Enhancing Tolerance to Combined Heat and Drought Stress in Cool-Season Grain Legumes: Mechanisms, Genetic Insights, and Future Directions

The increasing frequency of concurrent heat and drought stress poses a significant challenge to agricultural productivity, particularly for cool-season grain legumes, including broad bean (Vicia Faba L.), lupin (Lupinus spp.), lentil (Lens culinaris Medik), chickpea (Cicer arietinum L.), grasspea (Lathyrus sativus L.), pea (Pisum sativum L.), and common vetch (Vicia sativa L.). These legumes play a vital role in sustainable agricultural systems due to their nitrogen-fixing ability and high nutritional value. This review synthesizes current knowledge of the impacts and tolerance mechanisms associated with combined heat and drought stresses in these crops. We evaluate physiological and biochemical responses to combined heat and drought stress, focusing on their detrimental effects on growth, development, and yield. Key genetic and molecular mechanisms, such as the roles of osmolytes, antioxidants, and stress-responsive genes, are explored. We also discuss the intricate interplay between heat and drought stress signaling pathways, including the involvement of Ca²⁺ ions, reactive oxygen species, transcription factor DREB2A, and the endoplasmic reticulum in mediating stress responses. This comprehensive analysis offers new insights into developing resilient legume varieties to enhance agricultural sustainability under climate change. Future research should prioritize integrating omics technologies to unravel plant responses to combined abiotic stresses.