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Purpose Large parts of the Amazon rainforest grow on weathered soils depleted in phosphorus and rock-derived cations. We tested the hypothesis that in this ecosystem, fine roots stimulate decomposition and nutrient release from leaf litter biochemically by releasing enzymes, and by exuding labile carbon stimulating microbial decomposers. Methods We monitored leaf litter decomposition in a Central Amazon tropical rainforest, where fine roots were either present or excluded, over 188 days and added labile carbon substrates (glucose and citric acid) in a fully factorial design. We tracked litter mass loss, remaining carbon, nitrogen, phosphorus and cation concentrations, extracellular enzyme activity and microbial carbon and nutrient concentrations. Results Fine root presence did not affect litter mass loss but significantly increased the loss of phosphorus and cations from leaf litter. In the presence of fine roots, acid phosphatase activity was 43.2% higher, while neither microbial stoichiometry, nor extracellular enzyme activities targeting carbon- and nitrogen-containing compounds changed. Glucose additions increased phosphorus loss from litter when fine roots were present, and enhanced phosphatase activity in root exclusions. Citric acid additions reduced litter mass loss, microbial biomass nitrogen and phosphorus, regardless of fine root presence or exclusion. Conclusions We conclude that plant roots release significant amounts of acid phosphatases into the litter layer and mobilize phosphorus without affecting litter mass loss. Our results further indicate that added labile carbon inputs (i.e. glucose) can stimulate acid phosphatase production by microbial decomposers, highlighting the potential importance of plant-microbial feedbacks in tropical forest ecosystems.

Global terrestrial models currently predict that the Amazon rainforest will continue to act as a carbon sink in the future, primarily owing to the rising atmospheric carbon dioxide (CO2) concentration. Soil phosphorus impoverishment in parts of the Amazon basin largely controls its functioning, but the role of phosphorus availability has not been considered in global model ensembles—for example, during the Fifth Climate Model Intercomparison Project. Here we simulate the planned free-air CO2 enrichment experiment AmazonFACE with an ensemble of 14 terrestrial ecosystem models. We show that phosphorus availability reduces the projected CO2-induced biomass carbon growth by about 50% to 79 ± 63 g C m−2 yr−1 over 15 years compared to estimates from carbon and carbon–nitrogen models. Our results suggest that the resilience of the region to climate change may be much less than previously assumed. Variation in the biomass carbon response among the phosphorus-enabled models is considerable, ranging from 5 to 140 g C m−2 yr−1, owing to the contrasting plant phosphorus use and acquisition strategies considered among the models. The Amazon forest response thus depends on the interactions and relative contributions of the phosphorus acquisition and use strategies across individuals, and to what extent these processes can be upregulated under elevated CO2.

Many serious ecosystem consequences of climate change will take decades or even centuries to emerge. Long-term ecological responses to global change are strongly regulated by slow processes, such as changes in species composition, carbon dynamics in soil and by long-lived plants, and accumulation of nutrient capitals. Understanding and predicting these processes require experiments on decadal time scales. But decadal experiments by themselves may not be adequate because many of the slow processes have characteristic time scales much longer than experiments can be maintained. This article promotes a coordinated approach that combines long-term, large-scale global change experiments with process studies and modeling. Long-term global change manipulative experiments, especially in high-priority ecosystems such as tropical forests and high-latitude regions, are essential to maximize information gain concerning future states of the earth system. The long-term experiments should be conducted in tandem with complementary process studies, such as those using model ecosystems, species replacements, laboratory incubations, isotope tracers, and greenhouse facilities. Models are essential to assimilate data from long-term experiments and process studies together with information from long-term observations, surveys, and space-for-time studies along environmental and biological gradients. Future research programs with coordinated long-term experiments, process studies, and modeling have the potential to be the most effective strategy to gain the best information on long-term ecosystem dynamics in response to global change.

