Making an effective switch at the kinetochore by phosphorylation and dephosphorylation

Chromosoma

Volumn 122, Issue 3, 2013, Pages 135-158

  Funabiki, Hironori ^a   Wynne, David J ^a  

^a ROCKEFELLER UNIVERSITY   (United States)

Author keywords

Cell cycle; Checkpoint; Chromosome segregation; Feedback; Kinase; Kinetochore; Mitosis; Phosphatase; Spindle

Indexed keywords

AURORA B KINASE; BUB1 RELATED PROTEIN; BUB3 PROTEIN; CELL CYCLE PROTEIN 20; CENTROMERE PROTEIN E; CENTROMERE PROTEIN U; CYCLIN DEPENDENT KINASE 1; KINESIN; MITOGEN ACTIVATED PROTEIN KINASE; NDC80 COMPLEX; PHOSPHOPROTEIN PHOSPHATASE 1; PHOSPHOPROTEIN PHOSPHATASE 2A; POLO LIKE KINASE 1; PROTEIN CDC42; PROTEIN KIF2B; PROTEIN KINASE; PROTEIN KINASE BUB1; PROTEIN MAD2; PROTEIN MCAK; PROTEIN MPS1; SURVIVIN; UNCLASSIFIED DRUG;

BINDING AFFINITY; CATALYSIS; CENTROMERE; ENZYME ACTIVATION; ENZYME ACTIVITY; ENZYME INHIBITION; ENZYME REGULATION; ENZYME STABILITY; ENZYME SUBSTRATE; FEEDBACK SYSTEM; HUMAN; MICROTUBULE; MITOSIS; NONHUMAN; PROTEIN DEPHOSPHORYLATION; PROTEIN EXPRESSION; PROTEIN FUNCTION; PROTEIN LOCALIZATION; PROTEIN PHOSPHORYLATION; PROTEIN PROTEIN INTERACTION; REVIEW; SIGNAL TRANSDUCTION;

ANIMALS; CHROMOSOME SEGREGATION; HUMANS; KINETOCHORES; MICROTUBULES; MITOSIS; PHOSPHORYLATION; PROTEIN BINDING; SACCHAROMYCETALES; SPINDLE APPARATUS;

MAMMALIA; SACCHAROMYCETALES; SCHIZOSACCHAROMYCETACEAE;

EID: 84878587815     PISSN: 00095915     EISSN: 14320886     Source Type: Journal    
DOI: 10.1007/s00412-013-0401-5     Document Type: Review

References (266)

1. Abrieu A, Magnaghi-Jaulin L, Kahana JA, Peter M, Castro A, Vigneron S, Lorca T, Cleveland DW, Labbe JC (2001) Mps1 is a kinetochore-associated kinase essential for the vertebrate mitotic checkpoint. Cell 106:83-93 (Pubitemid 32709960)
2. Ahonen LJ, Kallio MJ, Daum JR, Bolton M, Manke IA, Yaffe MB, Stukenberg PT, Gorbsky GJ (2005) Polo-like kinase 1 creates the tension-sensing 3F3/2 phosphoepitope and modulates the association of spindle-checkpoint proteins at kinetochores. Curr Biol 15:1078-1089 (Pubitemid 40841350)
3. Akiyoshi B, Nelson CR, Ranish JA, Biggins S (2009a) Analysis of Ipl1-mediated phosphorylation of the Ndc80 kinetochore protein in Saccharomyces cerevisiae. Genetics 183:1591-1595
4. Akiyoshi B, Nelson CR, Ranish JA, Biggins S (2009b) Quantitative proteomic analysis of purified yeast kinetochores identifies a PP1 regulatory subunit. Genes Dev 23:2887-2899
5. Akiyoshi B, Sarangapani KK, Powers AF, Nelson CR, Reichow SL, Arellano-Santoyo H, Gonen T, Ranish JA, Asbury CL, Biggins S (2010) Tension directly stabilizes reconstituted kinetochore-microtubule attachments. Nature 468:576-579
6. Alexander J, Lim D, Joughin BA, Hegemann B, Hutchins JR, Ehrenberger T, Ivins F, Sessa F, Hudecz O, Nigg EA, Fry AM, Musacchio A, Stukenberg PT, Mechtler K, Peters JM, Smerdon SJ, Yaffe MB (2011) Spatial exclusivity combined with positive and negative selection of phosphorylation motifs is the basis for context-dependent mitotic signaling. Sci Signal 4:ra42
7. Alushin GM, Ramey VH, Pasqualato S, Ball DA, Grigorieff N, Musacchio A, Nogales E (2010) The Ndc80 kinetochore complex forms oligomeric arrays along microtubules. Nature 467:805-810
8. Andrews PD, Ovechkina Y, Morrice N, Wagenbach M, Duncan K, Wordeman L, Swedlow JR (2004) Aurora B regulates MCAK at the mitotic centromere. Dev Cell 6:253-268 (Pubitemid 38190365)
9. Bentley AM, Normand G, Hoyt J, King RW (2007) Distinct sequence elements of cyclin B1 promote localization to chromatin, centrosomes, and kinetochores during mitosis. Mol Biol Cell 18:4847-4858 (Pubitemid 350246687)
10. Bernard P, Maure JF, Javerzat JP (2001) Fission yeast Bub1 is essential in setting up the meiotic pattern of chromosome segregation. Nat Cell Biol 3:522-526 (Pubitemid 32422091)
11. Biggins S, Murray AW (2001) The budding yeast protein kinase Ipl1/Aurora allows the absence of tension to activate the spindle checkpoint. Genes Dev 15:3118-3129 (Pubitemid 33115676)
12. + in fission yeast mitotic control. Cell 57:1009-1016 (Pubitemid 19168381)
13. Boyarchuk Y, Salic A, Dasso M, Arnaoutov A (2007) Bub1 is essential for assembly of the functional inner centromere. J Cell Biol 176:919-928 (Pubitemid 46506966)
14. Burton JL, Solomon MJ (2007) Mad3p, a pseudosubstrate inhibitor of APCCdc20 in the spindle assembly checkpoint. Genes Dev 21:655-667 (Pubitemid 46440288)
15. Buttrick GJ, Lancaster TC, Meadows JC, Millar JB (2012) Plo1 phosphorylates Dam1 to promote chromosome bi-orientation in fission yeast. J Cell Sci 125:1645-1651
16. Cai S, O'Connell CB, Khodjakov A, Walczak CE (2009) Chromosome congression in the absence of kinetochore fibres. Nat Cell Biol 11:832-838
17. Carmena M, Pinson X, Platani M, Salloum Z, Xu Z, Clark A, Macisaac F, Ogawa H, Eggert U, Glover DM, Archambault V, Earnshaw WC (2012a) The chromosomal passenger complex activates Polo kinase at centromeres. PLoS Biol 10:e1001250
18. Carmena M, Wheelock M, Funabiki H, Earnshaw WC (2012b) The chromosomal passenger complex (CPC): from easy rider to the godfather of mitosis. Nat Rev Mol Cell Biol 13(12):789-803
19. Ceulemans H, Vulsteke V, De Maeyer M, Tatchell K, Stalmans W, Bollen M (2002) Binding of the concave surface of the Sds22 superhelix to the alpha 4/alpha 5/alpha 6-triangle of protein phosphatase-1. J Biol Chem 277:47331-47337 (Pubitemid 36159247)
20. Chan GK, Schaar BT, Yen TJ (1998) Characterization of the kinetochore binding domain of CENP-E reveals interactions with the kinetochore proteins CENP-F and hBUBR1. J Cell Biol 143:49-63 (Pubitemid 28480517)
21. Chan GK, Jablonski SA, Sudakin V, Hittle JC, Yen TJ (1999) Human BUBR1 is a mitotic checkpoint kinase that monitors CENP-E functions at kinetochores and binds the cyclosome/APC. J Cell Biol 146:941-954 (Pubitemid 29430438)
22. Chan YW, Jeyaprakash AA, Nigg EA, Santamaria A (2012) Aurora B controls kinetochore-microtubule attachments by inhibiting Ska complex-KMN network interaction. J Cell Biol 196:563-571
23. Chao WC, Kulkarni K, Zhang Z, Kong EH, Barford D (2012) Structure of the mitotic checkpoint complex. Nature 484:208-213
24. Cheeseman IM, Desai A (2008) Molecular architecture of the kinetochore-microtubule interface. Nat Rev Mol Cell Biol 9:33-46
25. Cheeseman IM, Anderson S, Jwa M, Green EM, Js K, Yates JR, Chan CSM, Drubin DG, Barnes G (2002) Phospho-regulation of kinetochore-microtubule attachments by the Aurora kinase Ipl1p. Cell 111:163-172 (Pubitemid 35292437)
26. Cheeseman IM, Niessen S, Anderson S, Hyndman F, Yates JR 3rd, Oegema K, Desai A (2004) A conserved protein network controls assembly of the outer kinetochore and its ability to sustain tension. Genes Dev 18:2255-2268 (Pubitemid 39209574)
