Pivotal role of the AREB/ABF-SnRK2 pathway in ABRE-mediated transcription in response to osmotic stress in plants

Water availability is one of the main limiting factors for plant growth and development. The phytohormone abscisic acid (ABA) fulfills a critical role in coordinating the responses to reduced water availability as well as in multiple developmental processes. Endogenous ABA levels increase in response to osmotic stresses such as drought and high salinity, and ABA activates the expression of many genes via ABA-responsive elements (ABREs) in their promoter regions. ABRE-binding protein/ABRE-binding factor (AREB/ABF) transcription factors (TFs) regulate the ABRE-mediated transcription of downstream target genes. Three subclass III sucrose non-fermenting-1 related protein kinase 2 (SnRK2) protein kinases (SRK2D/SnRK2.2, SRK2E/SnRK2.6/OST1 and SRK2I/SnRK2.3) phosphorylate and positively control the AREB/ABF TFs. Substantial progress has been made in our understanding of the ABA-sensing system mediated by Pyrabactin resistance1/PYR1-like/regulatory components of ABA receptor (PYR/PYL/RCAR)-protein phosphatase 2C complexes. In addition to PP2C-PYR/PYL/RCAR ABAreceptor complex, the AREB/ABF-SnRK2 pathway, which is well conserved in land plants, was recently shown to play a major role as a positive regulator of ABA/stress signaling through ABRE-mediated transcription of target genes implicated in the osmotic stress response. This review focuses on current progress in the study of the AREB/ABF-SnRK2 positive regulatory pathway in plants and describes additional signaling factors implicated in the AREB/ABF-SnRK2 pathway. Moreover, to help promote the link between basic and applied studies, the nomenclature and phylogenetic relationships between the AREB/ABFs and SnRK2s are summarized and discussed.