SHINE clade of ERF transcription factors: A significant player in abiotic and biotic stress tolerance in plants

SHINE (SHN) clade transcription factors (TFs) represents a subfamily of APETALA2/ethylene-responsive factor (AP2/ERF) proteins. The latter, is characterized by its responsiveness to the phytohormone ethylene and the presence of AP2 DNA-binding domain. They are involved in many biological processes and in responses to different environmental constraints. SHN TFs were among the first identified regulators of cuticle formation. Cuticle plays crucial role in plant tolerance to drought, salinity and high temperature as well as in defense against pathogens. In addition, SHN were shown to be involved in the regulation of stomatal development which influences resistance to drought and diseases. Interestingly, recent studies have also shown that SHN TFs are involved in mediating the beneficial effects of arbuscular mycorrhizal fungi (AMF) as well as disease resistance conferred by nanoparticles. To fulfill their roles, SHN TFs are controlled upstream by other TFs and they control, in their turn, different downstream genes. In this review, we highlight the role of SHN TFs in different abiotic and biotic stresses through their involvement in cuticle biosynthesis, stomatal development and molecular regulation of biochemical and physiological traits. In addition, we discuss the regulation of SHN TFs by plant hormones and their influence on hormone biosynthesis and signaling pathways. Knowledge of this complex regulation can be put into contribution to increase multiple abiotic stress tolerances through transgenesis, gene editing and classical breeding.