27. Cheeseman IM, Chappie JS, Wilson-Kubalek EM, Desai A (2006) The conserved KMN network constitutes the core microtubule-binding site of the kinetochore. Cell 127:983-997 (Pubitemid 44792248)
28. Chen RH (2002) BubR1 is essential for kinetochore localization of other spindle checkpoint proteins and its phosphorylation requires Mad1. J Cell Biol 158:487-496 (Pubitemid 34851707)
29. Chen RH (2004) Phosphorylation and activation of Bub1 on unattached chromosomes facilitate the spindle checkpoint. EMBO J 23:3113-3121 (Pubitemid 39093605)
30. Chen R-H, Waters JC, Salmon ED, Murray AW (1996) Association of spindle assembly checkpoint component XMAD2 with unattached kinetochores. Science 274:242-246 (Pubitemid 26353200)
31. Chen R-H, Shevchenko A, Mann M, Murray AW (1998) Metaphase arrest induced by an excess of the spindle checkpoint protein Xmad2 is independent of Xmad1. J Cell Biol 143:283-295 (Pubitemid 28487850)
32. Cheng L, Zhang J, Ahmad S, Rozier L, Yu H, Deng H, Mao Y (2011) Aurora B regulates formin mDia3 in achieving metaphase chromosome alignment. Dev Cell 20:342-352
33. Choi E, Choe H, Min J, Choi JY, Kim J, Lee H (2009) BubR1 acetylation at prometaphase is required for modulating APC/C activity and timing of mitosis. EMBO J 28:2077-2089
34. Chung E, Chen RH (2003) Phosphorylation of Cdc20 is required for its inhibition by the spindle checkpoint. Nat Cell Biol 5:748-753 (Pubitemid 36986783)
35. Ciferri C, Pasqualato S, Screpanti E, Varetti G, Santaguida S, Dos Reis G, Maiolica A, Polka J, De Luca JG, De Wulf P, Salek M, Rappsilber J, Moores CA, Salmon ED, Musacchio A (2008) Implications for kinetochore-microtubule attachment from the structure of an engineered Ndc80 complex. Cell 133:427-439 (Pubitemid 351622013)
36. Cimini D, Wan X, Hirel CB, Salmon ED (2006) Aurora kinase promotes turnover of kinetochore microtubules to reduce chromosome segregation errors. Curr Biol 16:1711-1718 (Pubitemid 44691779)
37. Clute P, Pines J (1999) Temporal and spatial control of cyclin B1 destruction in metaphase. Nat Cell Biol 1:82-87 (Pubitemid 129501948)
38. Coffman VC, Wu P, Parthun MR, Wu JQ (2011) CENP-A exceeds microtubule attachment sites in centromere clusters of both budding and fission yeast. J Cell Biol 195:563-572
39. D'Angiolella V, Mari C, Nocera D, Rametti L, Grieco D (2003) The spindle checkpoint requires cyclin-dependent kinase activity. Genes Dev 17:2520-2525 (Pubitemid 37271422)
40. Dawlaty MM, Malureanu L, Jeganathan KB, Kao E, Sustmann C, Tahk S, Shuai K, Grosschedl R, van Deursen JM (2008) Resolution of sister centromeres requires RanBP2-mediated SUMOylation of topoisomerase IIalpha. Cell 133:103-115 (Pubitemid 351442988)
41. De Antoni A, Maffini S, Knapp S, Musacchio A, Santaguida S (2012) A small-molecule inhibitor of Haspin alters the kinetochore functions of Aurora B. J Cell Biol 199:269-284
42. De Wulf P, McAinsh AD, Sorger PK (2003) Hierarchical assembly of the budding yeast kinetochore from multiple subcomplexes. Genes Dev 17:2902-2921 (Pubitemid 37523155)
43. DeLuca JG, Howell BJ, Canman JC, Hickey JM, Fang G, Salmon ED (2003) Nuf2 and Hec1 are required for retention of the checkpoint proteins Mad1 and Mad2 to kinetochores. Curr Biol 13:2103-2109 (Pubitemid 37502528)
44. DeLuca JG, Gall WE, Ciferri C, Cimini D, Musacchio A, Salmon ED (2006) Kinetochore microtubule dynamics and attachment stability are regulated by Hec1. Cell 127:969-982 (Pubitemid 44792254)
45. DeLuca KF, Lens SM, DeLuca JG (2011) Temporal changes in Hec1 phosphorylation control kinetochore-microtubule attachment stability during mitosis. J Cell Sci 124:622-634
46. Ditchfield C, Johnson VL, Tighe A, Ellston R, Haworth C, Johnson T, Mortlock A, Keen N, Taylor SS (2003) Aurora B couples chromosome alignment with anaphase by targeting BubR1, Mad2, and Cenp-E to kinetochores. J Cell Biol 161:267-280 (Pubitemid 36529846)
47. Dohadwala M, Da Cruz e Silva EF, Hall FL, Williams RT, Carbonaro-Hall DA, Nairn AC, Greengard P, Berndt N (1994) Phosphorylation and inactivation of protein phosphatase 1 by cyclin-dependent kinases. Proc Natl Acad Sci U S A 91:6408-6412 (Pubitemid 24200670)
48. Doonan JH, Morris NR (1989) The bimG gene of Aspergillus nidulans, required for completion of anaphase, encodes a homolog of mammalian phophoprotein phosphatase 1. Cell 57:987-996 (Pubitemid 19168379)
49. Dumont S, Salmon ED, Mitchison TJ (2012) Deformations within moving kinetochores reveal different sites of active and passive force generation. Science 337:355-358
50. Dunsch AK, Linnane E, Barr FA, Gruneberg U (2011) The astrin-kinastrin/SKAP complex localizes to microtubule plus ends and facilitates chromosome alignment. J Cell Biol 192:959-968
51. Elia AE, Cantley LC, Yaffe MB (2003) Proteomic screen finds pSer/pThr-binding domain localizing Plk1 to mitotic substrates. Science 299:1228-1231 (Pubitemid 36237463)
52. Elowe S, Hummer S, Uldschmid A, Li X, Nigg EA (2007) Tension-sensitive Plk1 phosphorylation on BubR1 regulates the stability of kinetochore microtubule interactions. Genes Dev 21:2205-2219 (Pubitemid 47360842)
53. Emanuele MJ, Lan W, Jwa M, Miller SA, Chan CSM, Stukenberg PT (2008) Aurora B kinase and protein phosphatase 1 have opposing roles in modulating kinetochore assembly. J Cell Biol 181:241-254 (Pubitemid 351574550)
54. Espeut J, Gaussen A, Bieling P, Morin V, Prieto S, Fesquet D, Surrey T, Abrieu A (2008) Phosphorylation relieves autoinhibition of the kinetochore motor Cenp-E. Mol Cell 29:637-643 (Pubitemid 351384643)
55. Espeut J, Cheerambathur DK, Krenning L, Oegema K, Desai A (2012) Microtubule binding by KNL-1 contributes to spindle checkpoint silencing at the kinetochore. J Cell Biol 196:469-482
56. Fernius J, Hardwick KG (2007) Bub1 kinase targets Sgo1 to ensure efficient chromosome biorientation in budding yeast mitosis. PLoS Genet 3:e213 (Pubitemid 350291589)
57. Firestone AJ, Weinger JS, Maldonado M, Barlan K, Langston LD, O'Donnell M, Gelfand VI, Kapoor TM, Chen JK (2012) Small-molecule inhibitors of the AAA + ATPase motor cytoplasmic dynein. Nature 484:125-129
58. Fisk HA, Winey M (2001) The mouse Mps1p-like kinase regulates centrosome duplication. Cell 106:95-104 (Pubitemid 32709961)
59. Foley EA, Kapoor TM (2012) Microtubule attachment and spindle assembly checkpoint signalling at the kinetochore. Nat Rev Mol Cell Biol 14:25-37
60. Foley EA, Maldonado M, Kapoor TM (2011) Formation of stable attachments between kinetochores and microtubules depends on the B56-PP2A phosphatase. Nat Cell Biol 13:1265-1271
61. Foster SA, Morgan DO (2012) The APC/C subunit Mnd2/Apc15 promotes Cdc20 autoubiquitination and spindle assembly checkpoint inactivation. Mol Cell 47:921-932
