.3&, and À4.8& to 6.4&, respectively, with no significant differences between Nothofagus species (Nothofagus pumilio, Nothofagus antarctica, Nothofagus betuloides). Principal component analysis and multiple regressions suggested that mainly water availability variables (mean annual precipitation), but not soil properties, explained between 42% and 79% of the variations in foliar and soil d 13 C and d 15 N natural abundance, which declined with increased moisture supply. We conclude that a decline in water use efficiency at wetter sites promotes both the depletion of heavy C and N isotopes in soil and plant biomass. Soil d 13 C values were higher than those of the plant tissues and this difference increased as annual precipitation increased. No such differences were apparent when d 15 N values in soil and plant were compared, which indicates that climatic differences contributed more to the overall C balance than to the overall N balance in these forest ecosystems.