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Molecular and Cellular Biology, December 2003, p. 8718-8728, Vol. 23, No. 23
0270-7306/03/$08.00+0     DOI: 10.1128/MCB.23.23.8718-8728.2003
Copyright © 2003, American Society for Microbiology. All Rights Reserved.

The Cdk-Activating Kinase Cak1p Promotes Meiotic S Phase through Ime2p{dagger}

Karen Schindler,1 Kirsten R. Benjamin,2,{ddagger} Allison Martin,1 Andrew Boglioli,1 Ira Herskowitz,2 and Edward Winter1*

Department of Biochemistry and Molecular Pharmacology, Thomas Jefferson University, Philadelphia, Pennsylvania 19107,1 Department of Biochemistry and Biophysics, University of California, San Francisco, California 941432

Received 7 July 2003/ Returned for modification 19 August 2003/ Accepted 3 September 2003

CAK1 encodes an essential protein kinase in Saccharomyces cerevisiae that is required for activation of the Cdc28p Cdk. CAK1 also has several CDC28-independent functions that are unique to meiosis. The earliest of these functions is to induce S phase, which is regulated differently in meiosis than in mitosis. In mitosis, Cdc28p controls its own S-phase-promoting activity by signaling the destruction of its inhibitor, Sic1p. In meiosis, Sic1p destruction is signaled by the meiosis-specific Ime2p protein kinase. Our data show that Cak1p is required to activate Ime2p through a mechanism that requires threonine 242 and tyrosine 244 in Ime2p's activation loop. This activation promotes autophosphorylation and accumulation of multiply phosphorylated forms of Ime2p during meiotic development. Consistent with Cak1p's role in activating Ime2p, cells lacking Cak1p are deficient in degrading Sic1p. Deletion of SIC1 or overexpression of IME2 can partially suppress the S-phase defect in cak1 mutant cells, suggesting that Ime2p is a key target of Cak1p regulation. These data show that Cak1p is required for the destruction of Sic1p in meiosis, as in mitosis, but in meiosis, it functions through a sporulation-specific kinase.

* Corresponding author. Mailing address: Department of Biochemistry and Molecular Pharmacology, Thomas Jefferson University, Philadelphia, PA 19107. Phone: (215) 503-4139. Fax: (215) 923-9162. E-mail: winter{at}lac.jci.tju.edu.

{dagger} This paper is dedicated to the memory of Ira Herskowitz (deceased 28 April 2003).

{ddagger} Present address: Molecular Sciences Institute, Berkeley, CA 94704.

Molecular and Cellular Biology, December 2003, p. 8718-8728, Vol. 23, No. 23
0022-538X/03/$08.00+0     DOI: 10.1128/MCB.23.23.8718-8728.2003
Copyright © 2003, American Society for Microbiology. All Rights Reserved.



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