62. Francisco L, Wang W, Chan CS (1994) Type 1 protein phosphatase acts in opposition to IpL1 protein kinase in regulating yeast chromosome segregation. Mol Cell Biol 14:4731-4740 (Pubitemid 24196823)
63. Gascoigne KE, Taylor SS (2008) Cancer cells display profound intra- and interline variation following prolonged exposure to antimitotic drugs. Cancer Cell 14:111-122
64. Gascoigne KE, Takeuchi K, Suzuki A, Hori T, Fukagawa T, Cheeseman IM (2011) Induced ectopic kinetochore assembly bypasses the requirement for CENP-A nucleosomes. Cell 145:410-422
65. Gassmann R, Holland AJ, Varma D, Wan X, Civril F, Cleveland DW, Oegema K, Salmon ED, Desai A (2010) Removal of Spindly from microtubule-attached kinetochores controls spindle checkpoint silencing in human cells. Genes Dev 24:957-971
66. Gestaut DR, Graczyk B, Cooper J, Widlund PO, Zelter A, Wordeman L, Asbury CL, Davis TN (2008) Phosphoregulation and depolymerization-driven movement of the Dam1 complex do not require ring formation. Nat Cell Biol 10:407-414
67. Good MC, Zalatan JG, Lim WA (2011) Scaffold proteins: hubs for controlling the flow of cellular information. Science 332:680-686
68. Gorbsky GJ, Ricketts WA (1993) Differential expression of a phosphoepitope at the kinetochores of moving chromosomes. J Cell Biol 122:1311-1321 (Pubitemid 23277300)
69. Goto H, Kiyono T, Tomono Y, Kawajiri A, Urano T, Furukawa K, Nigg EA, Inagaki M (2006) Complex formation of Plk1 and INCENP required for metaphase-anaphase transition. Nat Cell Biol 8:180-187
70. Guimaraes GJ, Dong Y, McEwen BF, DeLuca JG (2008) Kinetochore-microtubule attachment relies on the disordered N-terminal tail domain of Hec1. Curr Biol 18:1778-1784
71. Guo Y, Kim C, Ahmad S, Zhang J, Mao Y (2012) CENP-E-dependent BubR1 autophosphorylation enhances chromosome alignment and the mitotic checkpoint. J Cell Biol 198:205-217
72. Guse A, Carroll CW, Moree B, Fuller CJ, Straight AF (2011) In vitro centromere and kinetochore assembly on defined chromatin templates. Nature 477:354-358
73. Hached K, Xie SZ, Buffin E, Cladiere D, Rachez C, Sacras M, Sorger PK, Wassmann K (2011) Mps1 at kinetochores is essential for female mouse meiosis I. Development 138:2261-2271
74. Hauf S, Cole RW, LaTerra S, Zimmer C, Schnapp G, Walter R, Heckel A, van Meel J, Rieder CL, Peters JM (2003) The small molecule Hesperadin reveals a role for Aurora B in correcting kinetochore-microtubule attachment and in maintaining the spindle assembly checkpoint. J Cell Biol 161:281-294 (Pubitemid 36529847)
75. Hendrickx A, Beullens M, Ceulemans H, Den Abt T, Van Eynde A, Nicolaescu E, Lesage B, Bollen M (2009) Docking motif-guided mapping of the interactome of protein phosphatase-1. Chem Biol 16:365-371
76. Hewitt L, Tighe A, Santaguida S, White AM, Jones CD, Musacchio A, Green S, Taylor SS (2010) Sustained Mps1 activity is required in mitosis to recruit O-Mad2 to the Mad1-C-Mad2 core complex. J Cell Biol 190:25-34
77. Hood EA, Kettenbach AN, Gerber SA, Compton DA (2012) Plk1 regulates the kinesin-13 protein Kif2b to promote faithful chromosome segregation. Mol Biol Cell 23:2264-2274
78. Hori T, Okada M, Maenaka K, Fukagawa T (2008) CENP-O class proteins form a stable complex and are required for proper kinetochore function. Mol Biol Cell 19:843-854 (Pubitemid 351481802)
79. Howell BJ, McEwen BF, Canman JC, Hoffman DB, Farrar EM, Rieder CL, Salmon ED (2001) Cytoplasmic dynein/dynactin drives kinetochore protein transport to the spindle poles and has a role in mitotic spindle checkpoint inactivation. J Cell Biol 155:1159-1172 (Pubitemid 34286267)
80. Howell BJ, Moree B, Farrar EM, Stewart S, Fang G, Salmon ED (2004) Spindle checkpoint protein dynamics at kinetochores in living cells. Curr Biol 14:953-964 (Pubitemid 38722609)
81. Hsu JY, Sun ZW, Li X, Reuben M, Tatchell K, Bishop DK, Grushcow JM, Brame CJ, Caldwell JA, Hunt DF, Lin R, Smith MM, Allis CD (2000) Mitotic phosphorylation of histone H3 is governed by Ipl1/aurora kinase and Glc7/PP1 phosphatase in budding yeast and nematodes. Cell 102:279-291
82. Hua S, Wang Z, Jiang K, Huang Y, Ward T, Zhao L, Dou Z, Yao X (2011) CENP-U cooperates with Hec1 to orchestrate kinetochore-microtubule attachment. J Biol Chem 286:1627-1638
83. Huang H, Hittle J, Zappacosta F, Annan RS, Hershko A, Yen TJ (2008) Phosphorylation sites in BubR1 that regulate kinetochore attachment, tension, and mitotic exit. J Cell Biol 183:667-680
84. Hummer S, Mayer TU (2009) Cdk1 negatively regulates midzone localization of the mitotic kinesin Mklp2 and the chromosomal passenger complex. Curr Biol 19:607-612
85. Indjeian VB, Murray AW (2007) Budding yeast mitotic chromosomes have an intrinsic bias to biorient on the spindle. Curr Biol 17:1837-1846 (Pubitemid 350016924)
86. Ito D, Saito Y, Matsumoto T (2011) Centromere-tethered Mps1 pombe homolog (Mph1) kinase is a sufficient marker for recruitment of the spindle checkpoint protein Bub1, but not Mad1. Proc Natl Acad Sci U S A 109:209-214
87. Jelluma N, Brenkman AB, van den Broek NJ, Cruijsen CW, van Osch MH, Lens SM, Medema RH, Kops GJ (2008) Mps1 phosphorylates Borealin to control Aurora B activity and chromosome alignment. Cell 132:233-246 (Pubitemid 351172430)
88. Jeyaprakash AA, Santamaria A, Jayachandran U, Chan YW, Benda C, Nigg EA, Conti E (2012) Structural and functional organization of the Ska complex, a key component of the kinetochore-microtubule interface. Mol Cell 46:274-286
89. Jia L, Li B, Warrington RT, Hao X, Wang S, Yu H (2011) Defining pathways of spindle checkpoint silencing: functional redundancy between Cdc20 ubiquitination and p31(comet). Mol Biol Cell 22:4227-4235
90. Johnson VL, Scott MI, Holt SV, Hussein D, Taylor SS (2004) Bub1 is required for kinetochore localization of BubR1, Cenp-E, Cenp-F and Mad2, and chromosome congression. J Cell Sci 117:1577-1589 (Pubitemid 38559832)
91. Kallio MJ, McCleland ML, Stukenberg PT, Gorbsky GJ (2002) Inhibition of aurora B kinase blocks chromosome segregation, overrides the spindle checkpoint, and perturbs microtubule dynamics in mitosis. Curr Biol 12:900-905 (Pubitemid 34689224)
92. Kang YH, Park JE, Yu LR, Soung NK, Yun SM, Bang JK, Seong YS, Yu H, Garfield S, Veenstra TD, Lee KS (2006) Self-regulated Plk1 recruitment to kinetochores by the Plk1-PBIP1 interaction is critical for proper chromosome segregation. Mol Cell 24:409-422 (Pubitemid 44754942)
93. Kapoor TM, Lampson MA, Hergert P, Cameron L, Cimini D, Salmon ED, McEwen BF, Khodjakov A (2006) Chromosomes can congress to the metaphase plate before biorientation. Science 311:388-391 (Pubitemid 43132665)
94. Kasuboski JM, Bader JR, Vaughan PS, Tauhata SB, Winding M, Morrissey MA, Joyce MV, Boggess W, Vos L, Chan GK, Hinchcliffe EH, Vaughan KT (2011) Zwint-1 is a novel Aurora B substrate required for the assembly of a dynein-binding platform on kinetochores. Mol Biol Cell 22:3318-3330
