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coenzyme redox methanogens 7-mercaptoheptanoylthreoninephosphate thiol methane methanogenesis enzyme methyl reductase cofactor f430 prosthetic group electron-donor heterodisulfide mcr methanogenic archaea anaerobic microorganism strain methanobacterium thermoautotrophicum spectroscopic x-ray crystallography nmr spectroscopy porphyrin macrocyclic corphin tetrapyrrole carbocyclic nickel chlorophyll siroheme heme biosynthesis metabolite metabolism mechanism pathway biosynthetic substrate precursor metabolic enzyme endogenous molecule molecular formula skeletal 3d