Midday water use efficiency in sorghum is linked to faster stomatal closure rate, lower stomatal aperture and higher stomatal density.

Most studies assume midday gas exchange measurements capture the leaf's daytime performance. However, stomatal conductance (gs ) and photosynthesis (An ) fluctuate diurnally due to endogenous and environmental rhythms, which can affect intrinsic water use efficiency (iWUE). Six Sorghum lines with contrasting stomatal anatomical traits were grown in environmentally controlled conditions, and leaf gas exchange was measured three times a day. Stomatal anatomy and kinetic responses to light transients were also measured. The highest An and gs and the lowest iWUE were observed at midday for most lines. Diurnally averaged iWUE correlated positively with morning and midday iWUE and negatively with the time taken for stomata to close after transition to low light intensity (kclose ). There was significant variation among sorghum lines for kclose , and smaller kclose correlated with lower gs and higher stomatal density (SD) across the lines. In turn, gs was negatively correlated with SD and regulated by the operational stomatal aperture regardless of stomatal size. Altogether, our data suggest a common physiology to improve iWUE in sorghum related to the control of water loss without impacting photosynthesis relying on higher SD, lower stomatal aperture and faster stomatal closing in response to low light intensity.

Pushing the envelope: do narrowly and widely distributed Eucalyptus species differ in response to climate warming?

Contemporary climate change will push many tree species into conditions that are outside their current climate envelopes. Using the Eucalyptus genus as a model, we addressed whether species with narrower geographical distributions show constrained ability to cope with warming relative to species with wider distributions, and whether this ability differs among species from tropical and temperate climates. We grew seedlings of widely and narrowly distributed Eucalyptus species from temperate and tropical Australia in a glasshouse under two temperature regimes: the summer temperature at seed origin and +3.5°C. We measured physical traits and leaf-level gas exchange to assess warming influences on growth rates, allocation patterns, and physiological acclimation capacity. Warming generally stimulated growth, such that higher relative growth rates early in development placed seedlings on a trajectory of greater mass accumulation. The growth enhancement under warming was larger among widely than narrowly distributed species and among temperate rather than tropical provenances. The differential growth enhancement was primarily attributable to leaf area production and adjustments of specific leaf area. Our results suggest that tree species, including those with climate envelopes that will be exceeded by contemporary climate warming, possess capacity to physiologically acclimate but may have varying ability to adjust morphology.

Sugar sensing in C4 source leaves: a gap that needs to be filled.

Plant growth depends on sugar production and export by photosynthesizing source leaves and sugar allocation and import by sink tissues (grains, roots, stems, and young leaves). Photosynthesis and sink demand are tightly coordinated through metabolic (substrate, allosteric) feedback and signalling (sugar, hormones) mechanisms. Sugar signalling integrates sugar production with plant development and environmental cues. In C3 plants (e.g. wheat and rice), it is well documented that sugar accumulation in source leaves, due to source-sink imbalance, negatively feeds back on photosynthesis and plant productivity. However, we have a limited understanding about the molecular mechanisms underlying those feedback regulations, especially in C4 plants (e.g. maize, sorghum, and sugarcane). Recent work with the C4 model plant Setaria viridis suggested that C4 leaves have different sugar sensing thresholds and behaviours relative to C3 counterparts. Addressing this research priority is critical because improving crop yield requires a better understanding of how plants coordinate source activity with sink demand. Here we review the literature, present a model of action for sugar sensing in C4 source leaves, and suggest ways forward.

Anatomical drivers of stomatal conductance in sorghum lines with different leaf widths grown under different temperatures.

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.

Elevated [CO2] negatively impacts C4 photosynthesis under heat and water stress without penalizing biomass.

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.

The role of chloroplast movement in C4 photosynthesis: a theoretical analysis using a three-dimensional reaction-diffusion model for maize.

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.

Spatial expression patterns of genes encoding sugar sensors in leaves of C4 and C3 grasses.

