中心体周期相关蛋白磷酸化/去磷酸化及其功能的研究进展
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摘要
中心体(centrosome)是动物细胞和低等植物细胞中存在的一种重要的无膜细胞器。正常情况下,它参与细胞的有丝分裂,形成基于微管结构的纺锤体,牵引着复制后的染色体移向细胞的两极。这一过程保证了遗传物质DNA能够精确、有序和完整地遗传到子代的细胞中。与细胞有丝分裂及增殖过程中发挥作用的其他重要细胞器一样,它的行为和调控同样是由一系列与功能相关的蛋白质来完成的。在这些蛋白质中,一部分与中心体的复制、分裂相关,保证中心体增殖周期的实现;另一部分则在细胞周期中与中心体有着一定的联系,但同时在细胞生命活动中发挥着其他重要功能。无论是在正常细胞中,还是在异常细胞(如肿瘤细胞)中,蛋白质修饰的调节方式都占据了优势,其中尤以磷酸化修饰方式引人注目。中心体是细胞内重要的细胞器之一,大量与中心体周期相关的蛋白质同样也通过磷酸化修饰方式进行调节。主要对中心体周期相关蛋白的磷酸化修饰及功能进行综述,为中心体周期及其调节的深入研究提供参考。
Centrosome is a significant organelle without membrane in animal cells and some lower plant cells.Normally,centrosomes participate cell division(mitosis),form spindle which is based on microtubules and then pull the duplicated chromosomes to the opposite poles of the cell.This process ensures that genetic materials(DNA) can pass from one generation to the next accurately,orderly and perfectly.Like other essential organelles related with cell division and proliferation,the behavior and regulation of centrosome also depends on many function-related proteins.In these proteins,one part connects with centrosome duplication and separation,ensuring centrosome cycle achieved;the other part,not only associates with cell cycle and centrosome,but also plays vital roles in other aspects.Regardless of normal cells or abnormal cells(e.g.tumour cells),protein modifications,especially the phosphorylation modification,dominate in cell regulation.As one kind of the major organelles cells,centrosome contains many centrosomal proteins depend on phosphorylation regulation.To provide references for future research,the centrosome cycle related protein phosphorylation and function were reviewed.
Centrosome is a significant organelle without membrane in animal cells and some lower plant cells.Normally,centrosomes participate cell division(mitosis),form spindle which is based on microtubules and then pull the duplicated chromosomes to the opposite poles of the cell.This process ensures that genetic materials(DNA) can pass from one generation to the next accurately,orderly and perfectly.Like other essential organelles related with cell division and proliferation,the behavior and regulation of centrosome also depends on many function-related proteins.In these proteins,one part connects with centrosome duplication and separation,ensuring centrosome cycle achieved;the other part,not only associates with cell cycle and centrosome,but also plays vital roles in other aspects.Regardless of normal cells or abnormal cells(e.g.tumour cells),protein modifications,especially the phosphorylation modification,dominate in cell regulation.As one kind of the major organelles cells,centrosome contains many centrosomal proteins depend on phosphorylation regulation.To provide references for future research,the centrosome cycle related protein phosphorylation and function were reviewed.
引文
[1]Doxsey S.Re-evaluating centrosome function.Nature ReviewsMolecular Cell Biology,2001,2(9):688-698.
[2]Bornens M.Centrosome composition and microtubule anchoringmechanisms.Current Opinion in Cell Biology,2002,14(1):25-34.
[3]Lüders J,Stearns T.Microtubule-organizing centres:a re-evaluation.Nature Reviews Molecular Cell Biology,2007,8(2):161-167.
[4]Bettencourt-Dias M,Glover D M.Centrosome biogenesis andfunction:centrosomics brings new understanding.Nature ReviewsMolecular Cell Biology,2007,8(6):451-463.
[5]Yamashita Y M,Mahowald A P,Perlin J R,et al.Asymmetricinheritance of mother versus daughter centrosome in stem celldivision.Science,2007,315:518-521.
[6]Gard D L,Hafezi S,Zhang T,et al.Centrosome duplicationcontinues in cycloheximide-treated xenopus blastulae in theabsence of a detectable cell cycle.Journal of Cell Biology,1990,110(6):2033-2042.
