Kinetochore Regulation of Anaphase and Cytokinesis

  • Scott Thomas
  • Kenneth B. Kaplan


Since the initial discovery of kinetochore antigens, a plethora of kinetochore-associated proteins have been identified. Their roles in connecting chromosomes to the mitotic spindle have been the subject of intense investigation. However, a surprising number of kinetochore proteins perform non-centromeric functions during mitosis. This class of kinetochore proteins has been best characterized through studies of the so-called “chromosomal passengers,” proteins that associate with kinetochores at the start of mitosis and then redistribute to the anaphase spindle. The conserved behavior of chromosomal passengers suggests that redistribution of kinetochore-associated proteins is a commonly used strategy for cells to temporally and spatially orchestrate mitotic events. The activities of chromosomal passengers are closely linked to cell cycle regulation, placing them in a position to transmit regulatory changes to the cell division machinery. As cells enter mitosis, chromosomal passengers alter chromatin organization. At kinetochores, they ensure that sister chromatids form proper attachments with the mitotic spindle. During anaphase, they organize spindle structures to direct the cytokinetic machinery. In this chapter, we will discuss the expanding role for chromosomal passengers in regulating anaphase events and how the redistribution of other kinetochore-associated proteins might contribute to the orderly progression of mitosis.


Kinetochore Protein Anaphase Promote Complex Microtubule Attachment Cdc14 Phosphatase Spindle Elongation 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Adams, R.R., M. Carmena, and W.C. Earnshaw. 2001. Chromosomal passengers and the (aurora) ABCs of mitosis. Trends Cell Biol. 11:49–54.PubMedCrossRefGoogle Scholar
  2. Adams, R.R., S.P. Wheatley, A.M. Gouldsworthy, S.E. Kandels-Lewis, M. Carmena, C. Smythe, D.L. Gerloff, and W.C. Earnshaw. 2000. INCENP binds the Aurora-related kinase AIRK2 and is required to target it to chromosomes, the central spindle and cleavage furrow. Curr Biol. 10:1075–8.PubMedCrossRefGoogle Scholar
  3. Akhmanova, A., and C.C. Hoogenraad. 2005. Microtubule plus-end-tracking proteins: mechanisms and functions. Curr Opin Cell Biol. 17:47–54.PubMedCrossRefGoogle Scholar
  4. Arnaud, L., J. Pines, and E.A. Nigg. 1998. GFP tagging reveals human Polo-like kinase 1 at the kinetochore/centromere region of mitotic chromosomes. Chromosoma. 107:424–9.PubMedCrossRefGoogle Scholar
  5. Bardin, A.J., and A. Amon. 2001. Men and sin: what's the difference? Nat Rev Mol Cell Biol. 2:815–26.PubMedCrossRefGoogle Scholar
  6. Biggins, S., and A.W. Murray. 2001. The budding yeast protein kinase Ipl1/Aurora allows the absence of tension to activate the spindle checkpoint. Genes Dev. 15:3118–29.PubMedCrossRefGoogle Scholar
  7. Bishop, J.D., and J.M. Schumacher. 2002. Phosphorylation of the carboxyl terminus of inner centromere protein (INCENP) by the Aurora B Kinase stimulates Aurora B kinase activity. J Biol Chem. 277:27577–80.PubMedCrossRefGoogle Scholar
  8. Bouck, D.C., and K.S. Bloom. 2005. The kinetochore protein Ndc10p is required for spindle stability and cytokinesis in yeast. Proc Natl Acad Sci USA. 102:5408–13.PubMedCrossRefGoogle Scholar
