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Molecular and Cellular Biology, December 2004, p. 10857-10867, Vol. 24, No. 24
0270-7306/04/$08.00+0     DOI: 10.1128/MCB.24.24.10857-10867.2004
Copyright © 2004, American Society for Microbiology. All Rights Reserved.

Counting of Rif1p and Rif2p on Saccharomyces cerevisiae Telomeres Regulates Telomere Length{dagger}

Daniel L. Levy and Elizabeth H. Blackburn*

Department of Biochemistry and Biophysics, University of California, San Francisco, San Francisco, California

Received 10 August 2004/ Returned for modification 30 August 2004/ Accepted 16 September 2004

Telomere length is negatively regulated by proteins of the telomeric DNA-protein complex. Rap1p in Saccharomyces cerevisiae binds the telomeric TG1-3 repeat DNA, and the Rap1p C terminus interacts with Rif1p and Rif2p. We investigated how these three proteins negatively regulate telomere length. We show that direct tethering of each Rif protein to a telomere shortens that telomere proportionally to the number of tethered molecules, similar to previously reported counting of Rap1p. Surprisingly, Rif proteins could also regulate telomere length even when the Rap1p C terminus was absent, and tethered Rap1p counting was completely dependent on the Rif proteins. Thus, Rap1p counting is in fact Rif protein counting. In genetic settings that cause telomeres to be abnormally long, tethering even a single Rif2p molecule was sufficient for maximal effectiveness in preventing the telomere overelongation. We show that a heterologous protein oligomerization domain, the mammalian PDZ domain, when fused to Rap1p can confer telomere length control. We propose that a nucleation and spreading mechanism is involved in forming the higher-order telomere structure that regulates telomere length.

* Corresponding author. Mailing address: Department of Biochemistry and Biophysics, University of California, San Francisco, San Francisco, CA 94143-2200. Phone: (415) 476-4912. Fax: (415) 514-2913. E-mail: telomer{at}itsa.ucsf.edu.

{dagger} Supplemental material for this article may be found at http://mcb.asm.org.

Molecular and Cellular Biology, December 2004, p. 10857-10867, Vol. 24, No. 24
0022-538X/04/$08.00+0     DOI: 10.1128/MCB.24.24.10857-10867.2004
Copyright © 2004, American Society for Microbiology. All Rights Reserved.



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