Anatomical determinants of gas exchange and hydraulics vary with leaf shape in soybean.

BACKGROUND AND AIMS: Leaf shape in crops can impact light distribution and carbon capture at the whole plant and canopy level. Given similar leaf inclination, narrow leaves can allow a greater fraction of incident light to pass through to lower canopy leaves by reducing leaf area index, which can potentially increase canopy-scale photosynthesis. Soybean has natural variation in leaf shape which can be utilized to optimize canopy architecture. However, the anatomical and physiological differences underlying variation in leaf shape remain largely unexplored. METHODS: In this study, we selected 28 diverse soybean lines with leaf length to width ratios (leaf ratio) ranging between 1.1 and 3.2. We made leaf cross-sectional, gas exchange, vein density and hydraulic measurements and studied their interrelationships among these lines. KEY RESULTS: Our study shows that narrow leaves tend to be thicker, with an ~30 µm increase in leaf thickness for every unit increase in leaf ratio. Interestingly, thicker leaves had a greater proportion of spongy mesophyll while the proportions of palisade and paraveinal mesophyll decreased. In addition, narrow and thicker leaves had greater photosynthesis and stomatal conductance per unit area along with greater leaf hydraulic conductance. CONCLUSIONS: Our results suggest that selecting for narrow leaves can improve photosynthetic performance and potentially provide a yield advantage in soybean.

The genetic basis of transpiration sensitivity to vapor pressure deficit in wheat.

Genetic manipulation of whole-plant transpiration rate (TR) response to increasing atmospheric vapor pressure deficit (VPD) is a promising approach for crop adaptation to various drought regimes under current and future climates. Genotypes with a non-linear TR response to VPD are expected to achieve yield gains under terminal drought, thanks to a water conservation strategy, while those with a linear response exhibit a consumptive strategy that is more adequate for well-watered or transient-drought environments. In wheat, previous efforts indicated that TR has a genetic basis under naturally fluctuating conditions, but because TR is responsive to variation in temperature, photosynthetically active radiation, and evaporative demand, the genetic basis of its response VPD per se has never been isolated. To address this, we developed a controlled-environment gravimetric phenotyping approach where we imposed VPD regimes independent from other confounding environmental variables. We screened three nested association mapping populations totaling 150 lines, three times over a 3-year period. The resulting dataset, based on phenotyping nearly 1400 plants, enabled constructing 63-point response curves for each genotype, which were subjected to a genome-wide association study. The analysis revealed a hotspot for TR response to VPD on chromosome 5A, with SNPs explaining up to 17% of the phenotypic variance. The key SNPs were found in haploblocks that are enriched in membrane-associated genes, consistent with the hypothesized physiological determinants of the trait. These results indicate a promising potential for identifying new alleles and designing next-gen wheat cultivars that are better adapted to current and future drought regimes.

Overlapping and stress-specific transcriptomic and hormonal responses to flooding and drought in soybean.

Flooding and drought are serious constraints that reduce crop productivity worldwide. Previous studies identified genes conferring tolerance to both water extremes in various plants. However, overlapping responses to flooding and drought at the genome-scale remain obscure. Here, we defined overlapping and stress-specific transcriptomic and hormonal responses to submergence, drought and recovery from these stresses in soybean (Glycine max). We performed comparative RNA-sequencing and hormone profiling, identifying genes, hormones and biological processes that are differentially regulated in an overlapping or stress-specific manner. Overlapping responses included positive regulation of trehalose and sucrose metabolism and negative regulation of cellulose, tubulin, photosystem II and I, and chlorophyll biosynthesis, facilitating the economization of energy reserves under both submergence and drought. Additional energy-consuming pathways were restricted in a stress-specific manner. Downregulation of distinct pathways for energy saving under each stress suggests energy-consuming processes that are relatively unnecessary for each stress adaptation are turned down. Our newly developed transcriptomic-response analysis revealed that abscisic acid and ethylene responses were activated in common under both stresses, whereas stimulated auxin response was submergence-specific. The energy-saving strategy is the key overlapping mechanism that underpins adaptation to both submergence and drought in soybean. Abscisic acid and ethylene are candidate hormones that coordinate transcriptomic energy-saving processes under both stresses. Auxin may be a signaling component that distinguishes submergence-specific regulation of the stress response.