95. Kawashima SA, Yamagishi Y, Honda T, Ishiguro K, Watanabe Y (2010) Phosphorylation of H2A by Bub1 prevents chromosomal instability through localizing shugoshin. Science 327:172-177
96. Kelly AE, Funabiki H (2009) Correcting aberrant kinetochore microtubule attachments: an Aurora B-centric view. Curr Opin Cell Biol 21:51-58
97. Kelly AE, Sampath SC, Maniar TA, Woo EM, Chait BT, Funabiki H (2007) Chromosomal enrichment and activation of the aurora B pathway are coupled to spatially regulate spindle assembly. Dev Cell 12:31-43 (Pubitemid 46002596)
98. Kelly AE, Ghenoiu C, Xue JZ, Zierhut C, Kimura H, Funabiki H (2010) Survivin reads phosphorylated histone H3 threonine 3 to activate the mitotic kinase Aurora B. Science 330:235-239
99. Kemmler S, Stach M, Knapp M, Ortiz J, Pfannstiel J, Ruppert T, Lechner J (2009) Mimicking Ndc80 phosphorylation triggers spindle assembly checkpoint signalling. EMBO J 28:1099-1110
100. Kim Y, Holland AJ, Lan W, Cleveland DW (2010) Aurora kinases and protein phosphatase 1 mediate chromosome congression through regulation of CENP-E. Cell 142:444-455
101. King EM, Rachidi N, Morrice N, Hardwick KG, Stark MJ (2007a) Ipl1p-dependent phosphorylation of Mad3p is required for the spindle checkpoint response to lack of tension at kinetochores. Genes Dev 21:1163-1168 (Pubitemid 46789874)
102. King EM, van der Sar SJ, Hardwick KG (2007b) Mad3 KEN boxes mediate both Cdc20 and Mad3 turnover, and are critical for the spindle checkpoint. PLoS One 2:e342
103. Kitajima TS, Sakuno T, Ishiguro K, Iemura S, Natsume T, Kawashima SA, Watanabe Y (2006) Shugoshin collaborates with protein phosphatase 2A to protect cohesin. Nature 441:46-52
104. Kitajima TS, Ohsugi M, Ellenberg J (2011) Complete kinetochore tracking reveals error-prone homologous chromosome biorientation in mammalian oocytes. Cell 146:568-581
105. Kiyomitsu T, Obuse C, Yanagida M (2007) Human Blinkin/AF15q14 is required for chromosome alignment and the mitotic checkpoint through direct interaction with Bub1 and BubR1. Dev Cell 13:663-676 (Pubitemid 350019284)
106. Kiyomitsu T, Murakami H, Yanagida M (2011) Protein interaction domain mapping of human kinetochore protein Blinkin reveals a consensus motif for binding of spindle assembly checkpoint proteins Bub1 and BubR1. Mol Cell Biol 31:998-1011
107. Klebig C, Korinth D, Meraldi P (2009) Bub1 regulates chromosome segregation in a kinetochore-independent manner. J Cell Biol 185:841-858
108. Kline SL, Cheeseman IM, Hori T, Fukagawa T, Desai A (2006) The human Mis12 complex is required for kinetochore assembly and proper chromosome segregation. J Cell Biol 173:9-17
109. Knowlton AL, Lan W, Stukenberg PT (2006) Aurora B is enriched at merotelic attachment sites, where it regulates MCAK. Curr Biol 16:1705-1710 (Pubitemid 44691788)
110. Knowlton AL, Vorozhko VV, Lan W, Gorbsky GJ, Stukenberg PT (2009) ICIS and Aurora B coregulate the microtubule depolymerase Kif2a. Curr Biol 19:758-763
111. Koivomagi M, Valk E, Venta R, Iofik A, Lepiku M, Balog ER, Rubin SM, Morgan DO, Loog M (2011) Cascades of multisite phosphorylation control Sic1 destruction at the onset of S phase. Nature 480:128-131
112. Kops GJ, Shah JV (2012) Connecting up and clearing out: how kinetochore attachment silences the spindle assembly checkpoint. Chromosoma 121:509-525
113. Kulukian A, Han JS, Cleveland DW (2009) Unattached kinetochores catalyze production of an anaphase inhibitor that requires a Mad2 template to prime Cdc20 for BubR1 binding. Dev Cell 16:105-117
114. Labit H, Fujimitsu K, Bayin NS, Takaki T, Gannon J, Yamano H (2012) Dephosphorylation of Cdc20 is required for its C-box-dependent activation of the APC/C. EMBO J 31:3351-3362
115. Lampert F, Hornung P, Westermann S (2010) The Dam1 complex confers microtubule plus end-tracking activity to the Ndc80 kinetochore complex. J Cell Biol 189:641-649
116. Lampson MA, Cheeseman IM (2011) Sensing centromere tension: Aurora B and the regulation of kinetochore function. Trends Cell Biol 21(3):133-140
117. Lampson MA, Renduchitala K, Khodjakov A, Kapoor TM (2004) Correcting improper chromosome-spindle attachments during cell division. Nat Cell Biol 6:232-237 (Pubitemid 38344361)
118. Lan W, Zhang X, Kline-Smith SL, Rosasco SE, Barrett-Wilt GA, Shabanowitz J, Hunt DF, Walczak CE, Stukenberg PT (2004) Aurora B phosphorylates centromeric MCAK and regulates its localization and microtubule depolymerization activity. Curr Biol 14:273-286 (Pubitemid 38300205)
119. Lara-Gonzalez P, Scott MI, Diez M, Sen O, Taylor SS (2011) BubR1 blocks substrate recruitment to the APC/C in a KEN-box-dependent manner. J Cell Sci 124:4332-4345
120. Lawrimore J, Bloom KS, Salmon ED (2011) Point centromeres contain more than a single centromere-specific Cse4 (CENP-A) nucleosome. J Cell Biol 195:573-582
121. Lee S, Thebault P, Freschi L, Beaufils S, Blundell TL, Landry CR, Bolanos-Garcia VM, Elowe S (2012) Characterization of spindle checkpoint kinase Mps1 reveals domain with functional and structural similarities to tetratricopeptide repeat motifs of Bub1 and BubR1 checkpoint kinases. J Biol Chem 287:5988-6001
122. Lenart P, Petronczki M, Steegmaier M, Di Fiore B, Lipp JJ, Hoffmann M, Rettig WJ, Kraut N, Peters JM (2007) The small-molecule inhibitor BI 2536 reveals novel insights into mitotic roles of polo-like kinase 1. Curr Biol 17:304-315
123. Lens SM, Rodriguez JA, Vader G, Span SW, Giaccone G, Medema RH (2006) Uncoupling the central spindle-associated function of the chromosomal passenger complex from its role at centromeres. Mol Biol Cell 17:1897-1909 (Pubitemid 44011605)
124. Li Y, Benezra R (1996) Identification of a human mitotic checkpoint gene: hsMAD2. Science 274:246-248 (Pubitemid 26353201)
125. Li H, Liu XS, Yang X, Wang Y, Wang Y, Turner JR, Liu X (2010) Phosphorylation of CLIP-170 by Plk1 and CK2 promotes timely formation of kinetochore-microtubule attachments. EMBO J 29:2953-2965
126. Lin YT, Chen Y, Wu G, Lee WH (2006) Hec1 sequentially recruits Zwint-1 and ZW10 to kinetochores for faithful chromosome segregation and spindle checkpoint control. Oncogene 25:6901-6914 (Pubitemid 44684831)
127. Liu ST, Chan GK, Hittle JC, Fujii G, Lees E, Yen TJ (2003) Human MPS1 kinase is required for mitotic arrest induced by the loss of CENP-E from kinetochores. Mol Biol Cell 14:1638-1651 (Pubitemid 36547428)
128. Liu D, Vader G, Vromans MJM, Lampson MA, Lens SMA (2009) Sensing chromosome bi-orientation by spatial separation of aurora B kinase from kinetochore substrates. Science 323:1350-1353
129. Liu D, Vleugel M, Backer CB, Hori T, Fukagawa T, Cheeseman IM, Lampson MA (2010) Regulated targeting of protein phosphatase 1 to the outer kinetochore by KNL1 opposes Aurora B kinase. J Cell Biol 188:809-820
