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On the other hand, the very low pK a values of the nucleophilic Cys in reduced DsbA [pK a =3.5(82)], DsbC [pK a =4.1(99)], or Grx1 [pK a =3.5 (38, 69)] do not correspond to proportionally high nucleophilicity and are likely to have additional kinetic ( e.g., the effect of the leaving group pK a , see below) or thermodynamic ( i.e., an effect on the position of the corresponding thiol–disulfide equilibrium) purpose in the overall multistep thiol–disulfide exchange processes ( i.e., reduction of: DsbA by DsbB, protein disulfides by DsbC, and glutathionylated proteins by Grx1, respectively)