[7]Barros T P,Kinoshita K,Hyman A A,et al.Aurora a activatesD-TACC-Msps complexes exclusively at centrosomes to stabilizecentrosomal microtubules.Journal of Cell Biology,2005,170(7):1039-1046.
[8]Wei Y,Shen E Z,Zhao N,et al.Identification of a novelcentrosomal protein CrpF46 involved in cell cycle progression andmitosis.Experimental Cell Research,2008,314:1693-1707.
[9]Fukasawa K.Oncogenes and tumour suppressors take oncentrosomes.Nature Reviews Cancer,2007,7(12):911-924.
[10]Uetake Y,Sluder G.Cell cycle progression after cleavage failure:mammalian somatic cells do not possess a“tetraploidycheckpoint”.Journal of Cell Biology,2004,165(5):609-615.
[11]Meraldi P,Erich A N.Centrosome cohesion is regulated by abalance of kinase and phosphatase activities.Journal of CellBiology,2001,114(20):3749-3757.
[12]Lorca T,Bernis C,Vigneron S,et al.Constant regulation of boththe MPF amplification loop and the Greatwall-PP2A pathway isrequired for metaphase II arrest and correct entry into the firstembryonic cell cycle.Journal of Cell Biology,2010,123(13):2281-2291.
[13]Kitagawa D,Fluckiger I,Polanowska J,et al.PP2Aphosphatase acts upon SAS-5 to ensure centriole formation in C.elegans Embryos.Developmental Cell,2011,20(4):550-562.
[14]Song M H,Liu Y,Anderson D E,et al.Protein phosphatase 2A-SUR-6/B55 regulates centriole duplication in C.elegans bycontrolling the levels of centriole assembly factors.DevelopmentalCell,2011,20(4):563-571.
[15]Domingo-Sananes M R,Kapuy O,Hunt T,et al.Switches andlatches:a biochemical tug-of-war between the kinases andphosphatases that control mitosis.Philosophical Transactions ofthe Royal Society B-biological Sciences,2011,366(1584):3584-3594.
[16]Meraldi P,Nigg E A.The centrosome cycle.Febs Letters,2002,512(1-3):9-13.
[17]Tokuyama Y,Horn H F,Kawamura K,et al.Specificphosphorylation of nucleophosmin on Thr(199)by cyclin-dependent kinase 2-cyclin E and its role in centrosomeduplication.Journal of Biological Chemistry,2001,276(24):21529-21537.
[18]Ma Z,Kanai M,Kawamura K,et al.Interaction between ROCKII and nucleophosmin/B23 in the regulation of centrosomeduplication.Molecular and Cellular Biology,2006,26(23):9016-9034.
[19]Wang H F,Takenaka K,Nakanishi A,et al.BRCA2 andnucleophosmin coregulate centrosome amplification and form acomplex with the Rho effector kinase ROCK2.Cancer Research,2010,71(1):68-77.
[20]Chen Z H,Indjeian V B,McManus M,et al.CP110,a cellcycle-dependent CDK substrate,regulates centrosome duplicationin human cells.Developmental Cell,2002,3(3):339-350.
[21]D’Angiolella V,Donato V,Vijayakumar S,et al.SCF(CyclinF)controls centrosome homeostasis and mitotic fidelity throughCP110 degradation.Nature,2010,466(7302):138-U161.
[22]Kasbek C,Yang C H,Yusof A M,et al.Preventing thedegradation of Mps1 at centrosomes is sufficient to causecentrosome reduplication in human cells.Molecular Biology of theCell,2007,18(11):4457-4469.
[23]Mattison C P,Old W M,Steiner E,et al.Mps1 activation loopautophosphorylation enhances kinase activity.Journal ofBiological Chemistry,2007,282(42):30553-30561.
[24]Alvarado-Kristensson M,Rodriguez M J,Silio V,et al.SADBphosphorylation ofγ-tubulin regulates centrosome duplication.Nature Cell Biology,2009,11(9):1081-1086.
[25]Hergovich A,Kohler R S,Schmitz D,et al.The MST1 andhMOB1 tumor suppressors control human centrosome duplicationby regulating NDR kinase phosphorylation.Current Biology,2009,19(20):1692-1702.
[26]Cizmecioglu O,Warnke S,Arnold M,et al.Plk2 regulatedcentriole duplication is dependent on its localization to thecentrioles and a functional polo-box domain.Cell Cycle,2008,7(22):3548-3555.
