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The frmRAB operon codes for: a regulator, FrmR (EcFrmR); a formaldehyde dehydrogenase, FrmA; and an S-formylglutathione hydrolase, FrmB. In E. coli, and many other organisms, formaldehyde in the cytosol reacts with the major reductant glutathione, yielding S-hydroxymethylglutathione, which is then oxidized by FrmA to S-formylglutathione. In E. coli S-formylglutathione is hydrolyzed to formate and glutathione by the product of the final gene of the frmRAB operon, FrmB, or by a second, less-efficient hydrolase YeiG11; some bacteria, such as the closely related Salmonella enterica serovar Typhimurium, lack frmB but possess yeiG. The proteins coded by the frmRAB operon and yeiG can permit bacteria to sense and detoxify formaldehyde.