Climate change predictions derived from coupled carbon-climate models are highly dependent on assumptions about feedbacks between the biosphere and atmosphere. One critical feedback occurs if C uptake by the biosphere increases in response to the fossil-fuel driven increase in atmospheric [CO 2] (''CO2 fertilization''), thereby slowing the rate of increase in atmospheric [CO 2]. Carbon exchanges between the terrestrial biosphere and atmosphere are often first represented in models as net primary productivity (NPP). However, the contribution of CO 2 fertilization to the future global C cycle has been uncertain, especially in forest ecosystems that dominate global NPP, and models that include a feedback between terrestrial biosphere metabolism and atmospheric [CO 2] are poorly constrained by experimental evidence. We analyzed the response of NPP to elevated CO 2 (Ϸ550 ppm) in four free-air CO 2 enrichment experiments in forest stands. We show that the response of forest NPP to elevated [CO 2] is highly conserved across a broad range of productivity, with a stimulation at the median of 23 ؎ 2%. At low leaf area indices, a large portion of the response was attributable to increased light absorption, but as leaf area indices increased, the response to elevated [CO 2] was wholly caused by increased light-use efficiency. The surprising consistency of response across diverse sites provides a benchmark to evaluate predictions of ecosystem and global models and allows us now to focus on unresolved questions about carbon partitioning and retention, and spatial variation in NPP response caused by availability of other growth limiting resources.CO2 fertilization ͉ global change ͉ leaf area index ͉ net primary productivity A nalysis and prediction of the effects of human activities, particularly the combustion of fossil fuels, on climate and the biological, physical, and social responses to changing climate require an integrated view of the complex interactions between the biosphere and atmosphere. Carbon cycle models are now being coupled to atmosphere-ocean general circulation climate models to achieve a dynamic analysis of the relationships between C emissions, atmospheric chemistry, biosphere activity, and climatic change (1-3).Exchanges between the terrestrial biosphere and atmosphere are represented in models using empirical and theoretical expressions of net primary productivity (NPP), the net fixation of C by green plants into organic matter, or the difference between photosynthesis and plant respiration. Because the photosynthetic uptake of carbon that drives NPP is not saturated at current atmospheric concentrations (4), NPP should increase as fossilfuel combustion adds to the atmospheric [CO 2 ]. Increased C uptake into the biosphere in response to rising [CO 2 ] (''CO 2 fertilization'') can create a negative feedback that slows the rate of increase in atmospheric [CO 2 ] (3, 5). Hence, assumptions regarding CO 2 fertilization of the terrestrial biosphere greatly affect predictions of future atmospheric [CO 2 ] (3)...

Forest ecosystems are important sinks for rising concentrations of atmospheric CO 2. In previous research, we showed that net primary production (NPP) increased by 23 ؎ 2% when four experimental forests were grown under atmospheric concentrations of CO 2 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 CO 2 enrichment (FACE). In this study, experimental evidence demonstrates that the uptake of N increased under elevated CO 2 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 CO 2 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 CO 2. Regardless of the specific mechanism, this analysis shows that the larger quantities of C entering the below-ground system under elevated CO 2 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 CO 2 .global change ͉ net primary production

SUMMARYIn most natural ecosystems a significant portion of carbon fixed through photosynthesis is allocated to the production and maintenance of fine roots, the ephemeral portion of the root system that absorbs growth-limiting moisture and nutrients. In turn, senescence of fine roots can be the greatest source of C input to forest soils. Consequently, important questions in ecology entail the extent to which increasing atmospheric CO^ may alter the allocation of carbon to, and demography of, fine roots. Using microvideo and image analysis technology, we demonstrate that elevated atmospheric CO^ increases the rates of both fine root production and mortality. Rates of root mortality also increased substantially as soil nitrogen availability increased, regardless of COg concentration. Nitrogen greatly infiuenced the proportional allocation of carbon to leaves vs. fine roots. The amount of available nitrogen in the soil appears to be the most important factor regulating fine root demography in Populus trees.

Summary1. The impacts of elevated atmospheric CO 2 and/or O 3 have been examined over 4 years using an open-air exposure system in an aggrading northern temperate forest containing two different functional groups (the indeterminate, pioneer, O 3 -sensitive species Trembling Aspen, Populus tremuloides and Paper Birch, Betula papyrifera , and the determinate, late successional, O 3 -tolerant species Sugar Maple, Acer saccharum ). 2. The responses to these interacting greenhouse gases have been remarkably consistent in pure Aspen stands and in mixed Aspen/Birch and Aspen/Maple stands, from leaf to ecosystem level, for O 3 -tolerant as well as O 3 -sensitive genotypes and across various trophic levels. These two gases act in opposing ways, and even at low concentrations (1·5 × ambient, with ambient averaging 34 -36 nL L − 1 during the summer daylight hours), O 3 offsets or moderates the responses induced by elevated CO 2 . 3. After 3 years of exposure to 560 µ mol mol − 1 CO 2 , the above-ground volume of Aspen stands was 40% above those grown at ambient CO 2 , and there was no indication of a diminishing growth trend. In contrast, O 3 at 1·5 × ambient completely offset the growth enhancement by CO 2 , both for O 3 -sensitive and O 3 -tolerant clones. Implications of this finding for carbon sequestration, plantations to reduce excess CO 2 , and global models of forest productivity and climate change are presented.