BACKGROUND AND AIMS: The mechanisms of sugar sensing in grasses remain elusive, especially those using C4 photosynthesis even though a large proportion of the world's agricultural crops utilize this pathway. We addressed this gap by comparing the expression of genes encoding components of sugar sensors in C3 and C4 grasses, with a focus on source tissues of C4 grasses. Given C4 plants evolved into a two-cell carbon fixation system, it was hypothesized this may have also changed how sugars were sensed. METHODS: For six C3 and eight C4 grasses, putative sugar sensor genes were identified for target of rapamycin (TOR), SNF1-related kinase 1 (SnRK1), hexokinase (HXK) and those involved in the metabolism of the sugar sensing metabolite trehalose-6-phosphate (T6P) using publicly available RNA deep sequencing data. For several of these grasses, expression was compared in three ways: source (leaf) versus sink (seed), along the gradient of the leaf, and bundle sheath versus mesophyll cells. KEY RESULTS: No positive selection of codons associated with the evolution of C4 photosynthesis was identified in sugar sensor proteins here. Expressions of genes encoding sugar sensors were relatively ubiquitous between source and sink tissues as well as along the leaf gradient of both C4 and C3 grasses. Across C4 grasses, SnRK1ß1 and TPS1 were preferentially expressed in the mesophyll and bundle sheath cells, respectively. Species-specific differences of gene expression between the two cell types were also apparent. CONCLUSIONS: This comprehensive transcriptomic study provides an initial foundation for elucidating sugar-sensing genes within major C4 and C3 crops. This study provides some evidence that C4 and C3 grasses do not differ in how sugars are sensed. While sugar sensor gene expression has a degree of stability along the leaf, there are some contrasts between the mesophyll and bundle sheath cells.

Moderate heat stress prevented the observed biomass and yield stimulation caused by elevated CO2 in two well-watered wheat cultivars.

KEY MESSAGE: Heat stress (HS) under well-watered conditions was not detrimental to leaf photosynthesis or yield but modified the elevated CO2 response of photosynthesis and yield in two contrasting wheat cultivars. Climate change is increasing the frequency of extreme events such as heat waves, adversely affecting crop productivity. While positive impacts of elevated carbon dioxide (eCO2) on crop productivity are evident, the interactive effects of eCO2 and environmental stresses are still unclear. To investigate the interactive effects of elevated CO2 and heat stress (HS), we grew two contrasting wheat cultivars, early-maturing Scout and high-tillering Yitpi, under non-limiting water and nutrients at ambient (aCO2, 450 ppm) or elevated (eCO2, 650 ppm) CO2 and 22 °C in the glasshouse. Plants were exposed to two 3-day HS cycles at the vegetative (38.1 °C) and/or flowering (33.5 °C) stage. At aCO2, both wheat cultivars showed similar responses of photosynthesis and mesophyll conductance to temperature and produced similar grain yield. Relative to aCO2, eCO2 enhanced photosynthesis rate and reduced stomatal conductance and maximal carboxylation rate (Vcmax). During HS, high temperature stimulated photosynthesis at eCO2 in both cultivars, while eCO2 stimulated photosynthesis in Scout. Electron transport rate (Jmax) was unaffected by any treatment. eCO2 equally enhanced biomass and grain yield of both cultivars in control, but not HS, plants. HS reduced biomass and yield of Scout at eCO2. Yitpi, the cultivar with higher grain nitrogen, underwent a trade-off between grain yield and nitrogen. In conclusion, eCO2 improved photosynthesis of control and HS wheat, and improved biomass and grain yield of control plants only. Under well-watered conditions, HS was not detrimental to photosynthesis or growth but precluded a yield response to eCO2.

Coordination of stomata and vein patterns with leaf width underpins water-use efficiency in a C4 crop.

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.

High intrinsic water use efficiency is underpinned by high stomatal aperture and guard cell potassium flux in C3 and C4 grasses grown at glacial CO2 and low light.

We compared how stomatal morphology and physiology control intrinsic leaf water use efficiency (iWUE) in two C3 and six C4 grasses grown at ambient (400 µmol mol-1) or glacial CO2 (180 µmol mol-1) and high (1000 µmol m-2 s-1) or low light intensity (200 µmol m-2 s-1). C4 grasses tended to have higher iWUE and CO2 assimilation rates, and lower stomatal conductance (gs), operational stomatal aperture (aop), and guard cell K+ influx rate relative to C3 grasses, while stomatal size (SS) and stomatal density (SD) did not vary according to the photosynthetic type. Overall, iWUE and gs depended most on aop and density of open stomata. In turn, aop correlated with K+ influx, stomatal opening speed on transition to high light, and SS. Species with higher SD had smaller and faster-opening stomata. Although C4 grasses operated with lower gs and aop at ambient CO2, they showed a greater potential to open stomata relative to maximal stomatal conductance (gmax), indicating heightened stomatal sensitivity and control. We uncovered promising links between aop, gs, iWUE, and K+ influx among C4 grasses, and differential K+ influx responses of C4 guard cells to low light, revealing molecular targets for improving iWUE in C4 crops.

Leaf pigments and photosystems stoichiometry underpin photosynthetic efficiency of related C3 , C-C4 and C4 grasses under shade.