[27]Chang J,Cizmecioglu O,Hoffmann I,et al.PLK2phosphorylation is critical for CPAP function in procentrioleformation during the centrosome cycle.Embo Journal,2010,29(14):2395-2406.
[28]Kitagawa D,Busso C,Fluckiger I,et al.Phosphorylation of SAS-6 by ZYG-1 is critical for centriole formation in C.elegansembryos.Developmental Cell,2009,17(6):900-907.
[29]Bahtz R,Seidler J,Arnold M,et al.GCP6 is a substrate of Plk4and required for centriole duplication.Journal of Cell Science,2012,125(2):486-496.
[30]Verollet C,Colombie N,Daubon T,et al.Drosophilamelanogasterγ-TuRC is dispensable for targetingγ-tubulin to thecentrosome and microtubule nucleation.Journal of Cell Biology,2006,12(4):517-528.
[31]Haren L,Stearns T,Luders J.Plk1-dependent recruitment ofgamma-tubulin complexes to mitotic centrosomes involves multiplePCM components.Plos One,2009,4(6):e5976.
[32]Zhang X Y,Chen Q,Feng J,et al.Sequential phosphorylation ofNedd1 by Cdk1 and Plk1 is required for targeting of theγTuRC tothe centrosome.Journal of Cell Science,2009,122(13):2240-2251.
[33]Lee K,Rhee K.PLK1 phosphorylation of pericentrin initiatescentrosome maturation at the onset of mitosis.Journal of CellBiology,2011,195(7):1093-1101.
[34]Guarguaglini G,Duncan P I,Stierhof Y D,et al.The forkhead-associated domain protein Cep170 interacts with Polo-like kinase1 and serves as a marker for mature centrioles.Molecular Biologyof the Cell,2005,16(3):1095-1107.
[35]Hannak E,Kirkham M,Hyman A A,et al.Aurora-A kinase isrequired for centrosome maturation in caenorhabditis elegans.Journal of Cell Biology,2001,155(7):1109-1115.
[36]Berdnik D,Knoblich J A.Drosophila aurora-A is required forcentrosome maturation and actin-dependent asymmetric proteinlocalization during mitosis.Current Biology,2002,12(8):640-647.
[37]Maton G,Thibier C,Castro A,et al.Cdc2-cyclin B triggers H3kinase activation of Aurora-A in Xenopus oocytes.Journal ofBiological Chemistry,2003,278(24):21439-21449.
[38]Pascreau G,Delcros J G,Morin N,et al.Aurora-A kinaseSer349 phosphorylation is required during Xenopus laevis oocytematuration.Developmental Biology,2008,317(2):523-530.
[39]Kinoshita K,Noetzel T L,Pelletier L,et al.Aurora Aphosphorylation of TACC3/maskin is required for centrosome-dependent microtubule assembly in mitosis.Journal of CellBiology,2005,170(7):1047-1055.
[40]Mori D,Yano Y,Toyo-Oka K,et al.NDEL1 phosphorylation byaurora-A kinase is essential for centrosomal maturation,separation,and TACC3 recruitment.Molecular and CellularBiology,2007,27(1):352-367.
[41]Wu C C,Yang T Y,Yu C T,et al.p53 negatively regulatesAurora A via both transcriptional and posttranslational regulation.Cell Cycle.2012,11(18):3433-3442.
[42]Rosenblatt J,Cramer L P,Baum B,et al.Myosin II-dependentcortical movement is required for centrosome separation andpositioning during mitotic spindle assembly.Cell,2004,117(3):361-372.
[43]Cao J,Crest J,Fasulo B,et al.Cortical Actin dynamics facilitateearly-stage centrosome separation.Current Biology,2010,20(8):770-776.
[44]Wang W Q,Chen L Y,Ding Y B,et al.Centrosome separationdriven by actin-microfilaments during mitosis is mediated bycentrosome-associated tyrosine-phosphorylated cortactin.Journalof Cell Science,2008,121(8):1334-1343.
[45]Tehrani S,Tomasevic N,Weed S,et al.Src phosphorylation ofcortactin enhances actin assembly.Proceedings of the NationalAcademy of Sciences of the United States of America,2007,104(29):11933-11938.
[46]Tanenbaum M E,Macurek L,Galjart N,et al.Dynein,Lis1 andCLIP-170 counteract Eg5-dependent centrosome separation duringbipolar spindle assembly.Embo Journal,2008,27(24):3235-3245.
