Atmospheric CO(2) concentration ([CO(2)]) is now higher than it was at any time in the past 26 million years and is expected to nearly double during this century. Terrestrial plants with the C(3) photosynthetic pathway respond in the short term to increased [CO(2)] via increased net photosynthesis and decreased transpiration. In the longer term this increase is often offset by downregulation of photosynthetic capacity. But much of what is currently known about plant responses to elevated [CO(2)] comes from enclosure studies, where the responses of plants may be modified by size constraints and the limited life-cycle stages that are examined. Free-Air CO(2) Enrichment (FACE) was developed as a means to grow plants in the field at controlled elevation of CO(2) under fully open-air field conditions. The findings of FACE experiments are quantitatively summarized via meta-analytic statistics and compared to findings from chamber studies. Although trends agree with parallel summaries of enclosure studies, important quantitative differences emerge that have important implications both for predicting the future terrestrial biosphere and understanding how crops may need to be adapted to the changed and changing atmosphere.

Increasing the yield potential of the major food grain crops has contributed very significantly to a rising food supply over the past 50 years, which has until recently more than kept pace with rising global demand. Whereas improved photosynthetic efficiency has played only a minor role in the remarkable increases in productivity achieved in the last half century, further increases in yield potential will rely in large part on improved photosynthesis. Here we examine inefficiencies in photosynthetic energy transduction in crops from light interception to carbohydrate synthesis, and how classical breeding, systems biology, and synthetic biology are providing new opportunities to develop more productive germplasm. Near-term opportunities include improving the display of leaves in crop canopies to avoid light saturation of individual leaves and further investigation of a photorespiratory bypass that has already improved the productivity of model species. Longer-term opportunities include engineering into plants carboxylases that are better adapted to current and forthcoming CO(2) concentrations, and the use of modeling to guide molecular optimization of resource investment among the components of the photosynthetic apparatus, to maximize carbon gain without increasing crop inputs. Collectively, these changes have the potential to more than double the yield potential of our major crops.

Plant responses to the projected future levels of CO(2) were first characterized in short-term experiments lasting days to weeks. However, longer term acclimation responses to elevated CO(2) were subsequently discovered to be very important in determining plant and ecosystem function. Free-Air CO(2) Enrichment (FACE) experiments are the culmination of efforts to assess the impact of elevated CO(2) on plants over multiple seasons and, in the case of crops, over their entire lifetime. FACE has been used to expose vegetation to elevated concentrations of atmospheric CO(2) under completely open-air conditions for nearly two decades. This review describes some of the lessons learned from the long-term investment in these experiments. First, elevated CO(2) stimulates photosynthetic carbon gain and net primary production over the long term despite down-regulation of Rubisco activity. Second, elevated CO(2) improves nitrogen use efficiency and, third, decreases water use at both the leaf and canopy scale. Fourth, elevated CO(2) stimulates dark respiration via a transcriptional reprogramming of metabolism. Fifth, elevated CO(2) does not directly stimulate C(4) photosynthesis, but can indirectly stimulate carbon gain in times and places of drought. Finally, the stimulation of yield by elevated CO(2) in crop species is much smaller than expected. While many of these lessons have been most clearly demonstrated in crop systems, all of the lessons have important implications for natural systems.