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Anthocyanins are biosynthesized on the cytosolic surface of the endoplasmic reticulum and then transported into the vacuole for storage. Glutathione  S -transferases (GSTs) are considered to be responsible for the transport of anthocyanins into the vacuole. However, the regulatory mechanisms of GSTs in plants are still unclear. Here, we performed a genome-wide analysis and identified 69  GST  genes in apple. The expression of  MdGSTF6  was positively correlated with the anthocyanin content ( r  = 0.949) during ‘Yanfu 8’ fruit development. The overexpression of  MdGSTF6  in the  Arabidopsis thaliana tt19  mutant resulted in seedlings of 35S:: MdGSTF6 -GFP/ tt19  that could accumulate anthocyanin and rescue its phenotype, suggesting that MdGSTF6 was an anthocyanin transporter. The silencing of  MdGSTF6  affected anthocyanin accumulation in apple fruit. Moreover, the knockdown of  MdGSTF6  by RNA interference in cultured ‘Gala’ seedlings inhibited anthocyanin accumulation. The interaction experiments showed that MdMYB1 could bind directly to the  MdGSTF6  promoter to transcriptionally activate its expression. Collectively, our results demonstrate that  MdGSTF6  encodes an important GST transporter of anthocyanins in apple fruit and provide evidence for the associated regulatory mechanisms. Therefore, MdMYB1 can not only regulate anthocyanin synthesis, but also control the transport of anthocyanin in apples. This information may be useful for further clarifying the regulation of anthocyanin transport in apple.

MdGSTF6, activated by MdMYB1, plays an essential role in anthocyanin accumulation in apple