The quantum yield of photosynthesis (QY, CO2 fixed per light absorbed) depends on the efficiency of light absorption, the coupling between light absorption and electron transport, and the coupling between electron transport and carbon metabolism. QY is generally lower in C3 relative to C4 plants at warm temperatures and differs among the C4 subtypes. We investigated the acclimation to shade of light absorption and electron transport in six representative grasses with C3 , C3 -C4 and C4 photosynthesis. Plants were grown under full (control) or 25% (shade) sunlight. We measured the in vivo activity and stoichiometry of PSI and PSII, leaf spectral properties and pigment contents, and photosynthetic enzyme activities. Under control growth-light conditions, C4 species had higher CO2 assimilation rates, which declined to a greater extent relative to the C3 species. Whole leaf PSII/PSI ratios were highest in the C3 species, while QY and cyclic electron flow (CEF) were highest in the C4 , NADP-ME species. Shade significantly reduced leaf PSII/PSI, linear electron flow (LEF) and CEF of most species. Overall, shade reduced leaf absorptance, especially in the green region, as well as carotenoid and chlorophyll contents in C4 more than non-C4 species. The NAD-ME species underwent the greatest reduction in leaf absorptance and pigments under shade. In conclusion, shade compromised QY the least in the C3 and the most in the C4 -NAD-ME species. Different sensitivity to shade was associated with the ability to maintain leaf absorptance and pigments. This is important for maximising light absorption and minimising photoprotection under low light.

The transcriptomic responses of C4 grasses to subambient CO2 and low light are largely species specific and only refined by photosynthetic subtype.

Three subtypes of C4 photosynthesis exist (NADP-ME, NAD-ME and PEPCK), each known to be beneficial under specific environmental conditions. However, the influence of photosynthetic subtype on transcriptomic plasticity, as well as the genes underpinning this variability, remain largely unknown. Here, we comprehensively investigate the responses of six C4 grass species, spanning all three C4 subtypes, to two controlled environmental stresses: low light (200 µmol m-2  sec-1 ) and glacial CO2 (subambient; 180 ppm). We identify a susceptibility within NADP-ME species to glacial CO2 . Notably, although glacial CO2 phenotypes could be tied to C4 subtype, biochemical and transcriptomic responses to glacial CO2 were largely species specific. Nevertheless, we were able to identify subtype specific subsets of significantly differentially expressed transcripts which link resource acquisition and allocation to NADP-ME species susceptibility to glacial CO2 . Here, low light phenotypes were comparable across species with no clear subtype response, while again, transcriptomic responses to low light were largely species specific. However, numerous functional similarities were noted within the transcriptomic responses to low light, suggesting these responses are functionally relatively conserved. Additionally, PEPCK species exhibited heightened regulation of transcripts related to metabolism in response to both stresses, likely tied to their C4 metabolic pathway. These results highlight the influence that both species and subtype can have on plant responses to abiotic stress, building on our mechanistic understanding of acclimation within C4 grasses and highlighting avenues for future crop improvements.

Mehler reaction plays a role in C3 and C4 photosynthesis under shade and low CO2.

Alternative electron fluxes such as the cyclic electron flux (CEF) around photosystem I (PSI) and Mehler reaction (Me) are essential for efficient photosynthesis because they generate additional ATP and protect both photosystems against photoinhibition. The capacity for Me can be estimated by measuring O2 exchange rate under varying irradiance and CO2 concentration. In this study, mass spectrometric measurements of O2 exchange were made using leaves of representative species of C3 and C4 grasses grown under natural light (control; PAR ~ 800 µmol quanta m-2 s-1) and shade (~ 300 µmol quanta m-2 s-1), and in representative species of gymnosperm, liverwort and fern grown under natural light. For all control grown plants measured at high CO2, O2 uptake rates were similar between the light and dark, and the ratio of Rubisco oxygenation to carboxylation (Vo/Vc) was low, which suggests little potential for Me, and that O2 uptake was mainly due to photorespiration or mitochondrial respiration under these conditions. Low CO2 stimulated O2 uptake in the light, Vo/Vc and Me in all species. The C3 species had similar Vo/Vc, but Me was highest in the grass and lowest in the fern. Among the C4 grasses, shade increased O2 uptake in the light, Vo/Vc and the assimilation quotient (AQ), particularly at low CO2, whilst Me was only substantial at low CO2 where it may contribute 20-50% of maximum electron flow under high light.

Smart glass impacts stomatal sensitivity of greenhouse Capsicum through altered light.

