Phytohormone-Mediated Regulation of Plant Cold Stress Tolerance: Signaling, Hormonal Crosstalk, and Translational Perspectives

Cold stress (CS) represents a major environmental factor that adversely affects plant growth, development, and productivity. To cope with low-temperature conditions, plants have evolved sophisticated mechanisms for CS perception and response, mediated through complex cellular signaling networks and physiological processes. Central to these adaptive responses are phytohormones, which function either independently or through synergistic and antagonistic interactions to fine-tune CS tolerance. This review synthesizes current knowledge on the roles of major classical phytohormones and signaling metabolites in regulating CS tolerance in plants. We first outline the molecular mechanisms involved in CS sensing and signal transduction, highlighting the roles of membrane-associated sensors, calcium signaling, and downstream transcriptional networks. Then, we discuss the contributions of key classical phytohormones, including auxin, abscisic acid, ethylene, salicylic acid, cytokinin, jasmonic acid, brassinosteroids, gibberellic acid, strigolactones, and signaling metabolites, including melatonin and gamma-aminobutyric acid, to CS tolerance, highlighting their individual and interacting roles in modulating gene expression regulation, antioxidant defense and physiological adaptations. We also discuss the crosstalk between these hormones, emphasizing the dynamic and often context-dependent nature of their interactions in response to CS. Furthermore, the review highlights recent advances in CRISPR/Cas9-based genome editing strategies targeting phytohormone biosynthesis, signaling, and response pathways to improve CS tolerance in plants. By integrating hormonal signaling, molecular regulation, and modern biotechnological tools, this review provides a comprehensive framework for understanding phytohormone-mediated CS adaptation and offers perspectives for developing climate-resilient crops through genetic and agronomic approaches.