Brassinosteroids as a multidimensional regulator of plant physiological and molecular responses under various environmental stresses

Biotic and abiotic stresses, especially heavy metal toxicity, are becoming a big problem in agriculture, which pose serious threats to crop production. Plant hormones have recently been used to develop stress tolerance in a variety of plants. Brassinosteroids (BRs) are the sixth class of plant steroid hormones, with pleiotropic effects on plants. Exogenous application of BRs to boost plant tolerance mechanisms to various stresses has been a major research focus. Numerous studies have revealed the role of these steroidal hormones in the up-regulation of stress-related resistance genes, as well as their interactions with other metabolic pathways. BRs interact with other phytohormones such as auxin, cytokinin, ethylene, gibberellin, jasmonic acid, abscisic acid, salicylic acid, and polyamines to regulate a variety of physiological and developmental processes in plants. BRs regulate expressions of many BR-inducible genes by activating the brassinazole-resistant 1 (BZR1)/BRI1-EMS suppressor 1 (BES1) complex. Moreover, to improve plant development under a variety of stresses, BRs regulate antioxidant enzyme activity, chlorophyll concentration, photosynthetic capability, and glucose metabolism. This review will provide insights into the mechanistic role and actions of brassinosteroids in plants in response to various stresses.