130. Liu D, Davydenko O, Lampson MA (2012) Polo-like kinase-1 regulates kinetochore-microtubule dynamics and spindle checkpoint silencing. J Cell Biol 198:491-499
131. Logarinho E, Resende T, Torres C, Bousbaa H (2008) The human spindle assembly checkpoint protein bub3 is required for the establishment of efficient kinetochore-microtubule attachments. Mol Biol Cell 19:1798-1813 (Pubitemid 351805130)
132. London N, Ceto S, Ranish JA, Biggins S (2012) Phosphoregulation of Spc105 by Mps1 and PP1 regulates Bub1 localization to kinetochores. Curr Biol 22:900-906
133. Luo X, Yu H (2008) Protein metamorphosis: the two-state behavior of Mad2. Structure 16:1616-1625
134. Maciejowski J, George KA, Terret ME, Zhang C, Shokat KM, Jallepalli PV (2010) Mps1 directs the assembly of Cdc20 inhibitory complexes during interphase and mitosis to control M phase timing and spindle checkpoint signaling. J Cell Biol 190:89-100
135. Magidson V, O'Connell CB, Loncarek J, Paul R, Mogilner A, Khodjakov A (2011) The spatial arrangement of chromosomes during prometaphase facilitates spindle assembly. Cell 146:555-567
136. Maldonado M, Kapoor TM (2011) Constitutive Mad1 targeting to kinetochores uncouples checkpoint signalling from chromosome biorientation. Nat Cell Biol 13:475-482
137. Malureanu LA, Jeganathan KB, Hamada M, Wasilewski L, Davenport J, van Deursen JM (2009) BubR1 N terminus acts as a soluble inhibitor of cyclin B degradation by APC/C(Cdc20) in interphase. Dev Cell 16:118-131
138. Manning AL, Bakhoum SF, Maffini S, Correia-Melo C, Maiato H, Compton DA (2010) CLASP1, astrin and Kif2b form a molecular switch that regulates kinetochore-microtubule dynamics to promote mitotic progression and fidelity. EMBO J 29:3531-3543
139. Mansfeld J, Collin P, Collins MO, Choudhary JS, Pines J (2011) APC15 drives the turnover of MCC-CDC20 to make the spindle assembly checkpoint responsive to kinetochore attachment. Nat Cell Biol 13:1234-1243
140. Mao Y, Abrieu A, Cleveland DW (2003) Activating and silencing the mitotic checkpoint through CENP-E-dependent activation/inactivation of BubR1. Cell 114:87-98 (Pubitemid 36859843)
141. Mao Y, Desai A, Cleveland DW (2005) Microtubule capture by CENP-E silences BubR1-dependent mitotic checkpoint signaling. J Cell Biol 170:873-880 (Pubitemid 41306286)
142. Maresca TJ, Salmon ED (2009) Intrakinetochore stretch is associated with changes in kinetochore phosphorylation and spindle assembly checkpoint activity. J Cell Biol 184:373-381
143. Maresca TJ, Salmon ED (2010) Welcome to a new kind of tension: translating kinetochore mechanics into a wait-anaphase signal. J Cell Sci 123:825-835
144. Martinez-Exposito MJ, Kaplan KB, Copeland J, Sorger PK (1999) Retention of the BUB3 checkpoint protein on lagging chromosomes. Proc Natl Acad Sci U S A 96:8493-8498 (Pubitemid 29354819)
145. Martin-Lluesma S, Stucke VM, Nigg EA (2002) Role of Hec1 in spindle checkpoint signaling and kinetochore recruitment of Mad1/Mad2. Science 297:2267-2270
146. Matson DR, Demirel PB, Stukenberg PT, Burke DJ (2012) A conserved role for COMA/CENP-H/I/N kinetochore proteins in the spindle checkpoint. Genes Dev 26:542-547
147. Matsumura S, Toyoshima F, Nishida E (2007) Polo-like kinase 1 facilitates chromosome alignment during prometaphase through BubR1. J Biol Chem 282:15217-15227 (Pubitemid 47093386)
148. Maure JF, Kitamura E, Tanaka TU (2007) Mps1 kinase promotes sister-kinetochore bi-orientation by a tension-dependent mechanism. Curr Biol 17:2175-2182 (Pubitemid 350236864)
149. Mayer C, Filopei J, Batac J, Alford L, Paluh JL (2006) An extended anaphase signaling pathway for Mad2p includes microtubule organizing center proteins and multiple motor-dependent transitions. Cell Cycle 5:1456-1463 (Pubitemid 44153657)
150. Meadows JC, Shepperd LA, Vanoosthuyse V, Lancaster TC, Sochaj AM, Buttrick GJ, Hardwick KG, Millar JB (2011) Spindle checkpoint silencing requires association of PP1 to both Spc7 and kinesin-8 motors. Dev Cell 20:739-750
151. Meraldi P, Sorger PK (2005) A dual role for Bub1 in the spindle checkpoint and chromosome congression. EMBO J 24:1621-1633 (Pubitemid 40646451)
152. Meraldi P, Draviam VM, Sorger PK (2004) Timing and checkpoints in the regulation of mitotic progression. Dev Cell 7:45-60 (Pubitemid 39172687)
153. Miller SA, Johnson ML, Stukenberg PT (2008) Kinetochore attachments require an interaction between unstructured tails on microtubules and Ndc80(Hec1). Curr Biol 18:1785-1791
154. Minoshima Y, Hori T, Okada M, Kimura H, Haraguchi T, Hiraoka Y, Bao YC, Kawashima T, Kitamura T, Fukagawa T (2005) The constitutive centromere component CENP-50 is required for recovery from spindle damage. Mol Cell Biol 25:10315-10328 (Pubitemid 41681957)
155. Minshull J, Sun H, Tonks NK, Murray AW (1994) MAP-kinase dependent mitotic feedback arrest in Xenopus egg extracts. Cell 79:475-486 (Pubitemid 24346062)
156. Morrow CJ, Tighe A, Johnson VL, Scott MI, Ditchfield C, Taylor SS (2005) Bub1 and aurora B cooperate to maintain BubR1-mediated inhibition of APC/CCdc20. J Cell Sci 118:3639-3652 (Pubitemid 41309411)
157. Musacchio A, Salmon ED (2007) The spindle-assembly checkpoint in space and time. Nat Rev Mol Cell Biol 8:379-393 (Pubitemid 46643240)
158. Nagaoka SI, Hassold TJ, Hunt PA (2012) Human aneuploidy: mechanisms and new insights into an age-old problem. Nat Rev Genet 13:493-504
159. Nicklas RB, Ward SC, Gorbsky GJ (1995) Kinetochore chemistry is sensitive to tension and may link mitotic forces to a cell cycle checkpoint. J Cell Biol 130:929-939
160. Nishino M, Kurasawa Y, Evans R, Lin SH, Brinkley BR, Yu-Lee LY (2006) NudC is required for Plk1 targeting to the kinetochore and chromosome congression. Curr Biol 16:1414-1421 (Pubitemid 44066771)
161. Obuse C, Iwasaki O, Kiyomitsu T, Goshima G, Toyoda Y, Yanagida M (2004) A conserved Mis12 centromere complex is linked to heterochromatic HP1 and outer kinetochore protein Zwint-1. Nat Cell Biol 6:1135-1141 (Pubitemid 39468016)
162. Ohi R, Sapra T, Howard J, Mitchison TJ (2004) Differentiation of cytoplasmic and meiotic spindle assembly MCAK functions by Aurora B-dependent phosphorylation. Mol Biol Cell 15:2895-2906 (Pubitemid 38691877)
163. Ohkura H, Yanagida M (1991) S. pombe gene sds22+ essential for a midmitotic transition encodes a leucine-rich repeat protein that positively modulates protein phosphatase-1. Cell 64:149-157 (Pubitemid 121001146)
164. + gene required for chromosome disjoining encodes one of two putative type 1 protein phosphatases. Cell 57:997-1007 (Pubitemid 19168380)
165. Park JE, Erikson RL, Lee KS (2011) Feed-forward mechanism of converting biochemical cooperativity to mitotic processes at the kinetochore plate. Proc Natl Acad Sci U S A 108:8200-8205
