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In  Salmonella enterica  serovar Typhimurium five of the eight family codon boxes are decoded by a tRNA having the modified nucleoside uridine-5-oxyacetic acid (cmo 5 U) as a wobble nucleoside present in position 34 of the tRNA. In the proline family codon box, one (tRNA Pro  cmo5UGG ) of the three tRNAs that reads the four proline codons has cmo 5 U34. According to theoretical predictions and several results obtained in vitro, cmo 5 U34 should base pair with A, G, and U in the third position of the codon but not with C. To analyze the function of cmo 5 U34 in tRNA Pro  cmo5UGG  in vivo, we first identified two genes ( cmoA  and  cmoB ) involved in the synthesis of cmo 5 U34. The null mutation  cmoB2  results in tRNA having 5-hydroxyuridine (ho 5 U34) instead of cmo 5 U34, whereas the null mutation  cmoA1  results in the accumulation of 5-methoxyuridine (mo 5 U34) and ho 5 U34 in tRNA. The results suggest that the synthesis of cmo 5 U34 occurs as follows: U34 → ?  ho 5 U → CmoB  mo 5 U → CmoA?  cmo 5 U. We introduced the  cmoA1  or the  cmoB2  null mutations into a strain that only had tRNA Pro  cmo5UGG  and thus lacked the other two proline-specific tRNAs normally present in the cell. From analysis of growth rates of various strains and of the frequency of +1 frameshifting at a CCC-U site we conclude: (1) unexpectedly, tRNA Pro  cmo5UGG  is able to read all four proline codons; (2) the presence of ho 5 U34 instead of cmo 5 U34 in this tRNA reduces the efficiency with which it reads all four codons; and (3) the fully modified nucleoside is especially important for reading proline codons ending with U or C.

The modified wobble nucleoside uridine-5-oxyacetic acid in tRNAProcmo5UGG promotes reading of all four proline codons in vivo