Despite the importance of nitrogen (N) limitation of forest carbon (C) sequestration at rising atmospheric CO2 concentration, the mechanisms responsible are not well understood. To elucidate the interactive effects of elevated CO2 (eCO2) and soil N availability on forest productivity and C allocation, we hypothesized that (1) trees maximize fitness by allocating N and C to maximize their net growth and (2) that N uptake is controlled by soil N availability and root exploration for soil N. We tested this model using data collected in Free-Air CO2 Enrichment sites dominated by evergreen (Pinus taeda; Duke Forest) and deciduous [Liquidambar styraciflua; Oak Ridge National Laboratory (ORNL)] trees. The model explained 80 -95% of variation in productivity and N-uptake data among eCO2, N fertilization and control treatments over 6 years. The model explains why fine-root production increased, and why N uptake increased despite reduced soil N availability under eCO2 at ORNL and Duke. In agreement with observations at other sites, the model predicts that soil N availability reduced below a critical level diminishes all eCO2 responses. At Duke, a negative feedback between reduced soil N availability and N uptake prevented progressive reduction in soil N availability at eCO2. At ORNL, soil N availability progressively decreased because it did not trigger reductions in N uptake; N uptake was maintained at ORNL through a large increase in the production of fast turnover fine roots. This implies that species with fast root turnover could be more prone to progressive N limitation of carbon sequestration in woody biomass than species with slow root turnover, such as evergreens. However, longer term data are necessary for a thorough evaluation of this hypothesis. The success of the model suggests that the principle of maximization of net growth to control growth and allocation could serve as a basis for simplification and generalization of larger scale forest and ecosystem models, f

Forest ecosystems are important sinks for rising concentrations of atmospheric CO2. In previous research, we showed that net primary production (NPP) increased by 23 ± 2% when four experimental forests were grown under atmospheric concentrations of CO2 predicted for the latter half of this century. Because nitrogen (N) availability commonly limits forest productivity, some combination of increased N uptake from the soil and more efficient use of the N already assimilated by trees is necessary to sustain the high rates of forest NPP under free-air CO2 enrichment (FACE). In this study, experimental evidence demonstrates that the uptake of N increased under elevated CO2 at the Rhinelander, Duke, and Oak Ridge National Laboratory FACE sites, yet fertilization studies at the Duke and Oak Ridge National Laboratory FACE sites showed that tree growth and forest NPP were strongly limited by N availability. By contrast, nitrogen-use efficiency increased under elevated CO2 at the POP-EUROFACE site, where fertilization studies showed that N was not limiting to tree growth. Some combination of increasing fine root production, increased rates of soil organic matter decomposition, and increased allocation of carbon (C) to mycorrhizal fungi is likely to account for greater N uptake under elevated CO2. Regardless of the specific mechanism, this analysis shows that the larger quantities of C entering the below-ground system under elevated CO2 result in greater N uptake, even in N-limited ecosystems. Biogeochemical models must be reformulated to allow C transfers below ground that result in additional N uptake under elevated CO2.

Gap models have been used extensively in ecological studies of forest structure and succession, and they should be useful tools for studying potential responses of forests to climatic change. There is a wide variety of gap models with different degrees of physiological detail, and the manner in which the effects of climatic factors are analyzed varies across that range of detail. Here we consider how well the current suite of gap models can accommodate climatic-change issues, and we suggest what physiological attributes and responses should be added to better represent responses of aboveground growth and competition. Whether a gap model is based on highly empirical, aggregated growth functions or more mechanistic expressions of carbon uptake and allocation, the greatest challenge will be to express allocation correctly. For example, incorporating effects of elevated CO2 requires that the fixed allometry between stem volume and leaf area be made flexible. Simulation of the effects of climatic warming should incorporate the possibility of a longer growing season and acclimation of growth processes to changing temperature. To accommodate climatic-change factors, some of the simplicity of gap models must be sacrificed by increasing the amount of physiological detail, but it is important that the capability of the models to predict competition and successional dynamics not be sacrificed.