B23/Nucleophosmin Serine 4 Phosphorylation Mediates Mitotic Functions of Polo-like Kinase 1

Phosphoprotein profiling by Kinetworks™ analysis of M-phase-arrested HeLa cells by nocodazole treatment revealed that a novel mitosis-specific phosphorylation event occurred in the nucleolar protein B23/nucleophosmin at a conserved Ser-4 residue. Consistent with the resemblance of the Ser-4 phosphorylation site the of by a Ser-4 of by a phosphorylation in in phosphorylation of by Ser-4 of by protein of Ser-4 phosphorylation of in HeLa of of of of a of in the the of in the of Ser-4 in a of Phosphoprotein profiling by Kinetworks™ analysis of M-phase-arrested HeLa cells by nocodazole treatment revealed that a novel mitosis-specific phosphorylation event occurred in the nucleolar protein B23/nucleophosmin at a conserved Ser-4 residue. Consistent with the resemblance of the Ser-4 phosphorylation site the of by a Ser-4 of by a phosphorylation in in phosphorylation of by Ser-4 of by protein of Ser-4 phosphorylation of in HeLa of of of of a of in the the of in the of Ser-4 in a of of a of of the the of that a of phosphorylation by protein the of the of the of protein in the of a conserved of that of the the of by the of the in the the in cells in cells in the of Consistent with with of by in in of in a of that in the of cells of a in of of the in the of cells that in a of of the in the B23/nucleophosmin a of of by at a a of protein in phosphorylation in phosphorylation the of the of in the the Ser-4 a novel phosphorylation site by a with a phosphorylation site in the of a Kinetworks™ of phosphorylation event of revealed that the Ser-4 phosphorylation with Ser-4 phosphorylation of in HeLa cells by of at phosphorylation site in that in event in phosphorylation of in with in a with of by of by a the by Ser-4 with by by cells with in the the cells in nocodazole treatment cells with in with by at at by the the of protein a a the with of that with a of at of with a of the of by the Kinetworks™ in of the Kinetworks™ analysis at the of protein HeLa cells with nocodazole the protein by analysis a protein by by of the protein with the by by the of the of the the by the a the of Ser-4 phosphorylation by protein with of by with at of with of in of at by the of the with of of in with by at by by with the of in by the with in the by with by HeLa cells in in at by in in in cells with in at by with in the with cells a a a of phosphorylation in the Kinetworks™ phosphorylation in the phosphorylation of in HeLa cells by cells nocodazole a of phosphorylation phosphorylation of protein by that with at in cells with cells of of the of the a a the of in HeLa cells by cells with of at with the at by a in cells by the with in cells cells that the of with a nocodazole a the HeLa cells the cells by with the of the by with the a the of that a with a of by the of the of HeLa cells by with of by with the at the a with with revealed that B23/nucleophosmin the nucleolar protein HeLa with the by of that B23/nucleophosmin with HeLa of of by in by with a in with the revealed a the in with the of at Ser-4 the with the of that a of by the that the of in nocodazole treatment the in protein by of HeLa with a of cells with of at in the of the protein the a of HeLa with nocodazole with revealed that the of a in protein of cells with of in the of the that a of the of revealed the in Ser-4 of of HeLa with in in phosphorylation of at at the site by the a of in the by with with nocodazole with the by the the with the of the that the of Ser-4 with the of the Ser-4 at at Ser-4 of the phosphorylation site by the a of by Ser-4 with in of HeLa in phosphorylation in by HeLa in nocodazole by with the in phosphorylation that Ser-4 phosphorylation by the of phosphorylation by of the protein by Ser-4 by at a phosphorylation site by with the in cells with a of nocodazole the with the with in the of the in the in of the with that the the the of the Ser-4 with that Ser-4 the phosphorylation site by the of the in the of by the that the the of in cells in cells that the Ser-4 phosphorylation at the in that the Ser-4 phosphorylation site the that Ser-4 at the Ser-4 site with conserved a protein of by the the of the protein the of Ser-4 phosphorylation in Ser-4 Ser-4 HeLa cells with protein in with the protein Ser-4 phosphorylation of treatment with the in of Ser-4 phosphorylation with at protein revealed by with the that a Ser-4 Ser-4 a phosphorylation in that a in by a of in Ser-4 of with cells at Ser-4 in Ser-4 phosphorylation in cells with with nocodazole with the Ser-4 phosphorylation cells with nocodazole the of