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Role of N-terminal residues in the ubiquitin-independent degradation of human thymidylate synthase.

Pena MM, Xing YY, Koli S, Berger FG.

Thymidylate synthase (TS) catalyzes the reductive methylation of dUMP to form dTMP, a reaction that is essential for maintenance of nucleotide pools during cell growth. Because the enzyme is indispensable for DNA replication in actively dividing cells, it is an important target for cytotoxic drugs used in cancer chemotherapy. Ligand binding to TS results in stabilization of the enzyme and an increase in its intracellular concentration. Previously, we showed that degradation of the TS polypeptide is carried out by the 26S proteasome in a ubiquitin-independent manner. Such degradation is directed by the disordered N-terminal region of the TS polypeptide, and is abrogated by ligand binding. In the current study, we have verified this conclusion by showing that proteasomal degradation of TS is maintained when all lysine residues, which are typically the sites of ubiquitin ligation, are replaced by arginine. In addition, we have mapped the structural determinants of intracellular TS degradation in more detail, and show that residues at the N-terminal end of the molecule, particularly the penultimate amino acid Pro2, play an important role in governing the half-life of the enzyme. This region is capable on its own of destabilizing an evolutionarily distinct TS molecule that normally lacks this domain, indicating that it functions as a degradation signal. Interestingly, degradation of an intrinsically unstable mutant form of TS, containing a Pro->Leu substitution at residue 303, is directed by C-terminal, rather than N-terminal, sequences. The implications of these findings to the control of thymidylate synthase expression, and to the regulation of protein degradation in general, are discussed.

PMID: 16259621 [PubMed - as supplied by publisher]

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