166. Parry DH, Hickson GR, O'Farrell PH (2003) Cyclin B destruction triggers changes in kinetochore behavior essential for successful anaphase. Curr Biol 13:647-653 (Pubitemid 36453312)
167. Perera D, Taylor SS (2010) Sgo1 establishes the centromeric cohesion protection mechanism in G2 before subsequent Bub1-dependent recruitment in mitosis. J Cell Sci 123:653-659
168. Perpelescu M, Fukagawa T (2011) The ABCs of CENPs. Chromosoma 120:425-446
169. Peters JM (2006) The anaphase promoting complex/cyclosome: a machine designed to destroy. Nat Rev Mol Cell Biol 7:644-656 (Pubitemid 44268210)
170. Petersen J, Hagan IM (2003) S. pombe aurora kinase/survivin is required for chromosome condensation and the spindle checkpoint attachment response. Curr Biol 13:590-597 (Pubitemid 36391938)
171. Petrovic A, Pasqualato S, Dube P, Krenn V, Santaguida S, Cittaro D, Monzani S, Massimiliano L, Keller J, Tarricone A, Maiolica A, Stark H, Musacchio A (2010) The MIS12 complex is a protein interaction hub for outer kinetochore assembly. J Cell Biol 190:835-852
172. Pinsky BA, Kotwaliwale CV, Tatsutani SY, Breed CA, Biggins S (2006a) Glc7/protein phosphatase 1 regulatory subunits can oppose the Ipl1/aurora protein kinase by redistributing Glc7. Mol Cell Biol 26:2648-2660
173. Pinsky BA, Kung C, Shokat KM, Biggins S (2006b) The Ipl1-Aurora protein kinase activates the spindle checkpoint by creating unattached kinetochores. Nat Cell Biol 8:78-83 (Pubitemid 43064800)
174. Pinsky BA, Nelson CR, Biggins S (2009) Protein phosphatase 1 regulates exit from the spindle checkpoint in budding yeast. Curr Biol 19:1182-1187
175. Pomerening JR, Sontag ED, Ferrell JE Jr (2003) Building a cell cycle oscillator: hysteresis and bistability in the activation of Cdc2. Nat Cell Biol 5:346-351 (Pubitemid 36432300)
176. Posch M, Khoudoli GA, Swift S, King EM, DeLuca JG, Swedlow JR (2010) Sds22 regulates aurora B activity and microtubule-kinetochore interactions at mitosis. J Cell Biol 191:61-74
177. Przewloka MR, Glover DM (2009) The kinetochore and the centromere: a working long distance relationship. Annu Rev Genet 43:439-465
178. Qi W, Tang Z, Yu H (2006) Phosphorylation- and polo-box-dependent binding of Plk1 to Bub1 is required for the kinetochore localization of Plk1. Mol Biol Cell 17:3705-3716 (Pubitemid 44156460)
179. Reddy SK, Rape M, Margansky WA, Kirschner MW (2007) Ubiquitination by the anaphase-promoting complex drives spindle checkpoint inactivation. Nature 446:921-925 (Pubitemid 46633168)
180. Ricke RM, Jeganathan KB, Malureanu L, Harrison AM, van Deursen JM (2012) Bub1 kinase activity drives error correction and mitotic checkpoint control but not tumor suppression. J Cell Biol 199:931-949
181. Riedel CG, Katis VL, Katou Y, Mori S, Itoh T, Helmhart W, Galova M, Petronczki M, Gregan J, Cetin B, Mudrak I, Ogris E, Mechtler K, Pelletier L, Buchholz F, Shirahige K, Nasmyth K (2006) Protein phosphatase 2A protects centromeric sister chromatid cohesion during meiosis I. Nature 441:53-61
182. Rieder CL, Alexander SP (1990) Kinetochores are transported poleward along a single astral microtubule during chromosome attachment to the spindle in newt lung cells. J Cell Biol 110(1):81-95 (Pubitemid 20035880)
183. Rieder CL, Schultz A, Cole R, Sluder G (1994) Anaphase onset in vertebrate somatic cells is controlled by a checkpoint that monitors sister kinetochore attachment to the spindle. J Cell Biol 127:1301-1310 (Pubitemid 24362564)
184. Rieder CL, Cole RW, Khodjakov A, Sluder G (1995) The checkpoint delaying anaphase in response to chromosome monoorientation is mediated by an inhibitory signal produced by unattached kinetochores. J Cell Biol 130:941-948
185. Rischitor PE, May KM, Hardwick KG (2007) Bub1 is a fission yeast kinetochore scaffold protein, and is sufficient to recruit other spindle checkpoint proteins to ectopic sites on chromosomes. PLoS One 2:e1342 (Pubitemid 351736602)
186. Rosasco-Nitcher SE, Lan W, Khorasanizadeh S, Stukenberg PT (2008) Centromeric Aurora-B activation requires TD-60, microtubules, and substrate priming phosphorylation. Science 319:469-472 (Pubitemid 351160581)
187. Rosenberg JS, Cross FR, Funabiki H (2011) KNL1/Spc105 recruits PP1 to silence the spindle assembly checkpoint. Curr Biol 21:942-947
188. Ryu H, Furuta M, Kirkpatrick D, Gygi SP, Azuma Y (2010) PIASy-dependent SUMOylation regulates DNA topoisomerase IIalpha activity. J Cell Biol 191:783-794
189. Sakuno T, Tada K, Watanabe Y (2009) Kinetochore geometry defined by cohesion within the centromere. Nature 458:852-858
190. Salimian KJ, Ballister ER, Smoak EM, Wood S, Panchenko T, Lampson MA, Black BE (2011) Feedback control in sensing chromosome biorientation by the Aurora B kinase. Curr Biol 21:1158-1165
191. Santaguida S, Vernieri C, Villa F, Ciliberto A, Musacchio A (2011) Evidence that Aurora B is implicated in spindle checkpoint signalling independently of error correction. EMBO J 30:1508-1519
192. Saurin AT, van der Waal MS, Medema RH, Lens SM, Kops GJ (2011) Aurora B potentiates Mps1 activation to ensure rapid checkpoint establishment at the onset of mitosis. Nat Commun 2:316
193. Schaar BT, Chan GK, Maddox P, Salmon ED, Yen TJ (1997) CENP-E function at kinetochores is essential for chromosome alignment. J Cell Biol 139:1373-1382 (Pubitemid 28013659)
194. Schmidt JC, Kiyomitsu T, Hori T, Backer CB, Fukagawa T, Cheeseman IM (2010) Aurora B kinase controls the targeting of the Astrin-SKAP complex to bioriented kinetochores. J Cell Biol 191:269-280
195. Schmidt JC, Arthanari H, Boeszoermenyi A, Dashkevich NM, Wilson-Kubalek EM, Monnier N, Markus M, Oberer M, Milligan RA, Bathe M, Wagner G, Grishchuk EL, Cheeseman IM (2012) The kinetochore-bound Ska1 complex tracks depolymerizing microtubules and binds to curved protofilaments. Dev Cell 23(5):968-980
196. Sczaniecka M, Feoktistova A, May KM, Chen JS, Blyth J, Gould KL, Hardwick KG (2008) The spindle checkpoint functions of Mad3 and Mad2 depend on a Mad3 KEN box-mediated interaction with Cdc20-anaphase-promoting complex (APC/C). J Biol Chem 283:23039-23047
197. Shah JV, Botvinick E, Bonday Z, Furnari F, Berns M, Cleveland DW (2004) Dynamics of centromere and kinetochore proteins; implications for checkpoint signaling and silencing. Curr Biol 14:942-952 (Pubitemid 38722608)
198. Shapiro PS, Vaisberg E, Hunt AJ, Tolwinski NS, Whalen AM, McIntosh JR, Ahn NG (1998) Activation of the MKK/ERK pathway during somatic cell mitosis: direct interactions of active ERK with kinetochores and regulation of the mitotic 3F3/2 phosphoantigen. J Cell Biol 142:1533-1545 (Pubitemid 28452601)
