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.

Comparing photosynthetic and photovoltaic efficiencies is not a simple issue. Although both processes harvest the energy in sunlight, they operate in distinctly different ways and produce different types of products: biomass or chemical fuels in the case of natural photosynthesis and nonstored electrical current in the case of photovoltaics. In order to find common ground for evaluating energy-conversion efficiency, we compare natural photosynthesis with present technologies for photovoltaic-driven electrolysis of water to produce hydrogen. Photovoltaic-driven electrolysis is the more efficient process when measured on an annual basis, yet short-term yields for photosynthetic conversion under optimal conditions come within a factor of 2 or 3 of the photovoltaic benchmark. We consider opportunities in which the frontiers of synthetic biology might be used to enhance natural photosynthesis for improved solar energy conversion efficiency.

The yield potential ( Y p ) of a grain crop is the seed mass per unit ground area obtained under optimum growing conditions without weeds, pests and diseases. It is determined by the product of the available light energy and by the genetically determined properties: efficiency of light capture ( e i ), the efficiency of conversion of the intercepted light into biomass ( e c ) and the proportion of biomass partitioned into grain ( h ). Plant breeding brings h and e i close to their theoretical maxima, leaving e c , primarily determined by photosynthesis, as the only remaining major prospect for improving Y p . Leaf photosynthetic rate, however, is poorly correlated with yield when different genotypes of a crop species are compared. This led to the viewpoint that improvement of leaf photosynthesis has little value for improving Y p . By contrast, the many recent experiments that compare the growth of a genotype in current and future projected elevated [CO 2 ] environments show that increase in leaf photosynthesis is closely associated with similar increases in yield. Are there opportunities to achieve similar increases by genetic manipulation? Six potential routes of increasing e c by improving photosynthetic efficiency were explored, ranging from altered canopy architecture to improved regeneration of the acceptor molecule for CO 2 . Collectively, these changes could improve e c and, therefore, Y p by c . 50%. Because some changes could be achieved by transgenic technology, the time of the development of commercial cultivars could be considerably less than by conventional breeding and potentially, within 10-15 years.

Model projections suggest that although increased temperature and decreased soil moisture will act to reduce global crop yields by 2050, the direct fertilization effect of rising carbon dioxide concentration ([CO 2 ]) will offset these losses. The CO 2 fertilization factors used in models to project future yields were derived from enclosure studies conducted approximately 20 years ago. Free-air concentration enrichment (FACE) technology has now facilitated large-scale trials of the major grain crops at elevated [CO 2 ] under fully open-air field conditions. In those trials, elevated [CO 2 ] enhanced yield by È50% less than in enclosure studies. This casts serious doubt on projections that rising [CO 2 ] will fully offset losses due to climate change. M uch effort has been put into linking models of climate and crop growth to project future changes in crop yields and food supply across the globe (1-4). Projections reviewed by the Intergovernmental Panel on Climate Change (IPCC) suggest that increased temperature and decreased soil moisture, which would otherwise reduce crop yields, will be offset by the direct fertilization effect of rising carbon dioxide concentration (ECO 2^) (5-7). The IPCC projections suggest that total crop yield may rise when averaged across the globe, but this net gain will result from generally lower yields in the tropics and increased yields in temperate zones. The accuracy of these projections and thus future food security depend critically on the magnitude of the CO 2 fertilization effect under actual growing conditions. Atmospheric ECO 2^h as risen from È260 parts per million (ppm) approximately 150 years ago to 380 ppm today (8). Yet ECO 2^i s markedly uniform across the globe; so, in contrast to temperature and soil moisture, there is no consistent spatial variation on which to estimate yield responses to increasing ECO 2^. Similarly, it is not easy to alter ECO 2^e xperimentally around a crop in the field. As a result, most information about crop responses to elevated ECO 2^i s obtained from studies in greenhouses, laboratory controlled-environment chambers, and transparent field chambers, where released CO 2 may be retained and easily controlled. These settings have provided the basis for projecting CO 2 fertilization effects on the major food crops: maize, rice, sorghum, soybeans, and wheat.Crops sense and respond directly to rising ECO 2^t hrough photosynthesis and stomatal conductance, and this is the basis for the fertilization effect on yield (9). In C 3 plants, mesophyll cells containing ribulose-1,5-bisphosphate carboxylase-oxygenase (RuBisCO) are in direct contact with the intercellular air space that is connected to the atmosphere via stomatal pores in the epidermis. Hence, in C 3 crops, rising CO 2 increases net photosynthetic CO 2 uptake because RuBisCO is not CO 2 -saturated in today_s atmosphere and because CO 2 inhibits the competing oxygenation reaction leading to photorespiration. RuBisCO is highly conserved across terrestrial plants, so instantaneous responses to...

The world's crop productivity is stagnating whereas population growth, rising affluence, and mandates for biofuels put increasing demands on agriculture. Meanwhile, demand for increasing cropland competes with equally crucial global sustainability and environmental protection needs. Addressing this looming agricultural crisis will be one of our greatest scientific challenges in the coming decades, and success will require substantial improvements at many levels. We assert that increasing the efficiency and productivity of photosynthesis in crop plants will be essential if this grand challenge is to be met. Here, we explore an array of prospective redesigns of plant systems at various scales, all aimed at increasing crop yields through improved photosynthetic efficiency and performance. Prospects range from straightforward alterations, already supported by preliminary evidence of feasibility, to substantial redesigns that are currently only conceptual, but that may be enabled by new developments in synthetic biology. Although some proposed redesigns are certain to face obstacles that will require alternate routes, the efforts should lead to new discoveries and technical advances with important impacts on the global problem of crop productivity and bioenergy production.light capture/conversion | carbon capture/conversion | smart canopy | enabling plant biotechnology tools | sustainable crop production Increasing demands for global food production over the next several decades portend a huge burden on the world's shrinking farmlands. Increasing global affluence, population growth, and demands for a bioeconomy (including livestock feed, bioenergy, chemical feedstocks, and biopharmaceuticals) will all require increased agricultural productivity, perhaps by as much as 60-120% over 2005 levels (e.g., refs. 1 and 2), putting increased productivity on a collision course with environmental and sustainability goals (3). The 45 y from 1960 to 2005 saw global food production grow ∼160%, mostly (135%) by improved production on

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