Optical films that alter light transmittance may reduce energy consumption in high-tech greenhouses, but their impact on crop physiology remains unclear. We compared the stomatal responses of Capsicum plants grown hydroponically under control glass (70% diffuse light) or the smart glass (SG) film ULR-80, which blocked >50% of short-wave radiation and ~9% of photosynthetically active radiation (PAR). SG had no significant effects on steady-state (gs) or maximal (gmax) stomatal conductance. In contrast, SG reduced stomatal pore size and sensitivity to exogenous abscisic acid (ABA), thereby increasing rates of leaf water loss, guard cell K+ and Cl- efflux, and Ca2+ influx. SG induced faster stomatal closing and opening rates on transition between low (100 µmol m-2 s-1) and high PAR (1500 µmol m-2 s-1), which compromised water use efficiency relative to control plants. The fraction of blue light (0% or 10%) did not affect gs in either treatment. Increased expression of stomatal closure and photoreceptor genes in epidermal peels of SG plants is consistent with fast stomatal responses to light changes. In conclusion, stomatal responses of Capsicum to SG were more affected by changes in light intensity than spectral quality, and re-engineering of the SG should maximize PAR transmission, and hence CO2 assimilation.

Sugar sensing responses to low and high light in leaves of the C4 model grass Setaria viridis.

Although sugar regulates photosynthesis, the signalling pathways underlying this process remain elusive, especially for C4 crops. To address this knowledge gap and identify potential candidate genes, we treated Setaria viridis (C4 model) plants acclimated to medium light intensity (ML, 500 µmol m-2 s-1) with low (LL, 50 µmol m-2 s-1) or high (HL, 1000 µmol m-2 s-1) light for 4 d and observed the consequences on carbon metabolism and the transcriptome of source leaves. LL impaired photosynthesis and reduced leaf content of signalling sugars (glucose, sucrose, and trehalose-6-phosphate). In contrast, HL strongly induced sugar accumulation without repressing photosynthesis. LL more profoundly impacted the leaf transcriptome, including photosynthetic genes. LL and HL contrastingly altered the expression of hexokinase (HXK) and sucrose-non-fermenting 1 (Snf1)-related protein kinase 1 (SnRK1) sugar sensors and trehalose pathway genes. The expression of key target genes of HXK and SnRK1 were affected by LL and sugar depletion, while surprisingly HL and strong sugar accumulation only slightly repressed the SnRK1 signalling pathway. In conclusion, we demonstrate that LL profoundly impacted photosynthesis and the transcriptome of S. viridis source leaves, while HL altered sugar levels more than LL. We also present the first evidence that sugar signalling pathways in C4 source leaves may respond to light intensity and sugar accumulation differently from C3 source leaves.

The role of leaf width and conductances to CO2 in determining water use efficiency in C4 grasses.

C4 plants achieve higher photosynthesis (An ) and intrinsic water use efficiency (iWUE) than C3 plants, but processes underpinning the variability in An and iWUE across the three C4 subtypes remain unclear, partly because we lack an integrated framework for quantifying the contribution of diffusional and biochemical limitations to C4 photosynthesis. We exploited the natural diversity among C4 grasses to develop an original mathematical approach for estimating eight key processes of C4 photosynthesis and their relative limitations to An . We also developed a new formulation to estimate mesophyll conductance (gm ) based on actual hydration rates of CO2 by carbonic anhydrases. We found a positive relationship between gm and iWUE and an inverse correlation with gsw among C4 grasses. We also revealed subtype-specific regulatory processes of iWUE that may be related to known anatomical traits characterising each C4 subtype. Leaf width was an important determinant of iWUE and showed significant correlations with key limitations of An , especially among NADP-ME species. In conclusion, incorporating leaf width in breeding trials may unlock new opportunities for C4 crops because the revealed negative relationship between leaf width and iWUE may translate into higher crop and canopy WUE.

Responses of respiration in the light to warming in field-grown trees: a comparison of the thermal sensitivity of the Kok and Laisk methods.

The Kok and Laisk techniques can both be used to estimate light respiration Rlight . We investigated whether responses of Rlight to short- and long-term changes in leaf temperature depend on the technique used to estimate Rlight . We grew Eucalyptus tereticornis in whole-tree chambers under ambient temperature (AT) or AT + 3°C (elevated temperature, ET). We assessed dark respiration Rdark and light respiration with the Kok (RKok ) and Laisk (RLaisk ) methods at four temperatures to determine the degree of light suppression of respiration using both methods in AT and ET trees. The ET treatment had little impact on Rdark , RKok or RLaisk . Although the thermal sensitivities of RKok or RLaisk were similar, RKok was higher than RLaisk . We found negative values of RLaisk at the lowest measurement temperatures, indicating positive net CO2 uptake, which we propose may be related to phosphoenolpyruvate carboxylase activity. Light suppression of Rdark decreased with increasing leaf temperature, but the degree of suppression depended on the method used. The Kok and Laisk methods do not generate the same estimates of Rlight or light suppression of Rdark between 20 and 35°C. Negative rates of RLaisk imply that this method may become less reliable at low temperatures.