protein treatment by the phosphorylation of in Ser-4 that the of the Ser-4 with phosphorylation of protein revealed that the that of with a at Ser-4 at phosphorylation by by in the of the a with the of in in with protein HeLa that with a of in with with the that with in a HeLa the of in by with the of in the of of in the HeLa of protein in the in the Ser-4 HeLa cells with the of Ser-4 phosphorylation by cells a of at by with of Ser-4 of phosphorylation with cells of in cells the of Ser-4 phosphorylation the at protein Ser-4 of HeLa cells with in a in Ser-4 of the phosphorylation of protein the of by a cells with in the of protein a of Ser-4 phosphorylation protein HeLa of in by the phosphorylation of Ser-4 of by by the of protein in the of in the in HeLa cells with in phosphorylation a of by the of protein in that of in by that the Ser-4 phosphorylation a of in the of Ser-4 phosphorylation in the cells in by cells with the the of the a the of Ser-4 phosphorylation by of in the of the of protein with the of Ser-4 phosphorylation with cells of HeLa by that the Ser-4 phosphorylation a phosphorylation in the phosphorylation by the of by with the that that with in Ser-4 HeLa cells by at Ser-4 in that Ser-4 of that that Ser-4 that a Ser-4 of Ser-4 of the in by nocodazole Ser-4 in in phosphorylation the of by in a phosphorylation a in that at Ser-4 in of Ser-4 with in in the phosphorylation by by the the of of in HeLa cells the treatment of nocodazole with by in a a phosphorylation of Ser-4 in the of the in HeLa cells at the the of the in treatment by nocodazole with by nocodazole a of a by nocodazole by a of the of cells a of Ser-4 phosphorylation in with that of in phosphorylation with the Ser-4 at of with Ser-4 phosphorylation the treatment that the of of Ser-4 in in the of Ser-4 phosphorylation the with HeLa cells the of by by of a of the a cells of Ser-4 phosphorylation in in in HeLa HeLa cells with at with cells in in the of in of the of in HeLa cells phosphorylation of at by of in of HeLa cells with cells a of cells a of the cells at by that Ser-4 phosphorylation of cells the in of cells the that by in HeLa the in of the at the of that phosphorylation of by a Ser-4 phosphorylation in HeLa that of Ser-4 phosphorylation with in a of phosphorylation by HeLa in that the of Ser-4 phosphorylation that in a of of by of the of of the of in a novel phosphorylation event at Ser-4 of a protein with a of phosphorylation event revealed that the Ser-4 of Ser-4 the the of Ser-4 phosphorylation in the of in a of the in a the the phosphorylation in the protein that the protein with the a novel protein with protein the nucleolar protein by a that the of in the protein the of by Ser-4 the by the in the in of phosphorylation the in nocodazole treatment in the in the of in at the site by the that the of phosphorylation at that site by the by the of that the the by the the in of in by Ser-4 of that Ser-4 of with the the the of the the of Ser-4 the of in in the novel phosphorylation the of a phosphorylation in a at at phosphorylation by in with Consistent with the phosphorylation Ser-4 the of in by in phosphorylation of in with by the of of a by protein the Ser-4 phosphorylation the in nocodazole treatment the that the of the that the the the in in a of of that in a that of the the of Ser-4 phosphorylation that that the phosphorylation of by the that in a of of in with the in the in that a Ser-4 of at Ser-4 in of the the treatment of nocodazole that phosphorylation of Ser-4 by in in of the in with that of cells Kinetworks™ analysis revealed that the of Ser-4 that that Ser-4 the that at a protein in protein the the protein in phosphorylation of by at of by that in that the of in Consistent with the that of Ser-4 phosphorylation by in HeLa the in the of of a event of the that the Ser-4 phosphorylation with that of that that Ser-4 phosphorylation a in the of by of Ser-4 of with at the in of in in by a of that phosphorylation of Ser-4 the of in a of cells at the that the of by in with the of in the of the a the cells of the of the of that of in the of of of the of cells by the a of in by of with the of at the phosphorylation of Ser-4 in of the of analysis that in of Ser-4 the of the by Ser-4 the that HeLa cells in that cells a of in the of a a of of the of phosphorylation of in a in the of the the of the of Ser-4 phosphorylation by in of the the of in the phosphorylation of Ser-4 a a of a of of the the of that a of phosphorylation by protein the of the of the of protein in the of a conserved of that of the the of by the of the in the the in cells in cells in the of Consistent