From stress signaling to yield stability: physiological and molecular mechanisms of wheat resilience to heat and drought stress.

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conclusionWheat resilience depends on coordinated signaling, reproductive protection, and source-sink regulation, providing a framework to breed robust trait combinations that stabilize yield under combined heat and drought. Climate change is increasing the frequency and severity of heat and drought events, posing a major threat to wheat productivity, yield stability, and food security. Because these stresses often coincide in the field, their combined effects can impair growth, reproductive development, grain filling, and final yield more severely than either stress alone. Wheat resilience under such conditions depends on coordinated physiological adjustment and molecular regulation that sustain cellular homeostasis, protect reproductive tissues, and preserve yield-related traits. This review synthesizes current knowledge on the physiological and molecular bases of wheat resilience to heat and drought, with emphasis on their combined effects. We discuss major physiological responses, including photosynthetic adjustment, stomatal regulation, canopy cooling, osmotic balance, antioxidant defense, membrane stability, and source-sink coordination. We also examine key regulatory pathways involved in stress perception and adaptation, including calcium and reactive oxygen species signaling, mitogen-activated protein kinase cascades, phytohormonal crosstalk, transcriptional regulation, heat shock proteins, late embryogenesis abundant proteins, and osmoprotective and redox-associated pathways. In addition, we highlight the growing contribution of transcriptomics, proteomics, metabolomics, and phenomics to the identification of candidate genes, biomarkers, and adaptive traits. Finally, we consider how mechanistic insights can be translated into wheat improvement through molecular markers, genomic selection, gene editing, and climate-realistic phenotyping. An integrated understanding of stress signaling and adaptive trait deployment will be essential for developing wheat cultivars with improved resilience and yield stability under future climates.