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Subclass III members of the sucrose non-fermenting-1-related protein kinase 2 (SnRK2) play essential roles in both the abscisic acid signaling and abiotic stress responses of plants by phosphorylating the downstream ABA-responsive element (ABRE)-binding proteins (AREB/ABFs). This comprehensive study investigated the function of new candidate genes, namely  SmSnRK2.3 ,  SmSnRK2.6 , and  SmAREB1 , with a view to breeding novel varieties of  Salvia miltiorrhiza  with improved stress tolerance stresses and more content of bioactive ingredients. Exogenous ABA strongly induced the expression of these genes. PlantCARE predicted several hormones and stress response  cis -elements in their promoters.  SmSnRK2.6  and  SmAREB1  showed the highest expression levels in the leaves of  S. miltiorrhiza  seedlings, while  SmSnRK2.3  exhibited a steady expression in their roots, stems, and leaves. A subcellular localization assay revealed that both  SmSnRK2.3  and  SmSnRK2.6  were located in the cell membrane, cytoplasm, and nucleus, whereas  SmAREB1  was exclusive to the nucleus. Overexpressing  SmSnRK2.3  did not significantly promote the accumulation of rosmarinic acid (RA) and salvianolic acid B (Sal B) in the transgenic  S. miltiorrhiza  hairy roots. However, overexpressing  SmSnRK2.6  and  SmAREB1  increased the contents of RA and Sal B, and regulated the expression levels of structural genes participating in the phenolic acid-branched and side-branched pathways, including  SmPAL1 ,  SmC4H ,  Sm4CL1 ,  SmTAT ,  SmHPPR ,  SmRAS ,  SmCHS ,  SmCCR ,  SmCOMT , and  SmHPPD . Furthermore, SmSnRK2.3 and SmSnRK2.6 interacted physically with SmAREB1. In summary, our results indicate that  SmSnRK2.6  is involved in stress responses and can regulate structural gene transcripts to promote greater metabolic flux to the phenolic acid-branched pathway, via its interaction with  SmAREB1 , a transcription factor. In this way,  SmSnRK2.6  contributes to the positive regulation of phenolic acids in  S. miltiorrhiza  hairy roots.

The Protein Kinase SmSnRK2.6 Positively Regulates Phenolic Acid Biosynthesis in Salvia miltiorrhiza by Interacting with SmAREB1