In melanoma cell lines, a short-chain dehydrogenase/reductase family member, 3-hydroxybutyrate dehydrogenase 2 (BDH2) at MLCs, was found to generate 2,5-dihydroxybenzoic acid (2,5-DHBA), which shuttles iron from lysosomes to mitochondria to support oxidative phosphorylation (OXPHOS) and ATP production, which, in turn, is utilized by lysosomes to maintain a low pH through v-ATPase activity [79]. Here, BDH2 is linked to melanoma.