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J. Biol. Chem., Vol. 281, Issue 48, 36952-36959, December 1, 2006

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The Smc5-Smc6 DNA Repair Complex

BRIDGING OF THE Smc5-Smc6 HEADS BY THE KLEISIN, Nse4, AND NON-KLEISIN SUBUNITS^*

From the Genome Damage and Stability Centre, University of Sussex, Falmer, Brighton BN1 9RQ, United Kingdom

Structural maintenance of chromosomes (SMC) proteins play fundamental roles in many aspects of chromosome organization and dynamics. The SMC complexes form unique structures with long coiled-coil arms folded at a hinge domain, so that the globular N- and C-terminal domains are brought together to form a "head." Within the Smc5-Smc6 complex, we previously identified two subcomplexes containing Smc6-Smc5-Nse2 and Nse1-Nse3-Nse4. A third subcomplex containing Nse5 and -6 has also been identified recently. We present evidence that Nse4 is the kleisin component of the complex, which bridges the heads of Smc5 and -6. The C-terminal part of Nse4 interacts with the head domain of Smc5, and structural predictions for Nse4 proteins suggest similar motifs that are shared within the kleisin family. Specific mutations within a predicted winged helix motif of Nse4 destroy the interaction with Smc5. We propose that Nse4 and its orthologs form the -kleisin subfamily. We further show that Nse3, as well as Nse5 and Nse6, also bridge the heads of Smc5 and -6. The Nse1-Nse3-Nse4 and Nse5-Nse6 subcomplexes bind to the Smc5-Smc6 heads domain at different sites.

Received for publication, August 21, 2006 , and in revised form, September 22, 2006.

^* This work was supported by a Medical Research Council Programme Grant and by the European Community Risc-Rad Integrated Project. The costs of publication of this article were defrayed in part by the payment of page charges. This article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.

¹ Present address: Dept. of Functional Genomics and Proteomics, Masaryk University, Kamenice 5, 625 00 Brno, Czech Republic.

² To whom correspondence should be addressed: Genome Damage and Stability Centre, University of Sussex, Falmer, Brighton BN1 9RQ, United Kingdom. Tel.: 44-1273-678120; Fax: 44-1273-678121; E-mail: a.r.lehmann{at}sussex.ac.uk.

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