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coenzyme f420 8-hydroxy-5-deazaflavin cofactor redox methanogens actinomycetota bacterial flavin methylenetetrahydromethanopterin dehydrogenase mycobacterium smegmatis fmn drosophila melanogaster ostreococcus tauri metabolite metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic intermediate biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous contractor cell biology biochemistry biomolecule bioactive microorganism strain molecule molecular formula skeletal structural structure geometry 3d