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Dipartimento di Chimica Biologica and Istituto di Neuroscienze del CNR, Università di Padova and Venetian Institute for Molecular Medicine (VIMM), Padova, Italy
1Correspondence: Dipartimento di Chimica Biologica, Università di Padova, Viale G. Colombo 3, 35121 Padova, Italy. E-mail: lorenzo.pinna{at}unipd.it
CK2 (formerly termed "casein kinase 2") is a ubiquitous, highly pleiotropic and constitutively active Ser/Thr protein kinase whose implication in neoplasia, cell survival, and virus infection is supported by an increasing number of arguments. Here an updated inventory of 307 CK2 protein substrates is presented. More than one-third of these are implicated in gene expression and protein synthesis as being either transcriptional factors (60)
or effectors of DNA/RNA structure (50)
or translational elements. Also numerous are signaling proteins and proteins of viral origin or essential to virus life cycle. In comparison, only a minority of CK2 targets (a dozen or so) are classical metabolic enzymes. An analysis of 308 sites phosphorylated by CK2 highlights the paramount relevance of negatively charged side chains that are (by far) predominant over any other residues at positions n+3 (the most crucial one), n+1, and n+2. Based on this signature, it is predictable that proteins phosphorylated by CK2 are much more numerous than those identified to date, and it is possible that CK2 alone contributes to the generation of the eukaryotic phosphoproteome more so than any other individual protein kinase. The possibility that CK2 phosphosites play some global role, e.g., by destabilizing 
helices, counteracting caspase cleavage, and generating adhesive motifs, will be discussed.—Meggio, F., Pinna, L. A. One-thousand-and-one substrates of protein kinase CK2?
Key Words: CK2 • casein kinase 2 • phosphoacceptor sites • phosphoproteome
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M. Shimada, C. Namikawa-Yamada, M. Nakanishi, and H. Murakami Regulation of Cdc2p and Cdc13p Is Required for Cell Cycle Arrest Induced by Defective RNA Splicing in Fission Yeast J. Biol. Chem., September 23, 2005; 280(38): 32640 - 32648. [Abstract] [Full Text] [PDF]
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N. Watanabe, H. Arai, J.-i. Iwasaki, M. Shiina, K. Ogata, T. Hunter, and H. Osada Cyclin-dependent kinase (CDK) phosphorylation destabilizes somatic Wee1 via multiple pathways PNAS, August 16, 2005; 102(33): 11663 - 11668. [Abstract] [Full Text] [PDF]
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J. Modrof, K. Lymperopoulos, and P. Roy Phosphorylation of Bluetongue Virus Nonstructural Protein 2 Is Essential for Formation of Viral Inclusion Bodies J. Virol., August 1, 2005; 79(15): 10023 - 10031. [Abstract] [Full Text] [PDF]
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L. Gu, W. J. Esselman, and R. W. Henry Cooperation between Small Nuclear RNA-activating Protein Complex (SNAPC) and TATA-box-binding Protein Antagonizes Protein Kinase CK2 Inhibition of DNA Binding by SNAPC J. Biol. Chem., July 29, 2005; 280(30): 27697 - 27704. [Abstract] [Full Text] [PDF]
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 L. A. Usakin, G. L. Kogan, A. I. Kalmykova, and V. A. Gvozdev An Alien Promoter Capture as a Primary Step of the Evolution of Testes-Expressed Repeats in the Drosophila melanogaster Genome Mol. Biol. Evol., July 1, 2005; 22(7): 1555 - 1560. [Abstract] [Full Text] [PDF]
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A. Dong, Z. Liu, Y. Zhu, F. Yu, Z. Li, K. Cao, and W.-H. Shen Interacting Proteins and Differences in Nuclear Transport Reveal Specific Functions for the NAP1 Family Proteins in Plants Plant Physiology, July 1, 2005; 138(3): 1446 - 1456. [Abstract] [Full Text] [PDF]
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M. P. Lolkema, M. L. Gervais, C. M. Snijckers, R. P. Hill, R. H. Giles, E. E. Voest, and M. Ohh Tumor Suppression by the von Hippel-Lindau Protein Requires Phosphorylation of the Acidic Domain J. Biol. Chem., June 10, 2005; 280(23): 22205 - 22211. [Abstract] [Full Text] [PDF]
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M. Yamada, S. Katsuma, T. Adachi, A. Hirasawa, S. Shiojima, T. Kadowaki, Y. Okuno, T.-a. Koshimizu, S. Fujii, Y. Sekiya, et al. Inhibition of protein kinase CK2 prevents the progression of glomerulonephritis PNAS, May 24, 2005; 102(21): 7736 - 7741. [Abstract] [Full Text] [PDF]
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K. Yamane and T. J. Kinsella CK2 Inhibits Apoptosis and Changes Its Cellular Localization Following Ionizing Radiation Cancer Res., May 15, 2005; 65(10): 4362 - 4367. [Abstract] [Full Text] [PDF]