with with of by in in of in a of that in the of cells of a in of of the in the of cells that in a of of the in the B23/nucleophosmin a of of by at a a of protein in phosphorylation in phosphorylation the of the of in the the Ser-4 a novel phosphorylation site by a with a phosphorylation site in the of a Kinetworks™ of phosphorylation event of revealed that the Ser-4 phosphorylation with Ser-4 phosphorylation of in HeLa cells by of at phosphorylation site in that in event in phosphorylation of in with in a with of by of by a the by Ser-4 with by by cells with in the the cells in nocodazole treatment cells with in with by at at by the the of protein a a the with of that with a of at of with a of the of by the Kinetworks™ in of the Kinetworks™ analysis at the of protein HeLa cells with nocodazole the protein by analysis a protein by by of the protein with the by by the of the of the the by the a the of Ser-4 phosphorylation by protein with of by with at of with of in of at by the of the with of of in with by at by by with the of in by the with in the by with by HeLa cells in in at by in in in cells with in at by with in the with cells a a in with in a with of by of by a the by Ser-4 with by by cells with in the the cells in nocodazole treatment cells with in with by at at by the the of protein a a the with of that with a of at of with a of the of by the Kinetworks™ in of the Kinetworks™ analysis at the of protein HeLa cells with nocodazole the protein by analysis a protein by by of the protein with the by by the of the of the the by the a the of Ser-4 phosphorylation by protein with of by with at of with of in of at by the of the with of of in with by at by by with the of in by the with in the by with by HeLa cells in in at by in in in cells with in at by with in the with cells a a a of phosphorylation in the Kinetworks™ phosphorylation in the phosphorylation of in HeLa cells by cells nocodazole a of phosphorylation phosphorylation of protein by that with at in cells with cells of of the of the a a the of in HeLa cells by cells with of at with the at by a in cells by the with in cells cells that the of with a nocodazole the a the of that a with a of by the of the of HeLa cells by with of by with the at the a with with revealed that B23/nucleophosmin the nucleolar protein HeLa with the by of that B23/nucleophosmin with HeLa of of by in by with a in with the revealed a the in with the of at Ser-4 the with the of that a of by the that the of in nocodazole treatment the in protein by of HeLa with a of cells with of at in the of the protein the a of HeLa with nocodazole with revealed that the of a in protein of cells with of in the of the that a of the of revealed the in Ser-4 of of HeLa with in in phosphorylation of at at the site by the a of in the by with with nocodazole with the by the the with the of the that the of Ser-4 with the of the Ser-4 at at Ser-4 of the phosphorylation site by the a of by Ser-4 with in of HeLa in phosphorylation in by HeLa in nocodazole by with the in phosphorylation that Ser-4 phosphorylation by the of phosphorylation by of the protein by Ser-4 by at a phosphorylation site by with the in cells with a of nocodazole the with the with in the of the in the in of the with that the the the of the Ser-4 with that Ser-4 the phosphorylation site by the of the in the of by the that the the of in cells in cells that the Ser-4 phosphorylation at the in that the Ser-4 phosphorylation site the that Ser-4 at the Ser-4 site with conserved a protein of by the the of the protein the of Ser-4 phosphorylation in Ser-4 Ser-4 HeLa cells with protein in with the protein Ser-4 phosphorylation of treatment with the in of Ser-4 phosphorylation with at protein revealed by with the that a Ser-4 Ser-4 a phosphorylation in that a in by a of in Ser-4 of with cells at Ser-4 in Ser-4 phosphorylation in cells with with nocodazole with the Ser-4 phosphorylation cells with nocodazole the of protein treatment by the phosphorylation of in Ser-4 that the of the Ser-4 with phosphorylation of protein revealed that the that of with a at Ser-4 at phosphorylation by by in the of the a with the of in in with protein HeLa that with a of in with with the that with in a HeLa the of in by with the of in the of of in the HeLa of protein in the in the Ser-4 HeLa cells with the of Ser-4 phosphorylation by cells a of at by with of Ser-4 of phosphorylation with cells of in cells the of Ser-4 phosphorylation the at protein Ser-4 of HeLa cells with in a in Ser-4 of the phosphorylation of protein the of by a cells with in the of protein a of Ser-4 phosphorylation protein