199. Sharp-Baker H, Chen RH (2001) Spindle checkpoint protein Bub1 is required for kinetochore localization of Mad1, Mad2, Bub3, and CENP-E, independently of its kinase activity. J Cell Biol 153:1239-1250 (Pubitemid 34289262)
200. Shepperd LA, Meadows JC, Sochaj AM, Lancaster TC, Zou J, Buttrick GJ, Rappsilber J, Hardwick KG, Millar JB (2012) Phosphodependent recruitment of Bub1 and Bub3 to Spc7/KNL1 by Mph1 kinase maintains the spindle checkpoint. Curr Biol 22:891-899
201. Sliedrecht T, Zhang C, Shokat KM, Kops GJ (2010) Chemical genetic inhibition of Mps1 in stable human cell lines reveals novel aspects of Mps1 function in mitosis. PLoS One 5:e10251
202. Stegmeier F, Rape M, Draviam VM, Nalepa G, Sowa ME, Ang XL, McDonald ER 3rd, Li MZ, Hannon GJ, Sorger PK, Kirschner MW, Harper JW, Elledge SJ (2007) Anaphase initiation is regulated by antagonistic ubiquitination and deubiquitination activities. Nature 446:876-881 (Pubitemid 46633164)
203. Stone EM, Yamano H, Kinoshita N, Yanagida M (1993) Mitotic regulation of protein phosphatases by the fission yeast sds22 protein. Curr Biol 3:13-26
204. Stucke VM, Sillje HH, Arnaud L, Nigg EA (2002) Human Mps1 kinase is required for the spindle assembly checkpoint but not for centrosome duplication. EMBO J 21:1723-1732 (Pubitemid 34614628)
205. Stucke VM, Baumann C, Nigg EA (2004) Kinetochore localization and microtubule interaction of the human spindle checkpoint kinase Mps1. Chromosoma 113:1-15 (Pubitemid 39236717)
206. Stumpff J, von Dassow G, Wagenbach M, Asbury C, Wordeman L (2008) The kinesin-8 motor Kif18A suppresses kinetochore movements to control mitotic chromosome alignment. Dev Cell 14:252-262 (Pubitemid 351189173)
207. Sudakin V, Chan GK, Yen TJ (2001) Checkpoint inhibition of the APC/C in HeLa cells is mediated by a complex of BUBR1, BUB3, CDC20, and MAD2. J Cell Biol 154:925-936 (Pubitemid 34286173)
208. Suijkerbuijk SJ, van Dam TJ, Karagoz GE, von Castelmur E, Hubner NC, Duarte AM, Vleugel M, Perrakis A, Rudiger SG, Snel B, Kops GJ (2012a) The vertebrate mitotic checkpoint protein BUBR1 is an unusual pseudokinase. Dev Cell 22:1321-1329
209. Suijkerbuijk SJ, Vleugel M, Teixeira A, Kops GJ (2012b) Integration of kinase and phosphatase activities by BUBR1 ensures formation of stable kinetochore-microtubule attachments. Dev Cell 23:745-755
210. Suzuki A, Hori T, Nishino T, Usukura J, Miyagi A, Morikawa K, Fukagawa T (2011) Spindle microtubules generate tension-dependent changes in the distribution of inner kinetochore proteins. J Cell Biol 193:125-140
211. Takeda DY, Wohlschlegel JA, Dutta A (2001) A bipartite substrate recognition motif for cyclin-dependent kinases. J Biol Chem 276:1993-1997 (Pubitemid 32109678)
212. Takenaka K, Gotoh Y, Nishida E (1997) MAP kinase is required for the spindle assembly checkpoint but is dispensable for the normal M phase entry and exit in Xenopus egg cell cycle extracts. J Cell Biol 136:1091-1097 (Pubitemid 27131740)
213. Takeuchi K, Fukagawa T (2012) Molecular architecture of vertebrate kinetochores. Exp Cell Res 318:1367-1374
214. Tanaka TU (2010) Kinetochore-microtubule interactions: steps towards bi-orientation. EMBO J 29:4070-4082
215. Tanaka TU, Rachidi N, Janke C, Pereira G, Galova M, Schiebel E, Stark MJ, Nasmyth K (2002) Evidence that the Ipl1-Sli15 (Aurora kinase-INCENP) complex promotes chromosome bi-orientation by altering kinetochore-spindle pole connections. Cell 108:317-329 (Pubitemid 34178397)
216. Tanaka K, Mukae N, Dewar H, van Breugel M, James EK, Prescott AR, Antony C, Tanaka TU (2005) Molecular mechanisms of kinetochore capture by spindle microtubules. Nature 434:987-994 (Pubitemid 40586041)
217. Tang Z, Bharadwaj R, Li B, Yu H (2001) Mad2-independent inhibition of APCCdc20 by the mitotic checkpoint protein BubR1. Dev Cell 1:227-237 (Pubitemid 33653058)
218. Tang Z, Shu H, Oncel D, Chen S, Yu H (2004) Phosphorylation of Cdc20 by Bub1 provides a catalytic mechanism for APC/C inhibition by the spindle checkpoint. Mol Cell 16:387-397 (Pubitemid 39504793)
219. Tang Z, Shu H, Qi W, Mahmood NA, Mumby MC, Yu H (2006) PP2A is required for centromeric localization of Sgo1 and proper chromosome segregation. Dev Cell 10:575-585
220. Tanno Y, Kitajima TS, Honda T, Ando Y, Ishiguro K, Watanabe Y (2010) Phosphorylation of mammalian Sgo2 by Aurora B recruits PP2A and MCAK to centromeres. Genes Dev 24:2169-2179
221. Taylor SS, McKeon F (1997) Kinetochore localization of murine Bub1 is required for normal mitotic timing and checkpoint response to spindle damage. Cell 89:727-735 (Pubitemid 27516175)
222. Taylor SS, Ha E, McKeon F (1998) The human homologue of Bub3 is required for kinetochore localization of Bub1 and a Mad3/Bub1-related protein kinase. J Cell Biol 142:1-11 (Pubitemid 28341135)
223. Teichner A, Eytan E, Sitry-Shevah D, Miniowitz-Shemtov S, Dumin E, Gromis J, Hershko A (2011) p31comet promotes disassembly of the mitotic checkpoint complex in an ATP-dependent process. Proc Natl Acad Sci U S A 108:3187-3192
224. Tien JF, Umbreit NT, Gestaut DR, Franck AD, Cooper J, Wordeman L, Gonen T, Asbury CL, Davis TN (2010) Cooperation of the Dam1 and Ndc80 kinetochore complexes enhances microtubule coupling and is regulated by aurora B. J Cell Biol 189:713-723
225. Tighe A, Staples O, Taylor S (2008) Mps1 kinase activity restrains anaphase during an unperturbed mitosis and targets Mad2 to kinetochores. J Cell Biol 181:893-901 (Pubitemid 351846823)
226. Tooley JG, Miller SA, Stukenberg PT (2011) The Ndc80 complex uses a tripartite attachment point to couple microtubule depolymerization to chromosome movement. Mol Biol Cell 22:1217-1226
227. Trinkle-Mulcahy L, Andrews PD, Wickramasinghe S, Sleeman J, Prescott A, Lam YW, Lyon C, Swedlow JR, Lamond AI (2003) Time-lapse imaging reveals dynamic relocalization of PP1gamma throughout the mammalian cell cycle. Mol Biol Cell 14:107-117 (Pubitemid 36106023)
228. Trinkle-Mulcahy L, Andersen J, Lam YW, Moorhead G, Mann M, Lamond AI (2006) Repo-Man recruits PP1 gamma to chromatin and is essential for cell viability. J Cell Biol 172:679-692
229. Tseng BS, Tan L, Kapoor TM, Funabiki H (2010) Dual detection of chromosomes and microtubules by the chromosomal passenger complex drives spindle assembly. Dev Cell 18:903-912
230. Tsukahara T, Tanno Y, Watanabe Y (2010) Phosphorylation of the CPC by Cdk1 promotes chromosome bi-orientation. Nature 467:719-723
231. Ubersax JA, Woodbury EL, Quang PN, Paraz M, Blethrow JD, Shah K, Shokat KM, Morgan DO (2003) Targets of the cyclin-dependent kinase Cdk1. Nature 425:859-864 (Pubitemid 37351471)
232. Uchida KS, Takagaki K, Kumada K, Hirayama Y, Noda T, Hirota T (2009) Kinetochore stretching inactivates the spindle assembly checkpoint. J Cell Biol 184:383-390
