Tree responses to rising CO₂ in field experiments: implications for the future forest

The need to assess the role of forests in the global cycling of carbon and how that role will change as the atmospheric concentration of CO₂ increases has spawned many experiments over a range of scales. Experiments using open-top chambers have been established at many sites to test whether the short-term responses of tree seedlings described in controlled environments would be sustained over several growing seasons under field conditions. Here we review the results of those experiments, using the framework of the interacting cycles of carbon, water and nutrients, because that is the framework of the ecosystem models that are being used to address the decades-long response of forests. Our analysis suggests that most of what was learned in seedling studies was qualitatively correct. The evidence from field-grown trees suggests a continued and consistent stimulation of photosynthesis of about 60% for a 300 p.p.m. increase in [CO₂], and there is little evidence of the long-term loss of sensitivity to CO₂ that was suggested by earlier experiments with tree seedlings in pots. Despite the importance of respiration to a tree's carbon budget, no strong scientific consensus has yet emerged concerning the potential direct or acclimation response of woody plant respiration to CO₂ enrichment. The relative effect of CO₂ on above-ground dry mass was highly variable and greater than that indicated by most syntheses of seedling studies. Effects of CO₂ concentration on static measures of response are confounded with the acceleration of ontogeny observed in elevated CO₂. The trees in these open-top chamber experiments were in an exponential growth phase, and the large growth responses to elevated CO₂ resulted from the compound interest associated with an increasing leaf area. This effect cannot be expected to persist in a closed-canopy forest where growth potential is constrained by a steady-state leaf area index. A more robust and informative measure of tree growth in these experiments is the annual increment in wood mass per unit leaf area, which increased 27% in elevated CO₂. There is no support for the conclusion from many studies of seedlings that root-to-shoot ratio is increased by elevated CO₂; the production of fine roots may be enhanced, but it is not clear that this response would persist in a forest. Foliar nitrogen concentrations were lower in CO₂-enriched trees, but to a lesser extent than was indicated in seedling studies and only when expressed on a leaf mass basis. The prediction that leaf litter C/N ratio would increase was not supported in field experiments. Also contrasting with seedling studies, there is little evidence from the field studies that stomatal conductance is consistently affected by CO₂; however, this is a topic that demands more study. Experiments with trees in open-top chambers under field conditions have provided data on longer-term, larger-scale responses of trees to elevated CO₂ under field conditions, confirmed some of the conclusions from previous seedling studies, and challenged other conclusions. There remain important obstacles to using these experimental results to predict forest responses to rising CO₂, but the studies are valuable nonetheless for guiding ecosystem model development and revealing the critical questions that must be addressed in new, larger-scale CO₂ experiments.