Acts as a sulfur carrier required for 2-thiolation of mcm(5S(2U at tRNA wobble positions of cytosolic tRNA(Lys, tRNA(Glu and tRNA(Gln. Serves as sulfur donor in tRNA 2-thiolation reaction by being thiocarboxylated (-COSH at its C-terminus by MOCS3. The sulfur is then transferred to tRNA to form 2-thiolation of mcm(5S(2U. Also acts as a ubiquitin-like protein (UBL that is covalently conjugated via an isopeptide bond to lysine residues of target proteins such as MOCS3, ATPBD3, CTU2, USP15 and CAS. The thiocarboxylated form serves as substrate for conjugation and oxidative stress specifically induces the formation of UBL-protein conjugates.