Donald R. Ort

Food for thought: Lower-than-expected crop yield stimulation with rising CO2 concentrations

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 ([CO2]) will offset these losses. The CO2 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 [CO2] under fully open-air field conditions. In those trials, elevated [CO2] enhanced yield by ∼50% less than in enclosure studies. This casts serious doubt on projections that rising [CO2] will fully offset losses due to climate change.

Comparing Photosynthetic and Photovoltaic Efficiencies and Recognizing the Potential for Improvement

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.

Elevated CO2 effects on plant carbon, nitrogen, and water relations: six important lessons from FACE

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.

Improving Photosynthetic Efficiency for Greater Yield

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.

Impacts of chilling temperatures on photosynthesis in warm-climate plants

Photosynthesis in warm-climate plants is substantially reduced after chilling. Tropical and subtropical species offer the opportunity to study the effects of low temperature on photosynthetic processes undisguised by the myriad of protective responses observed in temperate species. In this article, we highlight the primary components of photosynthesis that are affected by a short chill, in both the dark and the light, and discuss what is known of the mechanisms involved. Recent work implicates impaired redox and circadian regulation among other processes.

What is the maximum efficiency with which photosynthesis can convert solar energy into biomass

Photosynthesis is the source of our food and fiber. Increasing world population, economic development, and diminishing land resources forecast that a doubling of productivity is critical in meeting agricultural demand before the end of this century. A starting point for evaluating the global potential to meet this goal is establishing the maximum efficiency of photosynthetic solar energy conversion. The potential efficiency of each step of the photosynthetic process from light capture to carbohydrate synthesis is examined. This reveals the maximum conversion efficiency of solar energy to biomass is 4.6% for C3 photosynthesis at 30 degrees C and todays 380 ppm atmospheric [CO2], but 6% for C4 photosynthesis. This advantage over C3 will disappear as atmospheric [CO2] nears 700 ppm.

A photoprotective role for O2 as an alternative electron sink in photosynthesis

Photoprotection of the photosynthetic apparatus has two essential elements: first, the thermal dissipation of excess excitation energy in the photosystem II antennae (i.e. non-photochemical quenching), and second, the ability of photosystem II to transfer electrons to acceptors within the chloroplast (i.e. photochemical quenching). Recent studies indicate that the proportion of absorbed photons that are thermally dissipated through the non-photochemical pathway often reaches a maximum well before saturating irradiances are reached. Hence, photochemical quenching is crucial for photoprotection at saturating light intensities. When plants are exposed to environmental stresses and the availability of CO(2) within the leaf is restricted, the reduction of oxygen by both the photorespiratory and the Mehler ascorbate peroxidase pathways appears to play a critical photoprotective role, substituting for CO(2) in sustaining electron flow. Induction of high activity of the Mehler ascorbate peroxidase pathway may be associated with acclimation to environmental stress.

Photosynthesis, productivity, and yield of maize are not affected by open-air elevation of CO2 concentration in the absence of drought

While increasing temperatures and altered soil moisture arising from climate change in the next 50 years are projected to decrease yield of food crops, elevated CO2 concentration ([CO2]) is predicted to enhance yield and offset these detrimental factors. However, C4 photosynthesis is usually saturated at current [CO2] and theoretically should not be stimulated under elevated [CO2]. Nevertheless, some controlled environment studies have reported direct stimulation of C4 photosynthesis and productivity, as well as physiological acclimation, under elevated [CO2]. To test if these effects occur in the open air and within the Corn Belt, maize (Zea mays) was grown in ambient [CO2] (376 μmol mol−1) and elevated [CO2] (550 μmol mol−1) using Free-Air Concentration Enrichment technology. The 2004 season had ideal growing conditions in which the crop did not experience water stress. In the absence of water stress, growth at elevated [CO2] did not stimulate photosynthesis, biomass, or yield. Nor was there any CO2 effect on the activity of key photosynthetic enzymes, or metabolic markers of carbon and nitrogen status. Stomatal conductance was lower (−34%) and soil moisture was higher (up to 31%), consistent with reduced crop water use. The results provide unique field evidence that photosynthesis and production of maize may be unaffected by rising [CO2] in the absence of drought. This suggests that rising [CO2] may not provide the full dividend to North American maize production anticipated in projections of future global food supply.

Biotic stress globally downregulates photosynthesis genes

To determine if damage to foliage by biotic agents, including arthropods, fungi, bacteria and viral pathogens, universally downregulates the expression of genes involved in photosynthesis, we compared transcriptome data from microarray experiments after twenty two different forms of biotic damage on eight different plant species. Transcript levels of photosynthesis light reaction, carbon reduction cycle and pigment synthesis genes decreased regardless of the type of biotic attack. The corresponding upregulation of genes coding for the synthesis of jasmonic acid and those involved in the responses to salicylic acid and ethylene suggest that the downregulation of photosynthesis-related genes was part of a defence response. Analysis of the sub-cellular targeting of co-expressed gene clusters revealed that the transcript levels of 84% of the genes that carry a chloroplast targeting peptide sequence decreased. The majority of these downregulated genes shared common regulatory elements, such as G-box (CACGTG), T-box (ACTTTG) and SORLIP (GCCAC) motifs. Strong convergence in the response of transcription suggests that the universal downregulation of photosynthesis-related gene expression is an adaptive response to biotic attack. We hypothesize that slow turnover of many photosynthetic proteins allows plants to invest resources in immediate defence needs without debilitating near term losses in photosynthetic capacity.

More than taking the heat: Crops and global change

Grain production per unit of land will need to more than double over this century to address rising population and demand. This at a time when the procedures that have delivered increased yields over the past 50 years may have reached their ceiling for some of the worlds most important crops. Rising global temperature and more frequent droughts will act to drive down yields. The projected rise in atmospheric [CO(2)] by mid-century could in theory increase crop photosynthesis by over 30%, but this is not realized in grain yields in current C(3) cultivars in the field. Emerging understanding of gene networks controlling responses to these environmental changes indicates biotechnological opportunities for adaptation. Considerably more basic research, particularly under realistic field conditions, is critical before these opportunities can be adequately understood and validated. Given the time needed between discovery in a model plant species and translation to traits or stacked changes in a commercial grain crop cultivar, there is an urgent need to vigorously pursue and develop these opportunities now.