  9. Bourhis, E., S.G. Hymowitz, and A.G. Cochran. 2007. The mitotic regulator Survivin binds as a monomer to its functional interactor Borealin. J Biol Chem. 282:35018–23.PubMedCrossRefGoogle Scholar
  10. Buvelot, S., S.Y. Tatsutani, D. Vermaak, and S. Biggins. 2003. The budding yeast Ipl1/Aurora protein kinase regulates mitotic spindle disassembly. J Cell Biol. 160:329–39.PubMedCrossRefGoogle Scholar
  11. Cao, L., X. Yan, Y. Wu, H. Hu, Q. Li, T. Zhou, S. Jiang, and L. Yu. 2006. Survivin mutant (Surv-DD70, 71AA) disrupts the interaction of Survivin with Aurora B and causes multinucleation in HeLa cells. Biochem Biophys Res Commun. 346:400–7.PubMedCrossRefGoogle Scholar
  12. Carvalho, P., J.S. Tirnauer, and D. Pellman. 2003. Surfing on microtubule ends. Trends Cell Biol. 13:229–37.PubMedCrossRefGoogle Scholar
  13. Chan, C.S., and D. Botstein. 1993. Isolation and characterization of chromosome-gain and increase-in-ploidy mutants in yeast. Genetics. 135:677–91.PubMedGoogle Scholar
  14. Cheeseman, I.M., M. Enquist-Newman, T. Muller-Reichert, D.G. Drubin, and G. Barnes. 2001. Mitotic spindle integrity and kinetochore function linked by the Duo1p/Dam1p complex. J Cell Biol. 152:197–212.PubMedCrossRefGoogle Scholar
  15. Chiroli, E., V. Rossio, G. Lucchini, and S. Piatti. 2007. The budding yeast PP2ACdc55 protein phosphatase prevents the onset of anaphase in response to morphogenetic defects. J Cell Biol. 177:599–611.PubMedCrossRefGoogle Scholar
  16. Cohen-Fix, O., and D. Koshland. 1999. Pds1p of budding yeast has dual roles: inhibition of anaphase initiation and regulation of mitotic exit. Genes Dev. 13:1950–9.PubMedCrossRefGoogle Scholar
  17. Cooke, C.A., M.M. Heck, and W.C. Earnshaw. 1987. The inner centromere protein (INCENP) antigens: movement from inner centromere to midbody during mitosis. J Cell Biol. 105:2053–67.PubMedCrossRefGoogle Scholar
  18. Crosio, C., G.M. Fimia, R. Loury, M. Kimura, Y. Okano, H. Zhou, S. Sen, C.D. Allis, and P. Sassone-Corsi. 2002. Mitotic phosphorylation of histone H3: spatio-temporal regulation by mammalian Aurora kinases. Mol Cell Biol. 22:874–85.PubMedCrossRefGoogle Scholar
  19. Delcros, J.G., C. Prigent, and R. Giet. 2006. Dynactin targets Pavarotti-KLP to the central spindle during anaphase and facilitates cytokinesis in Drosophila S2 cells. J Cell Sci. 119:4431–41.PubMedCrossRefGoogle Scholar
  20. DeLuca, J.G., W.E. Gall, C. Ciferri, D. Cimini, A. Musacchio, and E.D. Salmon. 2006. Kinetochore microtubule dynamics and attachment stability are regulated by Hec1. Cell. 127:969–82.PubMedCrossRefGoogle Scholar
  21. Dobbelaere, J., and Y. Barral. 2004. Spatial coordination of cytokinetic events by compartmentalization of the cell cortex. Science. 305:393–6.PubMedCrossRefGoogle Scholar
  22. Dobbelaere, J., M.S. Gentry, R.L. Hallberg, and Y. Barral. 2003. Phosphorylation-dependent regulation of septin dynamics during the cell cycle. Dev Cell. 4:345–57.PubMedCrossRefGoogle Scholar
  23. Draviam, V.M., I. Shapiro, B. Aldridge, and P.K. Sorger. 2006. Misorientation and reduced stretching of aligned sister kinetochores promote chromosome missegregation in EB1- or APC-depleted cells. Embo J. 25:2814–27.PubMedCrossRefGoogle Scholar
  24. Dumitrescu, T.P., and W.S. Saunders. 2002. The FEAR Before MEN: networks of mitotic exit. Cell Cycle. 1:304–7.PubMedCrossRefGoogle Scholar