Acclimation and adaptation components of the temperature dependence of plant photosynthesis at the global scale.

The temperature response of photosynthesis is one of the key factors determining predicted responses to warming in global vegetation models (GVMs). The response may vary geographically, owing to genetic adaptation to climate, and temporally, as a result of acclimation to changes in ambient temperature. Our goal was to develop a robust quantitative global model representing acclimation and adaptation of photosynthetic temperature responses. We quantified and modelled key mechanisms responsible for photosynthetic temperature acclimation and adaptation using a global dataset of photosynthetic CO2 response curves, including data from 141 C3 species from tropical rainforest to Arctic tundra. We separated temperature acclimation and adaptation processes by considering seasonal and common-garden datasets, respectively. The observed global variation in the temperature optimum of photosynthesis was primarily explained by biochemical limitations to photosynthesis, rather than stomatal conductance or respiration. We found acclimation to growth temperature to be a stronger driver of this variation than adaptation to temperature at climate of origin. We developed a summary model to represent photosynthetic temperature responses and showed that it predicted the observed global variation in optimal temperatures with high accuracy. This novel algorithm should enable improved prediction of the function of global ecosystems in a warming climate.

Cyclic electron flow and light partitioning between the two photosystems in leaves of plants with different functional types.

Cyclic electron flow (CEF) around photosystem I (PSI) is essential for generating additional ATP and enhancing efficient photosynthesis. Accurate estimation of CEF requires knowledge of the fractions of absorbed light by PSI (fI) and PSII (fII), which are only known for a few model species such as spinach. No measures of fI are available for C4 grasses under different irradiances. We developed a new method to estimate (1) fII in vivo by concurrently measuring linear electron flux through both photosystems [Formula: see text] in leaf using membrane inlet mass spectrometry (MIMS) and total electron flux through PSII (ETR2) using chlorophyll fluorescence by a Dual-PAM at low light and (2) CEF as ETR1-[Formula: see text]. For a C3 grass, fI was 0.5 and 0.4 under control (high light) and shade conditions, respectively. C4 species belonging to NADP-ME and NAD-ME subtypes had fI of 0.6 and PCK subtype had 0.5 under control. All shade-grown C4 species had fI of 0.6 except for NADP-ME grass which had 0.7. It was also observed that fI ranged between 0.3 and 0.5 for gymnosperm, liverwort and fern species. CEF increased with irradiance and was induced at lower irradiances in C4 grasses and fern relative to other species. CEF was greater in shade-grown plants relative to control plants except for C4 NADP-ME species. Our study reveals a range of CEF and fI values in different plant functional groups. This variation must be taken into account for improved photosynthetic calculations and modelling.

Elevated CO2 alleviates the negative impact of heat stress on wheat physiology but not on grain yield.

Hot days are becoming hotter and more frequent, threatening wheat yields worldwide. Developing wheat varieties ready for future climates calls for improved understanding of how elevated CO2 (eCO2) and heat stress (HS) interactively impact wheat yields. We grew a modern, high-yielding wheat cultivar (Scout) at ambient CO2 (aCO2, 419 µl l -1) or eCO2 (654 µl l-1) in a glasshouse maintained at 22/15 °C (day/night). Half of the plants were exposed to HS (40/24 °C) for 5 d at anthesis. In non-HS plants, eCO2 enhanced (+36%) CO2 assimilation rates (Asat) measured at growth CO2 despite down-regulation of photosynthetic capacity. HS reduced Asat (-42%) in aCO2- but not in eCO2-grown plants because eCO2 protected photosynthesis by increasing ribulose bisphosphate regeneration capacity and reducing photochemical damage under HS. eCO2 stimulated biomass (+35%) of all plants and grain yield (+30%) of non-HS plants only. Plant biomass initially decreased following HS but recovered at maturity due to late tillering. HS equally reduced grain yield (-40%) in aCO2- and eCO2-grown plants due to grain abortion and reduced grain filling. While eCO2 mitigated the negative impacts of HS at anthesis on wheat photosynthesis and biomass, grain yield was reduced by HS in both CO2 treatments.