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Elevated [CO2] (eCO2) and water stress reduce leaf stomatal conductance (gs), which may affect leaf thermoregulation during heat waves (heat stress). Two sorghum lines, with different leaf width were grown in a glasshouse at a mean day temperature of 30 °C, under different [CO2] and watering levels, and subjected to heat stress (43 °C) for 6 d at the start of the reproductive stage. We measured leaf photosynthetic and stomatal responses to light transients before harvesting the plants. Photosynthesis at growth conditions (Agrowth) and biomass accumulation were enhanced by eCO2 under control conditions. Heat stress increased gs, especially in wider leaves, and reduced the time constant of stomatal opening (kopen) at ambient [CO2] but not eCO2. However, heat stress reduced photosynthesis under water stress and eCO2 due to increased leaf temperature and reduced evaporative cooling. eCO2 prevented the reduction of biomass under both water and heat stress, possibly due to improved plant and soil water status as a result of reduced gs. Our results suggest that the response of the C4 crop sorghum to future climate conditions depends on the trade-off between low gs needed for high water use efficiency and drought tolerance, and the high gs needed for improved thermoregulation and heat tolerance under an eCO2 future.

Sustaining crop productivity and resilience in water-limited environments and under rising temperatures are matters of concern worldwide. We investigated the leaf anatomical traits that underpin our recently identified link between leaf width (LW) and intrinsic water use efficiency (iWUE), as traits of interest in plant breeding. Ten sorghum lines with varying LW were grown under three temperatures to expand the range of variation of both LW and gas exchange rates. Leaf gas exchange, surface morphology and cross-sectional anatomy were measured and analysed using structural equations modelling. Narrower leaves had lower stomatal conductance (gs ) and higher iWUE across growth temperatures. They also had smaller intercellular airspaces, stomatal size, percentage of open stomatal aperture relative to maximum, hydraulic pathway, mesophyll thickness, and leaf mass per area. Structural modelling revealed a developmental association among leaf anatomical traits that underpinned gs variation in sorghum. Growing temperature and LW both impacted leaf gas exchange rates, but only LW directly impacted leaf anatomy. Wider leaves may be more productive under well-watered conditions, but consume more water for growth and development, which is detrimental under water stress.

Chloroplasts movement within mesophyll cells in C4 plants is hypothesized to enhance the CO2 concentrating mechanism, but this is difficult to verify experimentally. A three-dimensional (3D) leaf model can help analyse how chloroplast movement influences the operation of the CO2 concentrating mechanism. The first volumetric reaction-diffusion model of C4 photosynthesis that incorporates detailed 3D leaf anatomy, light propagation, ATP and NADPH production, and CO2, O2 and bicarbonate concentration driven by diffusional and assimilation/emission processes was developed. It was implemented for maize leaves to simulate various chloroplast movement scenarios within mesophyll cells: the movement of all mesophyll chloroplasts towards bundle sheath cells (aggregative movement) and movement of only those of interveinal mesophyll cells towards bundle sheath cells (avoidance movement). Light absorbed by bundle sheath chloroplasts relative to mesophyll chloroplasts increased in both cases. Avoidance movement decreased light absorption by mesophyll chloroplasts considerably. Consequently, total ATP and NADPH production and net photosynthetic rate increased for aggregative movement and decreased for avoidance movement compared with the default case of no chloroplast movement at high light intensities. Leakiness increased in both chloroplast movement scenarios due to the imbalance in energy production and demand in mesophyll and bundle sheath cells. These results suggest the need to design strategies for coordinated increases in electron transport and Rubisco activities for an efficient CO2 concentrating mechanism at very high light intensities.

Despite its importance for crop water use and productivity, especially in drought-affected environments, the underlying mechanisms of variation in intrinsic water-use efficiency (iWUE = net photosynthesis/stomatal conductance for water vapour, gsw ) are not well understood, especially in C4 plants. Recently, we discovered that leaf width (LW) correlated negatively with iWUE and positively with gsw across several C4 grasses. Here, we confirmed these relationships within 48 field-grown genotypes differing in LW in Sorghum bicolor, a C4 crop adapted to dry and hot conditions. We measured leaf gas exchange and modelled leaf energy balance three times a day, alongside anatomical traits as potential predictors of iWUE. LW correlated negatively with iWUE and stomatal density, but positively with gsw , interveinal distance of longitudinal veins, and the percentage of stomatal aperture relative to maximum. Energy balance modelling showed that wider leaves needed to open their stomata more to generate a more negative leaf-to-air temperature difference, especially at midday when air temperatures exceeded 40°C. These results highlight the important role that LW plays in shaping iWUE through coordination of vein and stomatal traits and by affecting stomatal aperture. Therefore, LW could be used as a predictor of higher iWUE among sorghum genotypes.