[47]Smith E,Hegarat N,Vesely C,et al.Differential control of Eg5-dependent centrosome EMBO separation by Plk1 and Cdk1.Embo Journal,2011,30(11):2233-2245.
[48]Mardin B R,Lange C,Baxter J E,et al.Components of theHippo pathway cooperate with Nek2 kinase to regulate centrosomedisjunction.Nature Cell Biology,2010,12(12):1166-U106.
[49]Busch C,Barton O,Morgenstern E,et al.The G(2)/Mcheckpoint phosphatase cdc25C is located within centrosomes.International Journal of Biochemistry&Cell Biology,2007,39(9):1707-1713.
[50]Boutros R,Lorenzo C,Mondesert O,et al.CDC25B associateswith a centrin 2-containing complex and is involved inmaintaining centrosome integrity.Biology of the Cell,2011,103(2):55-68.
[51]Shreeram S,Hee W K,Bulavin D V.Cdc25A serine 123phosphorylation couples centrosome duplication with DNAreplication and regulates tumorigenesis.Molecular and CellularBiology,2008,28(24):7442-7450.
[52]Boutros R,Ducommun B.Asymmetric localization of the CDC25Bphosphatase to the mother centrosome during interphase.CellCycle,2008,7(3):401-406.
[53]Martin-Granados C,Philp A,Oxenham S K,et al.Depletion ofprotein phosphatase 4 in human cells reveals essential roles incentrosome maturation,cell migration and the regulation of RhoGTPases.International Journal of Biochemistry&Cell Biology,2008,40(10):2315-2332.
[54]Mardin B R,Agircan F G,Lange C,et al.Plk1 controls theNek2A-PP1 gamma antagonism in centrosome disjunction.Current Biology,2011,21(13):1145-1151.
[55]Eto M,Elliott E,Prickett T D,et al.Inhibitor-2 regulates proteinphosphatase-1 complexed with NimA-related kinase to inducecentrosome separation.Journal of Biological Chemistry,2002,277(46):44013-44020.
[56]Leach C,Shenolikar S,Brautigan D L.Phosphorylation ofphosphatase inhibitor-2 at centrosomes during mitosis.Journal ofBiological Chemistry,2003,278(28):26015-26020.
[57]Janssens V,Longin S,Goris J.PP2A holoenzyme assembly:incauda venenum(the sting is in the tail).Trends in BiochemicalSciences,2008,33(3):113-121.
[58]Mochida S,Ikeo S,Gannon J,et al.Regulated activity of PP2A-B55 delta is crucial for controlling entry into and exit from mitosisin Xenopus egg extracts.Embo Journal,2009,28(18):2777-2785.
[59]Mochida S,Maslen S L,Skehel M,et al.Greatwallphosphorylates an inhibitor of protein phosphatase 2A that isessential for mitosis.Science,2010,330(6011):1670-1673.
[60]Gharbi-Ayachi A,Labbe J C,Burgess A,et al.The substrate ofgreatwall kinase,Arpp19,controls mitosis by inhibiting proteinphosphatase 2A.Science,2010,330(6011):1673-1677.
[61]Brownlee C W,Klebba J E,Buster D W,et al.The proteinphosphatase 2A regulatory subunit twins stabilizes Plk4 to inducecentriole amplification.Journal of Cell Biology,2011,195(2):231-243.
[62]Voets E,Wolthuis R M F.MASTL is the human orthologue ofgreatwall kinase that facilitates mitotic entry,anaphase andcytokinesis.Cell Cycle,2010,9(17):3591-3601.
[63]Boutros R,Lobjois V,Ducommun B.CDC25B involvement in thecentrosome duplication cycle and in microtubule nucleation.Cancer Research,2007,67:11557-11564.
[64]Sumiyoshi E,Sugimoto A,Yamamoto M.Protein phosphatase 4 isrequired for centrosome maturation in mitosis and sperm meiosisin C.elegans.Journal of Cell Science,2002,115(7):1403-1410.
[65]Lingle W L,Lutz W H,Ingle J N,et al.Centrosome hypertrophyin human breast tumors:implications for genomic stability andcell polarity.Proc Natl Acad Sci.USA,1998,95:2950-2955.
