Field-based dissection of stomatal anatomy and conductance reveals stable QTL under drought and heat in wheat

2026 · doi:10.64898/2026.03.30.715413
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This paper examined how wheat stomatal physiological and anatomical traits, along with yield, respond under water-limitation and delayed sowing–induced heat exposure using two years of large-scale field trials and genome-wide association mapping. Stomatal conductance declined under both stresses (water-limitation and heat), with the adaxial leaf surface dominating genotype variation and showing the strongest stress responsiveness; despite increased theoretical anatomical gas-exchange capacity, stomatal conductance operating efficiency decreased, and anatomical plasticity differed by stress (drought reduced stomatal size and increased density, while heat increased size). The authors reported moderate-to-high heritability for anatomical traits and lower, less stable heritability for stomatal conductance, and they identified 169 putative QTLs—mostly for anatomical traits— including stable co-localised pleiotropic loci on chromosomes 2B, 3B, and 7B across seasons/studies, with the main caveat being variability in heritability and stability across different trait types and environments. The paper does not explicitly discuss endometriosis or adenomyosis; it was included in the corpus via a keyword match in the upstream search index.

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Abstract Stomatal traits balance carbon gain with water loss, yet their breeding potential in wheat remains underexploited. This study investigated physiological and anatomical stomatal responses alongside yield across two years of large-scale field trials under water-limitation and delayed sowing-induced heat exposure. Across both seasons, stomatal conductance (gs) declined under stress, reflecting strong environmental constraint on gas-exchange (water-limitation:-26.9%; heat:-13.8%). Partitioning responses by leaf surface and genotype identified the adaxial surface as the dominant contributor to gs variation and the most stress responsive. Despite increases in theoretical anatomical gas-exchange capacity (gsmax), gs-efficiency declined, indicating partial decoupling between structural potential and realised conductance. Drought reduced stomatal size while increasing density whereas heat increased size, suggesting stress-specific anatomical plasticity. Moderate-to-high heritability was observed for anatomical traits (Water-limitation: 0.13-0.57; Heat: 0.42-0.71), contrasting with lower and less stable heritability for gs (water-limitation: 0.13-0.41; heat: 0.13-0.50). Genome-wide-association-mapping identified 169 putative QTLs, predominantly for anatomical traits, including stable and co-localised pleiotropic loci. Fourteen sets of closely positioned markers were detected across seasons or studies, with stable regions on chromosomes 2B, 3B and 7B emerging as key loci. Focusing on stable loci controlling adaxial stomatal anatomy offers a realistic strategy to enhance wheat photosynthetic efficiency and climate resilience. Highlight Adaxial stomatal traits dominate gas exchange responses to heat and drought in wheat, with stable anatomical QTL identified on chromosomes 2B, 3B and 7B. Their stability across environments supports their relevance for crop improvement in water-limited and high temperature systems. Competing Interest Statement The authors have declared no competing interest. Footnotes Addition of Supplementary Figures and Tables File. Abbreviations - CAIGE - CIMMYT Australia ICARDA Germplasm Evaluation - CIMMYT - International Maize and Wheat Improvement Center - CO2 - Carbon Dioxide - EDPIE - Elite Diversity International Experiment - ESWYT - Elite Selection Wheat Yield Trial - GCL - Guard Cell Length - GCW - Guard Cell Width - gs - Stomatal conductance - gse - gsop/gsmax - Stomatal conductance operating efficiency (unitless) - gsmax - Maximum anatomical stomatal conductance - gsop - Operating rate of stomatal conductance - H2O - Water - HeDWIC - Heat and Drought Wheat Improvement Consortium Project - HTWYT - High Temperature Wheat Yield Trials - ICARDA - International Centre for Agricultural Research in the Dry Areas - IRGA - Infra-red gas analysers - mAP - Mean average precision - Mbp - Mega base pair - LOD - Logarithm of the odds - PSII - Photosystem II - QTL - Quantitative Trait Loci - RH - Relative Humidity - SATYN - Stress Adapted Trait Yield Nursery - SAWYT - Semi-Arid Wheat Yield Trial - S1 - Season 1; 2023 - S2 - Season 2; 2024 - SA - Stomatal Area - SD - Stomatal density (stomata m-2) - TKW - Thousand Kernel Weight - TOS - Time of Sowing - VPD - Vapour Pressure Deficit - WUE - Water Use Efficiency - YOLO - You Only Look Once

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