SummaryAccurate representation of photosynthesis in terrestrial biosphere models (TBMs) is essential for robust projections of global change. However, current representations vary markedly between TBMs, contributing uncertainty to projections of global carbon fluxes. Here we compared the representation of photosynthesis in seven TBMs by examining leaf and canopy level responses of photosynthetic CO 2 assimilation (A) to key environmental variables: light, temperature, CO 2 concentration, vapor pressure deficit and soil water content. We identified research areas where limited process knowledge prevents inclusion of physiological phenomena in current TBMs and research areas where data are urgently needed for model parameterization or evaluation. We provide a roadmap for new science needed to improve the representation of photosynthesis in the next generation of terrestrial biosphere and Earth system models.
The ratio of plant carbon gain to water use, known as water use efficiency (WUE), has long been recognized as a key constraint on crop production and an important target for crop improvement. WUE is a physiologically and genetically complex trait that can be defined at a range of scales. Many component traits directly influence WUE, including photosynthesis, stomatal and mesophyll conductances, and canopy structure. Interactions of carbon and water relations with diverse aspects of the environment and crop development also modulate WUE. As a consequence, enhancing WUE by breeding or biotechnology has proven challenging but not impossible. This review aims to synthesize new knowledge of WUE arising from advances in phenotyping, modeling, physiology, genetics, and molecular biology in the context of classical theoretical principles. In addition, we discuss how rising atmospheric CO2 concentration has created and will continue to create opportunities for enhancing WUE by modifying the trade-off between photosynthesis and transpiration.
High-throughput, noninvasive field phenotyping has revealed genetic variation in crop morphological, developmental, and agronomic traits, but rapid measurements of the underlying physiological and biochemical traits are needed to fully understand genetic variation in plant-environment interactions. This study tested the application of leaf hyperspectral reflectance (l = 500-2,400 nm) as a high-throughput phenotyping approach for rapid and accurate assessment of leaf photosynthetic and biochemical traits in maize (Zea mays). Leaf traits were measured with standard wet-laboratory and gas-exchange approaches alongside measurements of leaf reflectance. Partial least-squares regression was used to develop a measure of leaf chlorophyll content, nitrogen content, sucrose content, specific leaf area, maximum rate of phosphoenolpyruvate carboxylation, [CO 2 ]-saturated rate of photosynthesis, and leaf oxygen radical absorbance capacity from leaf reflectance spectra. Partial least-squares regression models accurately predicted five out of seven traits and were more accurate than previously used simple spectral indices for leaf chlorophyll, nitrogen content, and specific leaf area. Correlations among leaf traits and statistical inferences about differences among genotypes and treatments were similar for measured and modeled data. The hyperspectral reflectance approach to phenotyping was dramatically faster than traditional measurements, enabling over 1,000 rows to be phenotyped during midday hours over just 2 to 4 d, and offers a nondestructive method to accurately assess physiological and biochemical trait responses to environmental stress.
The C 4 grass Zea mays (maize or corn) is the third most important food crop globally in terms of production and demand is predicted to increase 45% from 1997 to 2020. However, the effects of rising [CO 2 ] upon C 4 plants, and Z. mays specifically, are not sufficiently understood to allow accurate predictions of future crop production. A rainfed, field experiment utilizing free-air concentration enrichment (FACE) technology in the primary area of global corn production (US Corn Belt) was undertaken to determine the effects of elevated [CO 2 ] on corn. FACE technology allows experimental treatments to be imposed upon a complete soil-plant-atmosphere continuum with none of the effects of experimental enclosures on plant microclimate. Crop performance was compared at ambient [CO 2 ] (354 lmol mol À1 ) and the elevated [CO 2 ] (549 lmol mol À1 ) predicted for 2050. Previous laboratory studies suggest that under favorable growing conditions C 4 photosynthesis is not typically enhanced by elevated [CO 2 ]. However, stomatal conductance and transpiration are decreased, which can indirectly increase photosynthesis in dry climates. Given the deep soils and relatively high rainfall of the US Corn Belt, it was predicted that photosynthesis would not be enhanced by elevated [CO 2 ]. The diurnal course of gas exchange of upper canopy leaves was measured in situ across the growing season of 2002. Contrary to the prediction, growth at elevated [CO 2 ] significantly increased leaf photosynthetic CO 2 uptake rate (A) by up to 41%, and 10% on average. Greater A was associated with greater intercellular [CO 2 ], lower stomatal conductance and lower transpiration. Summer rainfall during 2002 was very close to the 50-year average for this site, indicating that the year was not atypical or a drought year. The results call for a reassessment of the established view that C 4 photosynthesis is insensitive to elevated [CO 2 ] under favorable growing conditions and that the production potential of corn in the US Corn Belt will not be affected by the global rise in [CO 2 ].
Insufficient water availability for crop production is a mounting barrier to achieving the 70% increase in food production that will be needed by 2050. One solution is to develop crops that require less water per unit mass of production. Water vapor transpires from leaves through stomata, which also facilitate the influx of CO2 during photosynthetic assimilation. Here, we hypothesize that Photosystem II Subunit S (PsbS) expression affects a chloroplast-derived signal for stomatal opening in response to light, which can be used to improve water-use efficiency. Transgenic tobacco plants with a range of PsbS expression, from undetectable to 3.7 times wild-type are generated. Plants with increased PsbS expression show less stomatal opening in response to light, resulting in a 25% reduction in water loss per CO2 assimilated under field conditions. Since the role of PsbS is universal across higher plants, this manipulation should be effective across all crops.
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