Trans-translation is a quality-control process, activated upon premature te
rmination of protein elongation, which recycles stalled ribosomes and degra
des incomplete polypeptides. These functions are facilitated by transfer-me
ssenger RNA (tmRNA, also called 10Sa RNA or SsrA RNA), a small stable RNA m
olecule encoded by the SsrA gene found in bacteria, chloroplasts and mitoch
ondria. Most tmRNAs consist of a tRNA- and an mRNA-like domain connected by
up to four pseudoknots. Comparative sequence analysis provided the first i
nsight into tmRNA secondary and three-dimensional structure. Studies of the
E. coli tmRNA in vitro and in vivo demonstrated that tmRNA functions as a
ribonucleoprotein (RNP) complex with elongation factor Tu (EF-Tu), protein
SmpB and ribosomal protein S1. The tRNA-like and mRNA-like activities of tm
RNA mark prematurely terminated proteins for degradation by attaching to th
eir C-termini peptide tags, which are recognized by numerous proteases. Stu
dies aimed at understanding the details of the molecular mechanisms of tran
s-translation are ongoing.