[66]Weber R G,Bridger J M,Benner A,et al.Centrosomeamplification as a possible mechanism for numerical chromosomeaberrations in cerebral primitive neuroectodermal tumors withTP53 mutations.Cytogenet Cell Genet,1998,83:266-269.
[67]Sato N,Mizumoto K,Nakamura M,et al.Correlation betweencentrosome abnormalities and chromosomal instability in humanpancreatic cancer cells.Cancer Genet Cytogenet,2001,126:13-19.
[68]Pihan G A,Purohit A,Wallace J,et al.Centrosome defects canaccount for cellular and genetic changes that characterize prostatecancer progression.Cancer Res,2001,61:2212-2219.
[69]Takayama K,Horie-Inoue K,Suzuki T,et al.TACC2 is anandrogen-responsive cell cycle regulator promoting androgen-mediated and castration-resistant growth of prostate cancer.MolEndocrinol,2012,26(5):748-761.
[70]Wang S,Li W,Liu N,et al.Nek2A contributes to tumorigenicgrowth and possibly functions as potential therapeutic target forhuman breast cancer.J Cell Biochem,2012,113(6):1904-1914.
[71]Shen E Z,Lei Y,Liu Q,et al.Identification and characterizationof INMAP,a novel interphase nucleus and mitotic apparatusprotein that is involved in spindle formation and cell cycleprogression.Exp Cell Res,2009,315:1100-1116.
[72]Rangasamy V,Mishra R,Sondarva G,et al.Mixed-lineagekinase 3 phosphorylates prolyl-isomerase pinl to regulate itsnuclear translocation and cellular function.Proc Natl Acad SciUSA,2012,109(21):8149-8154.
[2]Bornens M.Centrosome composition and microtubule anchoringmechanisms.Current Opinion in Cell Biology,2002,14(1):25-34.
[3]Lüders J,Stearns T.Microtubule-organizing centres:a re-evaluation.Nature Reviews Molecular Cell Biology,2007,8(2):161-167.
[4]Bettencourt-Dias M,Glover D M.Centrosome biogenesis andfunction:centrosomics brings new understanding.Nature ReviewsMolecular Cell Biology,2007,8(6):451-463.
[5]Yamashita Y M,Mahowald A P,Perlin J R,et al.Asymmetricinheritance of mother versus daughter centrosome in stem celldivision.Science,2007,315:518-521.
[6]Gard D L,Hafezi S,Zhang T,et al.Centrosome duplicationcontinues in cycloheximide-treated xenopus blastulae in theabsence of a detectable cell cycle.Journal of Cell Biology,1990,110(6):2033-2042.
[7]Barros T P,Kinoshita K,Hyman A A,et al.Aurora a activatesD-TACC-Msps complexes exclusively at centrosomes to stabilizecentrosomal microtubules.Journal of Cell Biology,2005,170(7):1039-1046.
[8]Wei Y,Shen E Z,Zhao N,et al.Identification of a novelcentrosomal protein CrpF46 involved in cell cycle progression andmitosis.Experimental Cell Research,2008,314:1693-1707.
[9]Fukasawa K.Oncogenes and tumour suppressors take oncentrosomes.Nature Reviews Cancer,2007,7(12):911-924.
[10]Uetake Y,Sluder G.Cell cycle progression after cleavage failure:mammalian somatic cells do not possess a“tetraploidycheckpoint”.Journal of Cell Biology,2004,165(5):609-615.
[11]Meraldi P,Erich A N.Centrosome cohesion is regulated by abalance of kinase and phosphatase activities.Journal of CellBiology,2001,114(20):3749-3757.
[12]Lorca T,Bernis C,Vigneron S,et al.Constant regulation of boththe MPF amplification loop and the Greatwall-PP2A pathway isrequired for metaphase II arrest and correct entry into the firstembryonic cell cycle.Journal of Cell Biology,2010,123(13):2281-2291.
[13]Kitagawa D,Fluckiger I,Polanowska J,et al.PP2Aphosphatase acts upon SAS-5 to ensure centriole formation in C.elegans Embryos.Developmental Cell,2011,20(4):550-562.
[14]Song M H,Liu Y,Anderson D E,et al.Protein phosphatase 2A-SUR-6/B55 regulates centriole duplication in C.elegans bycontrolling the levels of centriole assembly factors.DevelopmentalCell,2011,20(4):563-571.
