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glutathione antioxidant radicals peroxides lipid heavy metals reactive oxygen species tripeptide gamma peptide linkage cysteine glutamate peptide glutamate–cysteine ligase glutathione synthetase knockout mice cytosol organelles thiol gsh gssg oxidative stress oxidized reduced reducing equivalent neurotransmitter neuromodulators monochlorobimane thiolquant green realthiol winemaking metabolite metabolism biosynthesis cycle mechanism pathway path catabolism biosynthetic biogenesis substrate precursor synthesis derivative bioprecursor metabolic enzyme endogenous 3d