Sheathing the blade: Significant contribution of sheaths to daytime and nighttime gas exchange in a grass crop.

Despite representing a sizeable fraction of the canopy, very little is known about leaf sheath gas exchange in grasses. Specifically, estimates of sheath stomatal conductance, transpiration and photosynthesis along with their responses to light, CO2 and vapour pressure deficit (VPD) are unknown. Furthermore, the anatomical basis of these responses is poorly documented. Here, using barley as a model system, and combining leaf-level gas exchange, whole-plant gravimetric measurements, transpiration inhibitors, anatomical observations, and biophysical modelling, we found that sheath and blade stomatal conductance and transpiration were similar, especially at low light, in addition to being genotypically variable. Thanks to high abaxial stomata densities and surface areas nearly half those of the blades, sheaths accounted for up to 17% of the daily whole-plant water use, which -surprisingly- increased to 45% during the nighttime. Sheath photosynthesis was on average 17-25% that of the blade and was associated with lower water use efficiency. Finally, sheaths responded differently to the environment, exhibiting a lack of response to CO2 but a strong sensitivity to VPD. Overall, these results suggest a key involvement of sheaths in feedback loops between canopy architecture and gas exchange with potentially significant implications on adaptation to current and future climates in grasses.

Variability in temperature-independent transpiration responses to evaporative demand correlate with nighttime water use and its circadian control across diverse wheat populations.

MAIN CONCLUSION: Nocturnal transpiration, through its circadian control, plays a role in modulating daytime transpiration response to increasing evaporative demand, to potentially enable drought tolerance in wheat. Limiting plant transpiration rate (TR) in response to increasing vapor pressure deficit (VPD) has been suggested to enable drought tolerance through water conservation. However, there is very little information on the extent of diversity of TR response curves to "true" VPD (i.e., independent from temperature). Furthermore, new evidence indicate that water-saving could operate by modulating nocturnal TR (TRN), and that this response might be coupled to daytime gas exchange. Based on 3 years of experimental data on a diverse group of 77 genotypes from 25 countries and 5 continents, a first goal of this study was to characterize the functional diversity in daytime TR responses to VPD and TRN in wheat. A second objective was to test the hypothesis that these traits could be coupled through the circadian clock. Using a new gravimetric phenotyping platform that allowed for independent temperature and VPD control, we identified three and fourfold variation in daytime and nighttime responses, respectively. In addition, TRN was found to be positively correlated with slopes of daytime TR responses to VPD, and we identified pre-dawn variation in TRN that likely mediated this relationship. Furthermore, pre-dawn increase in TRN positively correlated with the year of release among drought-tolerant Australian cultivars and with the VPD threshold at which they initiated water-saving. Overall, the study indicates a substantial diversity in TR responses to VPD that could be leveraged to enhance fitness under water-limited environments, and that TRN and its circadian control may play an important role in the expression of water-saving.

Physiological and transcriptomic characterization of submergence and reoxygenation responses in soybean seedlings.

Complete inundation at the early seedling stage is a common environmental constraint for soybean production throughout the world. As floodwaters subside, submerged seedlings are subsequently exposed to reoxygenation stress in the natural progression of a flood event. Here, we characterized the fundamental acclimation responses to submergence and reoxygenation in soybean at the seedling establishment stage. Approximately 90% of seedlings succumbed during 3 d of inundation under constant darkness, whereas 10 d of submergence were lethal to over 90% of seedlings under 12 h light/12 h dark cycles, indicating the significance of underwater photosynthesis in seedling survival. Submergence rapidly decreased the abundance of carbohydrate reserves and ATP in aerial tissue of seedlings although chlorophyll breakdown was not observed. The carbohydrate and ATP contents were recovered upon de-submergence, but sudden exposure to oxygen also induced lipid peroxidation, confirming that reoxygenation induced oxidative stress. Whole transcriptome analysis recognized genome-scale reconfiguration of gene expression that regulates various signalling and metabolic pathways under submergence and reoxygenation. Comparative analysis of differentially regulated genes in shoots and roots of soybean and other plants defines conserved, organ-specific and species-specific adjustments which enhance adaptability to submergence and reoxygenation through different metabolic pathways.