[15]Domingo-Sananes M R,Kapuy O,Hunt T,et al.Switches andlatches:a biochemical tug-of-war between the kinases andphosphatases that control mitosis.Philosophical Transactions ofthe Royal Society B-biological Sciences,2011,366(1584):3584-3594.
[16]Meraldi P,Nigg E A.The centrosome cycle.Febs Letters,2002,512(1-3):9-13.
[17]Tokuyama Y,Horn H F,Kawamura K,et al.Specificphosphorylation of nucleophosmin on Thr(199)by cyclin-dependent kinase 2-cyclin E and its role in centrosomeduplication.Journal of Biological Chemistry,2001,276(24):21529-21537.
[18]Ma Z,Kanai M,Kawamura K,et al.Interaction between ROCKII and nucleophosmin/B23 in the regulation of centrosomeduplication.Molecular and Cellular Biology,2006,26(23):9016-9034.
[19]Wang H F,Takenaka K,Nakanishi A,et al.BRCA2 andnucleophosmin coregulate centrosome amplification and form acomplex with the Rho effector kinase ROCK2.Cancer Research,2010,71(1):68-77.
[20]Chen Z H,Indjeian V B,McManus M,et al.CP110,a cellcycle-dependent CDK substrate,regulates centrosome duplicationin human cells.Developmental Cell,2002,3(3):339-350.
[21]D’Angiolella V,Donato V,Vijayakumar S,et al.SCF(CyclinF)controls centrosome homeostasis and mitotic fidelity throughCP110 degradation.Nature,2010,466(7302):138-U161.
[22]Kasbek C,Yang C H,Yusof A M,et al.Preventing thedegradation of Mps1 at centrosomes is sufficient to causecentrosome reduplication in human cells.Molecular Biology of theCell,2007,18(11):4457-4469.
[23]Mattison C P,Old W M,Steiner E,et al.Mps1 activation loopautophosphorylation enhances kinase activity.Journal ofBiological Chemistry,2007,282(42):30553-30561.
[24]Alvarado-Kristensson M,Rodriguez M J,Silio V,et al.SADBphosphorylation ofγ-tubulin regulates centrosome duplication.Nature Cell Biology,2009,11(9):1081-1086.
[25]Hergovich A,Kohler R S,Schmitz D,et al.The MST1 andhMOB1 tumor suppressors control human centrosome duplicationby regulating NDR kinase phosphorylation.Current Biology,2009,19(20):1692-1702.
[26]Cizmecioglu O,Warnke S,Arnold M,et al.Plk2 regulatedcentriole duplication is dependent on its localization to thecentrioles and a functional polo-box domain.Cell Cycle,2008,7(22):3548-3555.
[27]Chang J,Cizmecioglu O,Hoffmann I,et al.PLK2phosphorylation is critical for CPAP function in procentrioleformation during the centrosome cycle.Embo Journal,2010,29(14):2395-2406.
[28]Kitagawa D,Busso C,Fluckiger I,et al.Phosphorylation of SAS-6 by ZYG-1 is critical for centriole formation in C.elegansembryos.Developmental Cell,2009,17(6):900-907.
[29]Bahtz R,Seidler J,Arnold M,et al.GCP6 is a substrate of Plk4and required for centriole duplication.Journal of Cell Science,2012,125(2):486-496.
[30]Verollet C,Colombie N,Daubon T,et al.Drosophilamelanogasterγ-TuRC is dispensable for targetingγ-tubulin to thecentrosome and microtubule nucleation.Journal of Cell Biology,2006,12(4):517-528.
[31]Haren L,Stearns T,Luders J.Plk1-dependent recruitment ofgamma-tubulin complexes to mitotic centrosomes involves multiplePCM components.Plos One,2009,4(6):e5976.
[32]Zhang X Y,Chen Q,Feng J,et al.Sequential phosphorylation ofNedd1 by Cdk1 and Plk1 is required for targeting of theγTuRC tothe centrosome.Journal of Cell Science,2009,122(13):2240-2251.
[33]Lee K,Rhee K.PLK1 phosphorylation of pericentrin initiatescentrosome maturation at the onset of mitosis.Journal of CellBiology,2011,195(7):1093-1101.
