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Vol. 14, No. 18, pp. 2298-2313, September 15, 2000
1 Department of Biochemistry and Molecular Biology,
2 Department of Molecular Physiology and Biophysics,
3 Department of Molecular and Cellular Biology, Baylor
College of Medicine, Houston, Texas 77030, USA; 4 Department
of Biochemistry and Molecular Genetics, 5 Department of
Pathology, University of Alabama, Birmingham, Alabama 35294, USA;
6 CRC Centre for Cancer Therapeutics, Institute of Cancer
Research, Sutton, SM2 5NG, United Kingdom; 7 Fox Chase Cancer
Center, Philadelphia, Pennsylvania 19111, USA
Cyclin E/Cdk2 acts at the G1/S-phase transition to promote the E2F
transcriptional program and the initiation of DNA synthesis. To explore
further how cyclin E/Cdk2 controls S-phase events, we examined the
subcellular localization of the cyclin E/Cdk2 interacting protein
p220NPAT and its regulation by phosphorylation. p220 is
localized to discrete nuclear foci. Diploid fibroblasts in Go and G1
contain two p220 foci, whereas S- and G2-phase cells contain primarily
four p220 foci. Cells in metaphase and telophase have no detectable
focus. p220 foci contain cyclin E and are coincident with Cajal bodies (CBs), subnuclear organelles that associate with histone gene clusters
on chromosomes 1 and 6. Interestingly, p220 foci associate with
chromosome 6 throughout the cell cycle and with chromosome 1 during S
phase. Five cyclin E/Cdk2 phosphorylation sites in p220 were
identified. Phospho-specific antibodies against two of these sites
react with p220 within CBs in a cell cycle-specific manner. The timing
of p220 phosphorylation correlates with the appearance of cyclin E in
CBs at the G1/S boundary, and this phosphorylation is maintained until
prophase. Expression of p220 activates transcription of the histone H2B
promoter. Importantly, mutation of Cdk2 phosphorylation sites to
alanine abrogates the ability of p220 to activate the histone H2B
promoter. Collectively, these results strongly suggest that
p220NPAT links cyclical cyclin E/Cdk2 kinase activity to
replication-dependent histone gene transcription.
[Key Words: Cyclin-dependent kinases; phosphorylation; Cajal (coiled) bodies; histone transcription]
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