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D. A. Canton, M. E. K. Olsten, K. Kim, A. Doherty-Kirby, G. Lajoie, J. A. Cooper, and D. W. Litchfield The Pleckstrin Homology Domain-Containing Protein CKIP-1 Is Involved in Regulation of Cell Morphology and the Actin Cytoskeleton and Interaction with Actin Capping Protein Mol. Cell. Biol., May 1, 2005; 25(9): 3519 - 3534. [Abstract] [Full Text] [PDF]
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 M. L. DE MARCHIS, A. GIORGI, M. E. SCHININA, I. BOZZONI, and A. FATICA Rrp15p, a novel component of pre-ribosomal particles required for 60S ribosome subunit maturation RNA, April 1, 2005; 11(4): 495 - 502. [Abstract] [Full Text] [PDF]
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F. Rodriguez, C. C. Allende, and J. E. Allende Protein kinase casein kinase 2 holoenzyme produced ectopically in human cells can be exported to the external side of the cellular membrane PNAS, March 29, 2005; 102(13): 4718 - 4723. [Abstract] [Full Text] [PDF]
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 K. Yamane and T. J. Kinsella Casein Kinase 2 Regulates Both Apoptosis and the Cell Cycle Following DNA Damage Induced by 6-Thioguanine Clin. Cancer Res., March 15, 2005; 11(6): 2355 - 2363. [Abstract] [Full Text] [PDF]
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N. Franck, J. Le Seyec, C. Guguen-Guillouzo, and L. Erdtmann Hepatitis C Virus NS2 Protein Is Phosphorylated by the Protein Kinase CK2 and Targeted for Degradation to the Proteasome J. Virol., March 1, 2005; 79(5): 2700 - 2708. [Abstract] [Full Text] [PDF]
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 A. V. Ljubimov, S. Caballero, A. M. Aoki, L. A. Pinna, M. B. Grant, and R. Castellon Involvement of Protein Kinase CK2 in Angiogenesis and Retinal Neovascularization Invest. Ophthalmol. Vis. Sci., December 1, 2004; 45(12): 4583 - 4591. [Abstract] [Full Text] [PDF]
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A. C. B. Lim, Z. Hou, C.-P. Goh, and R. Z. Qi Protein Kinase CK2 Is an Inhibitor of the Neuronal Cdk5 Kinase J. Biol. Chem., November 5, 2004; 279(45): 46668 - 46673. [Abstract] [Full Text] [PDF]
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P. Malik and J. B. Clements Protein kinase CK2 phosphorylation regulates the interaction of Kaposi's sarcoma-associated herpesvirus regulatory protein ORF57 with its multifunctional partner hnRNP K Nucleic Acids Res., October 14, 2004; 32(18): 5553 - 5569. [Abstract] [Full Text] [PDF]
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H. Luo, D. W. Chan, T. Yang, M. Rodriguez, B. P.-C. Chen, M. Leng, J.-J. Mu, D. Chen, Z. Songyang, Y. Wang, et al. A New XRCC1-Containing Complex and Its Role in Cellular Survival of Methyl Methanesulfonate Treatment Mol. Cell. Biol., October 1, 2004; 24(19): 8356 - 8365. [Abstract] [Full Text] [PDF]
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 D. D. Mruk and C. Y. Cheng Sertoli-Sertoli and Sertoli-Germ Cell Interactions and Their Significance in Germ Cell Movement in the Seminiferous Epithelium during Spermatogenesis Endocr. Rev., October 1, 2004; 25(5): 747 - 806. [Abstract] [Full Text] [PDF]
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M. E. K. Olsten, D. A. Canton, C. Zhang, P. A. Walton, and D. W. Litchfield The Pleckstrin Homology Domain of CK2 Interacting Protein-1 Is Required for Interactions and Recruitment of Protein Kinase CK2 to the Plasma Membrane J. Biol. Chem., October 1, 2004; 279(40): 42114 - 42127. [Abstract] [Full Text] [PDF]
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S. Roosbeek, F. Peelman, A. Verhee, C. Labeur, H. Caster, M. F. Lensink, C. Cirulli, J. Grooten, C. Cochet, J. Vandekerckhove, et al. Phosphorylation by Protein Kinase CK2 Modulates the Activity of the ATP Binding Cassette A1 Transporter J. Biol. Chem., September 3, 2004; 279(36): 37779 - 37788. [Abstract] [Full Text] [PDF]
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S. A. Beausoleil, M. Jedrychowski, D. Schwartz, J. E. Elias, J. Villen, J. Li, M. A. Cohn, L. C. Cantley, and S. P. Gygi Large-scale characterization of HeLa cell nuclear phosphoproteins PNAS, August 17, 2004; 101(33): 12130 - 12135. [Abstract] [Full Text] [PDF]
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G. L. Russo, M. Tosto, A. Mupo, I. Castellano, A. Cuomo, and E. Tosti Biochemical and Functional Characterization of Protein Kinase CK2 in Ascidian Ciona intestinalis Oocytes at Fertilization: CLONING AND SEQUENCE ANALYSIS OF cDNA FOR {alpha} AND {beta} SUBUNITS J. Biol. Chem., July 30, 2004; 279(31): 33012 - 33023. [Abstract] [Full Text] [PDF]
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