[34]Guarguaglini G,Duncan P I,Stierhof Y D,et al.The forkhead-associated domain protein Cep170 interacts with Polo-like kinase1 and serves as a marker for mature centrioles.Molecular Biologyof the Cell,2005,16(3):1095-1107.
[35]Hannak E,Kirkham M,Hyman A A,et al.Aurora-A kinase isrequired for centrosome maturation in caenorhabditis elegans.Journal of Cell Biology,2001,155(7):1109-1115.
[36]Berdnik D,Knoblich J A.Drosophila aurora-A is required forcentrosome maturation and actin-dependent asymmetric proteinlocalization during mitosis.Current Biology,2002,12(8):640-647.
[37]Maton G,Thibier C,Castro A,et al.Cdc2-cyclin B triggers H3kinase activation of Aurora-A in Xenopus oocytes.Journal ofBiological Chemistry,2003,278(24):21439-21449.
[38]Pascreau G,Delcros J G,Morin N,et al.Aurora-A kinaseSer349 phosphorylation is required during Xenopus laevis oocytematuration.Developmental Biology,2008,317(2):523-530.
[39]Kinoshita K,Noetzel T L,Pelletier L,et al.Aurora Aphosphorylation of TACC3/maskin is required for centrosome-dependent microtubule assembly in mitosis.Journal of CellBiology,2005,170(7):1047-1055.
[40]Mori D,Yano Y,Toyo-Oka K,et al.NDEL1 phosphorylation byaurora-A kinase is essential for centrosomal maturation,separation,and TACC3 recruitment.Molecular and CellularBiology,2007,27(1):352-367.
[41]Wu C C,Yang T Y,Yu C T,et al.p53 negatively regulatesAurora A via both transcriptional and posttranslational regulation.Cell Cycle.2012,11(18):3433-3442.
[42]Rosenblatt J,Cramer L P,Baum B,et al.Myosin II-dependentcortical movement is required for centrosome separation andpositioning during mitotic spindle assembly.Cell,2004,117(3):361-372.
[43]Cao J,Crest J,Fasulo B,et al.Cortical Actin dynamics facilitateearly-stage centrosome separation.Current Biology,2010,20(8):770-776.
[44]Wang W Q,Chen L Y,Ding Y B,et al.Centrosome separationdriven by actin-microfilaments during mitosis is mediated bycentrosome-associated tyrosine-phosphorylated cortactin.Journalof Cell Science,2008,121(8):1334-1343.
[45]Tehrani S,Tomasevic N,Weed S,et al.Src phosphorylation ofcortactin enhances actin assembly.Proceedings of the NationalAcademy of Sciences of the United States of America,2007,104(29):11933-11938.
[46]Tanenbaum M E,Macurek L,Galjart N,et al.Dynein,Lis1 andCLIP-170 counteract Eg5-dependent centrosome separation duringbipolar spindle assembly.Embo Journal,2008,27(24):3235-3245.
[47]Smith E,Hegarat N,Vesely C,et al.Differential control of Eg5-dependent centrosome EMBO separation by Plk1 and Cdk1.Embo Journal,2011,30(11):2233-2245.
[48]Mardin B R,Lange C,Baxter J E,et al.Components of theHippo pathway cooperate with Nek2 kinase to regulate centrosomedisjunction.Nature Cell Biology,2010,12(12):1166-U106.
[49]Busch C,Barton O,Morgenstern E,et al.The G(2)/Mcheckpoint phosphatase cdc25C is located within centrosomes.International Journal of Biochemistry&Cell Biology,2007,39(9):1707-1713.
[50]Boutros R,Lorenzo C,Mondesert O,et al.CDC25B associateswith a centrin 2-containing complex and is involved inmaintaining centrosome integrity.Biology of the Cell,2011,103(2):55-68.
[51]Shreeram S,Hee W K,Bulavin D V.Cdc25A serine 123phosphorylation couples centrosome duplication with DNAreplication and regulates tumorigenesis.Molecular and CellularBiology,2008,28(24):7442-7450.
[52]Boutros R,Ducommun B.Asymmetric localization of the CDC25Bphosphatase to the mother centrosome during interphase.CellCycle,2008,7(3):401-406.
[53]Martin-Granados C,Philp A,Oxenham S K,et al.Depletion ofprotein phosphatase 4 in human cells reveals essential roles incentrosome maturation,cell migration and the regulation of RhoGTPases.International Journal of Biochemistry&Cell Biology,2008,40(10):2315-2332.
