Identification and Characterization of the AREB/ABF/ABI5 Gene Family in Sandalwood (Santalum album L.) and Its Potential Role in Drought Stress and ABA Treatment

AREB/ABF/ABI5 (ABA-responsive element-binding protein/ABRE binding factors and ABA INSENSITIVE 5) transcription factors are involved in regulating the expression of ABA (abscisic acid)-related genes and improving plant adaptability to environmental stress. To explore the influence of AREB/ABF transcription factors on santalol synthesis, we conducted a genome-wide analysis of the AREB gene family in sandalwood, identified 10 SaAREB genes, and divided them into five subfamilies. We found that all SaAREB genes encoded unstable hydrophilic proteins and the subcellular localization prediction of SaAREBs was that they are located in the nucleus. AREB/ABF genes belong to the bZIP-A subfamily and we found that the 10 AREB proteins all contained bZIP (basic region leucine zipper) and four potential phosphorylation sites (RXXS/T). According to the collinearity analysis results, four of the SaAREB genes were involved in two fragment duplication events. Through qRT-PCR (real-time fluorescence quantitative PCR), we explored the expression profile of SaAREB in different tissues; the effects of ABA treatment and drought treatment on AREB transcription factors were predicted. From the expression of different tissues, we found that SaAREB1 not only responded to prolonged drought but also was highly expressed in stems. Moreover, SaAREB3, SaAREB7, and SaAREB8 specifically respond to ABA treatment. Based on RNA-seq (RNA sequencing) data, we found that SaAREB6 and SaAREB8 were highly expressed in the sapwood and transition regions. Regarding SaCYP736A167, as a key gene in santalol synthesis, its promoter contains the most ABRE cis-reactive elements. These results provide a basis for further analysis of the role of the Santalum album L. (S. album) ABRE/ABF/ABI5 genes in the formation of santalols.