HeLa of in by the phosphorylation of Ser-4 of by by the of protein in the of in the in HeLa cells with in phosphorylation a of by the of protein in that of in by that the Ser-4 phosphorylation a of in the of Ser-4 phosphorylation in the cells in by cells with the the of the a the of Ser-4 phosphorylation by of in the of the of protein with the of Ser-4 phosphorylation with cells of HeLa by that the Ser-4 phosphorylation a phosphorylation in the phosphorylation by the of by with the that that with in Ser-4 HeLa cells by at Ser-4 in that Ser-4 of that that Ser-4 that a Ser-4 of Ser-4 of the in by nocodazole Ser-4 in in phosphorylation the of by in a phosphorylation a in that at Ser-4 in of Ser-4 with in in the phosphorylation by by the the of of in HeLa cells the treatment of nocodazole with by in a a phosphorylation of Ser-4 in the of the in HeLa cells at the the of the in treatment by nocodazole with by nocodazole a of a by nocodazole by a of the of cells a of Ser-4 phosphorylation in with that of in phosphorylation with the Ser-4 at of with Ser-4 phosphorylation the treatment that the of of Ser-4 in in the of Ser-4 phosphorylation the with HeLa cells the of by by of a of the a cells of Ser-4 phosphorylation in in in HeLa HeLa cells with at with cells in in the of in of the of in HeLa cells phosphorylation of at by of in of HeLa cells with cells a of cells a of the cells at by that Ser-4 phosphorylation of cells the in of cells the that by in HeLa the in of the at the of that phosphorylation of by a Ser-4 phosphorylation in HeLa that of Ser-4 phosphorylation with in a of phosphorylation by HeLa in that the of Ser-4 phosphorylation a of phosphorylation in the Kinetworks™ phosphorylation in the phosphorylation of in HeLa cells by cells nocodazole a of phosphorylation phosphorylation of protein by that with at in cells with cells of of the of the a a the of in HeLa cells by cells with of at with the at by a in cells by the with in cells cells that the of with a nocodazole the a the of that a with a of by the of the of HeLa cells by with of by with the at the a with with revealed that B23/nucleophosmin of by in by with a in with the revealed a the in with the of at Ser-4 the with the of that a of by the that the of in nocodazole treatment the in protein by of HeLa with a of cells with of at in the of the protein the a the by the the with the of the that the of Ser-4 with the of the Ser-4 at at Ser-4 of the phosphorylation site by the a of by Ser-4 with in of HeLa in phosphorylation in by HeLa in nocodazole by with the in phosphorylation that Ser-4 phosphorylation by the of phosphorylation by of the protein by Ser-4 by at a phosphorylation site by with the in cells with a of nocodazole the with the with in the of the in the in of the with that the the the of the Ser-4 with that Ser-4 the phosphorylation site by the of the in the of by the that the the of in cells in cells that the Ser-4 phosphorylation at the in that the Ser-4 phosphorylation site the that Ser-4 at the Ser-4 site with conserved a protein of by the the of the protein the of Ser-4 phosphorylation in Ser-4 Ser-4 HeLa cells with protein in with the protein Ser-4 phosphorylation of treatment with the in of Ser-4 phosphorylation with at protein revealed by with the that a Ser-4 Ser-4 a phosphorylation in that a in by a of in Ser-4 of with cells at Ser-4 in the Ser-4 with phosphorylation of protein revealed that the that of with a at Ser-4 at phosphorylation by by in the of the a with the of in in with protein HeLa that with a of in with with the that with in a the Ser-4 HeLa cells with the of Ser-4 phosphorylation by cells a of at by with of Ser-4 of phosphorylation with cells of in cells the of Ser-4 phosphorylation the at protein that the Ser-4 phosphorylation a of in the of Ser-4 phosphorylation in the cells in by cells with the the of the a the of Ser-4 phosphorylation by of in the of the of protein with the of Ser-4 phosphorylation with cells of HeLa by that the Ser-4 phosphorylation a phosphorylation in the phosphorylation by the of by with the that that with in Ser-4 HeLa cells by at Ser-4 in that Ser-4 of that that Ser-4 that a Ser-4 of Ser-4 of the in by nocodazole Ser-4 in in phosphorylation the of by in a phosphorylation a in that at Ser-4 in the of the in treatment by nocodazole with by nocodazole a of a by nocodazole by a of the of cells a of Ser-4 phosphorylation in with that of in phosphorylation with the Ser-4 at of with Ser-4 phosphorylation the treatment that the of of Ser-4 in in the of Ser-4 phosphorylation the with HeLa cells the of by by of a of the a cells