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Sandalwood ( Santalum album  L.) is highly valued for its fragrant heartwood and extracted oil. Santalols, which are the main components of that oil, are terpenoids, and these are biosynthesized via the mevalonic acid (MVA) pathway. Mevalonate kinase (MK) and phosphomevalonate kinase (PMK) are key enzymes in the MVA pathway. Little is known about the genes that encode MK and PMK in  S. album  or the mechanism that regulates their expression. To isolate and identify the functional genes involved in santalol biosynthesis in  S. album , an  MK  gene designated as  SaMK , and a  PMK  gene designated as  SaPMK , were cloned from  S. album . The sequences of these genes were analyzed. A bioinformatics analysis was conducted to assess the homology of  SaMK  and  SaPMK  with  MK  and  PMK  genes from other plants. The subcellular localization of  SaMK  and  SaPMK  proteins was also investigated, as was the functional complementation of  SaMK  and  SaPMK  in yeast. Our results show that the full-length cDNA sequences of  SaMK  and  SaPMK  were 1409 bp and 1679 bp long, respectively.  SaMK  contained a 1381 bp open reading frame (ORF) encoding a polypeptide of 460 amino acids and  SaPMK  contained a 1527 bp ORF encoding a polypeptide of 508 amino acids.  SaMK  and  SaPMK  showed high homology with  MK  and  PMK  genes of other plant species. Functional complementation of  SaMK  in a MK-deficient mutant yeast strain YMR208W and  SaPMK  in a PMK-deficient mutant yeast strain YMR220W confirmed that cloned  SaMK  and  SaPMK  cDNA encode a functional MK and PMK, respectively, mediating MVA biosynthesis in yeast. An analysis of tissue expression patterns revealed that  SaMK  and  SaPMK  were constitutively expressed in all the tested tissues.  SaMK  was highly expressed in young leaves but weakly expressed in sapwood.  SaPMK  was highly expressed in roots and mature leaves, but weakly expressed in young leaves. Induction experiments with several elicitors showed that  SaMK  and  SaPMK  expression was upregulated by methyl jasmonate. These results will help to further study the role of  MK  and  PMK  genes during santalol biosynthesis in  S. album .

Cloning and expression analysis of mevalonate kinase and phosphomevalonate kinase genes associated with the MVA pathway in Santalum album