[54]Mardin B R,Agircan F G,Lange C,et al.Plk1 controls theNek2A-PP1 gamma antagonism in centrosome disjunction.Current Biology,2011,21(13):1145-1151.
[55]Eto M,Elliott E,Prickett T D,et al.Inhibitor-2 regulates proteinphosphatase-1 complexed with NimA-related kinase to inducecentrosome separation.Journal of Biological Chemistry,2002,277(46):44013-44020.
[56]Leach C,Shenolikar S,Brautigan D L.Phosphorylation ofphosphatase inhibitor-2 at centrosomes during mitosis.Journal ofBiological Chemistry,2003,278(28):26015-26020.
[57]Janssens V,Longin S,Goris J.PP2A holoenzyme assembly:incauda venenum(the sting is in the tail).Trends in BiochemicalSciences,2008,33(3):113-121.
[58]Mochida S,Ikeo S,Gannon J,et al.Regulated activity of PP2A-B55 delta is crucial for controlling entry into and exit from mitosisin Xenopus egg extracts.Embo Journal,2009,28(18):2777-2785.
[59]Mochida S,Maslen S L,Skehel M,et al.Greatwallphosphorylates an inhibitor of protein phosphatase 2A that isessential for mitosis.Science,2010,330(6011):1670-1673.
[60]Gharbi-Ayachi A,Labbe J C,Burgess A,et al.The substrate ofgreatwall kinase,Arpp19,controls mitosis by inhibiting proteinphosphatase 2A.Science,2010,330(6011):1673-1677.
[61]Brownlee C W,Klebba J E,Buster D W,et al.The proteinphosphatase 2A regulatory subunit twins stabilizes Plk4 to inducecentriole amplification.Journal of Cell Biology,2011,195(2):231-243.
[62]Voets E,Wolthuis R M F.MASTL is the human orthologue ofgreatwall kinase that facilitates mitotic entry,anaphase andcytokinesis.Cell Cycle,2010,9(17):3591-3601.
[63]Boutros R,Lobjois V,Ducommun B.CDC25B involvement in thecentrosome duplication cycle and in microtubule nucleation.Cancer Research,2007,67:11557-11564.
[64]Sumiyoshi E,Sugimoto A,Yamamoto M.Protein phosphatase 4 isrequired for centrosome maturation in mitosis and sperm meiosisin C.elegans.Journal of Cell Science,2002,115(7):1403-1410.
[65]Lingle W L,Lutz W H,Ingle J N,et al.Centrosome hypertrophyin human breast tumors:implications for genomic stability andcell polarity.Proc Natl Acad Sci.USA,1998,95:2950-2955.
[66]Weber R G,Bridger J M,Benner A,et al.Centrosomeamplification as a possible mechanism for numerical chromosomeaberrations in cerebral primitive neuroectodermal tumors withTP53 mutations.Cytogenet Cell Genet,1998,83:266-269.
[67]Sato N,Mizumoto K,Nakamura M,et al.Correlation betweencentrosome abnormalities and chromosomal instability in humanpancreatic cancer cells.Cancer Genet Cytogenet,2001,126:13-19.
[68]Pihan G A,Purohit A,Wallace J,et al.Centrosome defects canaccount for cellular and genetic changes that characterize prostatecancer progression.Cancer Res,2001,61:2212-2219.
[69]Takayama K,Horie-Inoue K,Suzuki T,et al.TACC2 is anandrogen-responsive cell cycle regulator promoting androgen-mediated and castration-resistant growth of prostate cancer.MolEndocrinol,2012,26(5):748-761.
[70]Wang S,Li W,Liu N,et al.Nek2A contributes to tumorigenicgrowth and possibly functions as potential therapeutic target forhuman breast cancer.J Cell Biochem,2012,113(6):1904-1914.
[71]Shen E Z,Lei Y,Liu Q,et al.Identification and characterizationof INMAP,a novel interphase nucleus and mitotic apparatusprotein that is involved in spindle formation and cell cycleprogression.Exp Cell Res,2009,315:1100-1116.
[72]Rangasamy V,Mishra R,Sondarva G,et al.Mixed-lineagekinase 3 phosphorylates prolyl-isomerase pinl to regulate itsnuclear translocation and cellular function.Proc Natl Acad SciUSA,2012,109(21):8149-8154.