233. Uzunova K, Dye BT, Schutz H, Ladurner R, Petzold G, Toyoda Y, Jarvis MA, Brown NG, Poser I, Novatchkova M, Mechtler K, Hyman AA, Stark H, Schulman BA, Peters JM (2012) APC15 mediates CDC20 autoubiquitylation by APC/C(MCC) and disassembly of the mitotic checkpoint complex. Nat Struct Mol Biol 19:1116-1123
234. van der Waal MS, Hengeveld RC, van der Horst A, Lens SM (2012) Cell division control by the chromosomal passenger complex. Exp Cell Res 318:1407-1420
235. Vanoosthuyse V, Hardwick KG (2009) A novel protein phosphatase 1-dependent spindle checkpoint silencing mechanism. Curr Biol 19:1176-1181
236. Varma D, Monzo P, Stehman SA, Vallee RB (2008) Direct role of dynein motor in stable kinetochore-microtubule attachment, orientation, and alignment. J Cell Biol 182:1045-1054
237. Vigneron S, Prieto S, Bernis C, Labbe JC, Castro A, Lorca T (2004) Kinetochore localization of spindle checkpoint proteins: who controls whom? Mol Biol Cell 15:4584-4596 (Pubitemid 39299106)
238. Vorozhko VV, Emanuele MJ, Kallio MJ, Stukenberg PT, Gorbsky GJ (2008) Multiple mechanisms of chromosome movement in vertebrate cells mediated through the Ndc80 complex and dynein/dynactin. Chromosoma 117:169-179
239. Wan X, O'Quinn RP, Pierce HL, Joglekar AP, Gall WE, DeLuca JG, Carroll CW, Liu ST, Yen TJ, McEwen BF, Stukenberg PT, Desai A, Salmon ED (2009) Protein architecture of the human kinetochore microtubule attachment site. Cell 137:672-684
240. Wang XM, Zhai Y, Ferrell JE Jr (1997) A role for mitogen-activated protein kinase in the spindle assembly checkpoint in XTC cells. J Cell Biol 137:433-443 (Pubitemid 27181295)
241. Wang HW, Ramey VH, Westermann S, Leschziner AE, Welburn JP, Nakajima Y, Drubin DG, Barnes G, Nogales E (2007) Architecture of the Dam1 kinetochore ring complex and implications for microtubule-driven assembly and force-coupling mechanisms. Nat Struct Mol Biol 14:721-726 (Pubitemid 47220060)
242. Wang F, Dai J, Daum JR, Niedzialkowska E, Banerjee B, Stukenberg PT, Gorbsky GJ, Higgins JM (2010) Histone H3 Thr-3 phosphorylation by Haspin positions Aurora B at centromeres in mitosis. Science 330:231-235
243. Wang F, Ulyanova NP, van der Waal MS, Patnaik D, Lens SM, Higgins JM (2011) A positive feedback loop involving Haspin and Aurora B promotes CPC accumulation at centromeres in mitosis. Curr Biol 21:1061-1069
244. Wang F, Ulyanova NP, Daum JR, Patnaik D, Kateneva AV, Gorbsky GJ, Higgins JM (2012) Haspin inhibitors reveal centromeric functions of Aurora B in chromosome segregation. J Cell Biol 199:251-268
245. Warren CD, Brady DM, Johnston RC, Hanna JS, Hardwick KG, Spencer FA (2002) Distinct chromosome segregation roles for spindle checkpoint proteins. Mol Biol Cell 13:3029-3041 (Pubitemid 35034243)
246. Waters JC, Chen RH, Murray AW, Salmon ED (1998) Localization of Mad2 to kinetochores depends on microtubule attachment, not tension. J Cell Biol 141:1181-1191 (Pubitemid 28265612)
247. Weaver BA, Cleveland DW (2006) Does aneuploidy cause cancer? Curr Opin Cell Biol 18:658-667 (Pubitemid 44692476)
248. Wei RR, Al-Bassam J, Harrison SC (2007) The Ndc80/HEC1 complex is a contact point for kinetochore-microtubule attachment. Nat Struct Mol Biol 14:54-59 (Pubitemid 46067423)
249. Wei R, Ngo B, Wu G, Lee WH (2011) Phosphorylation of the Ndc80 complex protein, HEC1, by Nek2 kinase modulates chromosome alignment and signaling of the spindle assembly checkpoint. Mol Biol Cell 22:3584-3594
250. Welburn JPI, Vleugel M, Liu D, Iii JRY, Lampson MA, Fukagawa T, Cheeseman IM (2010) Aurora B phosphorylates spatially distinct targets to differentially regulate the kinetochore-microtubule interface. Mol Cell 38:383-392
251. Westhorpe FG, Tighe A, Lara-Gonzalez P, Taylor SS (2011) p31comet-mediated extraction of Mad2 from the MCC promotes efficient mitotic exit. J Cell Sci 124:3905-3916
252. Wong OK, Fang G (2007) Cdk1 phosphorylation of BubR1 controls spindle checkpoint arrest and Plk1-mediated formation of the 3F3/2 epitope. J Cell Biol 179:611-617 (Pubitemid 350146248)
253. Wood KW, Sakowicz R, Goldstein LS, Cleveland DW (1997) CENP-E is a plus end-directed kinetochore motor required for metaphase chromosome alignment. Cell 91:357-366 (Pubitemid 27467966)
254. Wurzenberger C, Held M, Lampson MA, Poser I, Hyman AA, Gerlich DW (2012) Sds22 and Repo-Man stabilize chromosome segregation by counteracting Aurora B on anaphase kinetochores. J Cell Biol 198:173-183
255. Yamagishi Y, Honda T, Tanno Y, Watanabe Y (2010) Two histone marks establish the inner centromere and chromosome bi-orientation. Science 330:239-243
256. Yamagishi Y, Yang CH, Tanno Y, Watanabe Y (2012) MPS1/Mph1 phosphorylates the kinetochore protein KNL1/Spc7 to recruit SAC components. Nat Cell Biol 14:746-752
257. Yamano H, Ishii K, Yanagida M (1994) Phosphorylation of dis2 protein phosphatase at the C-terminal cdc2 consensus and its potential role in cell cycle regulation. EMBO J 13:5310-5318 (Pubitemid 24351809)
258. Yang M, Li B, Liu CJ, Tomchick DR, Machius M, Rizo J, Yu H, Luo X (2008) Insights into mad2 regulation in the spindle checkpoint revealed by the crystal structure of the symmetric mad2 dimer. PLoS Biol 6:e50 (Pubitemid 351485607)
259. Yasuda S, Oceguera-Yanez F, Kato T, Okamoto M, Yonemura S, Terada Y, Ishizaki T, Narumiya S (2004) Cdc42 and mDia3 regulate microtubule attachment to kinetochores. Nature 428:767-771 (Pubitemid 38508288)
260. Yeh E, Skibbens RV, Cheng JW, Salmon ED, Bloom K (1995) Spindle dynamics and cell cycle regulation of dynein in the budding yeast, Saccharomyces cerevisiae. J Cell Biol 130:687-700
261. Yue Z, Carvalho A, Xu Z, Yuan X, Cardinale S, Ribeiro S, Lai F, Ogawa H, Gudmundsdottir E, Gassmann R, Morrison CG, Ruchaud S, Earnshaw WC (2008) Deconstructing Survivin: comprehensive genetic analysis of Survivin function by conditional knockout in a vertebrate cell line. J Cell Biol 183:279-296
262. Zecevic M, Catling AD, Eblen ST, Renzi L, Hittle JC, Yen TJ, Gorbsky GJ, Weber MJ (1998) Active MAP kinase in mitosis: localization at kinetochores and association with the motor protein CENP-E. J Cell Biol 142:1547-1558 (Pubitemid 28452602)
263. Zhang J, Ahmad S, Mao Y (2007) BubR1 and APC/EB1 cooperate to maintain metaphase chromosome alignment. J Cell Biol 178:773-784 (Pubitemid 47347363)
264. Zhao Y, Chen RH (2006) Mps1 phosphorylation by MAP kinase is required for kinetochore localization of spindle-checkpoint proteins. Curr Biol 16:1764-1769 (Pubitemid 44691789)
265. Zhou T, Aumais JP, Liu X, Yu-Lee LY, Erikson RL (2003) A role for Plk1 phosphorylation of NudC in cytokinesis. Dev Cell 5:127-138 (Pubitemid 36827347)
266. Zich J, Sochaj AM, Syred HM, Milne L, Cook AG, Ohkura H, Rappsilber J, Hardwick KG (2012) Kinase activity of fission yeast Mph1 is required for Mad2 and Mad3 to stably bind the anaphase promoting complex. Curr Biol 22:296-301