phosphorylation of at by of in of HeLa cells with cells a of cells a of the cells at by that Ser-4 phosphorylation of cells the in of cells the that by in HeLa the in of the at the of that phosphorylation of by a Ser-4 phosphorylation in HeLa that of Ser-4 phosphorylation with in a of phosphorylation by HeLa in that the of Ser-4 phosphorylation that in a of of by of the of of the of in a novel phosphorylation event at Ser-4 of a protein with a of phosphorylation event revealed that the Ser-4 of Ser-4 the the of Ser-4 phosphorylation in the of in a of the in a the the phosphorylation in the protein that the protein with the a novel protein with protein the nucleolar protein by a that the of in the protein the of by Ser-4 the by the in the in of phosphorylation the in nocodazole treatment in the in the of in at the site by the that the of phosphorylation at that site by the by the of that the the by the the in of in by Ser-4 of that Ser-4 of with the the the of the the of Ser-4 the of in in the novel phosphorylation the of a phosphorylation in a at at phosphorylation by in with Consistent with the phosphorylation Ser-4 the of in by in phosphorylation of in with by the of of a by protein the Ser-4 phosphorylation the in nocodazole treatment the that the of the that the the the in in a of of that in a that of the the of Ser-4 phosphorylation that that the phosphorylation of by the that in a of of in with the in the in that a Ser-4 of at Ser-4 in of the the treatment of nocodazole that phosphorylation of Ser-4 by in in of the in with that of cells Kinetworks™ analysis revealed that the of Ser-4 that that Ser-4 the that at a protein in protein the the protein in phosphorylation of by at of by that in that the of in Consistent with the that of Ser-4 phosphorylation by in HeLa the in the of of a event of the that the Ser-4 phosphorylation with that of that that Ser-4 phosphorylation a in the of by of Ser-4 of with at the in of in in by a of that phosphorylation of Ser-4 the of in a of cells at the that the of by in with the of in the of the a the cells of the of the of that of in the of of of the of cells by the a of in by of with the of at the phosphorylation of Ser-4 in of the of analysis that in of Ser-4 the of the by Ser-4 the that HeLa cells in that cells a of in the of a a of of the of phosphorylation of in a in the of the the of the of Ser-4 phosphorylation by in of the the of in the phosphorylation of Ser-4 a a that in a of of by of the of of the of in a novel phosphorylation event at Ser-4 of a protein with a of phosphorylation event revealed that the Ser-4 of Ser-4 the the of Ser-4 phosphorylation in the of in a of the in a the the phosphorylation in the protein that the protein with the a novel protein with protein the nucleolar protein by a that the of in the protein the of by Ser-4 the by the in the in of phosphorylation the in nocodazole treatment in the in the of in at the site by the that the of phosphorylation at that site by the by the of that the the by the the in of in by Ser-4 of that Ser-4 of with the the the of the the of Ser-4 the of in in the novel phosphorylation the of a phosphorylation in a at at phosphorylation by in with Consistent with the phosphorylation Ser-4 the of in by in phosphorylation of in with by the of of a by protein the Ser-4 phosphorylation the in nocodazole treatment the that the of the that the the the in in a of of that in a that of the the of Ser-4 phosphorylation that that the phosphorylation of by the that in a of of in with the in the in that a Ser-4 of at Ser-4 in of the the treatment of nocodazole that phosphorylation of Ser-4 by in in of the in with that of cells Kinetworks™ analysis revealed that the of Ser-4 that that Ser-4 the that at a protein in protein the the protein in phosphorylation of by at of by that in that the of in Consistent with the that of Ser-4 phosphorylation by in HeLa the in the of of a event of the that the Ser-4 phosphorylation with that of that that Ser-4 phosphorylation a in the of by of Ser-4 of Consistent with at the in of in in by a of that phosphorylation of Ser-4 the of in a of cells at the that the of by in with the of in the of the a the cells of the of the of that of in the of of of the of cells by the a of in by of with the of at the phosphorylation of Ser-4 in of the of analysis that in of Ser-4 the of the by Ser-4 the that HeLa cells in that cells a of in the of a a of of the of phosphorylation of in a in the of the the of the of Ser-4 phosphorylation by in of the the of in the phosphorylation of Ser-4 a a of the

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