  25. Earnshaw, W.C., and R.L. Bernat. 1991. Chromosomal passengers: toward an integrated view of mitosis. Chromosoma. 100:139–46.PubMedCrossRefGoogle Scholar
  26. Eckley, D.M., A.M. Ainsztein, A.M. Mackay, I.G. Goldberg, and W.C. Earnshaw. 1997. Chromosomal proteins and cytokinesis: patterns of cleavage furrow formation and inner centromere protein positioning in mitotic heterokaryons and mid-anaphase cells. J Cell Biol. 136:1169–83.PubMedCrossRefGoogle Scholar
  27. Fortugno, P., E. Beltrami, J. Plescia, J. Fontana, D. Pradhan, P.C. Marchisio, W.C. Sessa, and D.C. Altieri. 2003. Regulation of survivin function by Hsp90. Proc Natl Acad Sci USA. 100:13791–6.PubMedCrossRefGoogle Scholar
  28. Gadea, B.B., and J.V. Ruderman. 2006. Aurora B is required for mitotic chromatin-induced phosphorylation of Op18/Stathmin. Proc Natl Acad Sci USA. 103:4493–8.PubMedCrossRefGoogle Scholar
  29. Gassmann, R., A. Carvalho, A.J. Henzing, S. Ruchaud, D.F. Hudson, R. Honda, E.A. Nigg, D.L. Gerloff, and W.C. Earnshaw. 2004. Borealin: a novel chromosomal passenger required for stability of the bipolar mitotic spindle. J Cell Biol. 166:179–91.PubMedCrossRefGoogle Scholar
  30. Giet, R., and D.M. Glover. 2001. Drosophila aurora B kinase is required for histone H3 phosphorylation and condensin recruitment during chromosome condensation and to organize the central spindle during cytokinesis. J Cell Biol. 152:669–82.PubMedCrossRefGoogle Scholar
  31. Gillis, A.N., S. Thomas, S.D. Hansen, and K.B. Kaplan. 2005. A novel role for the CBF3 kinetochore-scaffold complex in regulating septin dynamics and cytokinesis. J Cell Biol. 171:773–84.PubMedCrossRefGoogle Scholar
  32. Glotzer, M. 2005. The molecular requirements for cytokinesis. Science. 307:1735–9.PubMedCrossRefGoogle Scholar
  33. Goto, H., T. Kiyono, Y. Tomono, A. Kawajiri, T. Urano, K. Furukawa, E.A. Nigg, and M. Inagaki. 2006. Complex formation of Plk1 and INCENP required for metaphase-anaphase transition. Nat Cell Biol. 8:180–7.PubMedCrossRefGoogle Scholar
  34. Green, R.A., R. Wollman, and K.B. Kaplan. 2005. APC and EB1 Function Together in Mitosis to Regulate Spindle Dynamics and Chromosome Alignment. Mol Biol Cell. 16:4609–22.PubMedCrossRefGoogle Scholar
  35. Gruneberg, U., R. Neef, R. Honda, E.A. Nigg, and F.A. Barr. 2004. Relocation of Aurora B from centromeres to the central spindle at the metaphase to anaphase transition requires MKlp2. J Cell Biol. 166:167–72.PubMedCrossRefGoogle Scholar
  36. Hsu, J.Y., Z.W. Sun, X. Li, M. Reuben, K. Tatchell, D.K. Bishop, J.M. Grushcow, C.J. Brame, J.A. Caldwell, D.F. Hunt, R. Lin, M.M. Smith, and C.D. Allis. 2000. Mitotic phosphorylation of histone H3 is governed by Ipl1/aurora kinase and Glc7/PP1 phosphatase in budding yeast and nematodes. Cell. 102:279–91.PubMedCrossRefGoogle Scholar
  37. Jelluma, N., A.B. Brenkman, N.J. van den Broek, C.W. Cruijsen, M.H. van Osch, S.M. Lens, R.H. Medema, and G.J. Kops. 2008. Mps1 phosphorylates Borealin to control Aurora B activity and chromosome alignment. Cell. 132:233–46.PubMedCrossRefGoogle Scholar
  38. Jeyaprakash, A.A., U.R. Klein, D. Lindner, J. Ebert, E.A. Nigg, and E. Conti. 2007. Structure of a Survivin-Borealin-INCENP core complex reveals how chromosomal passengers travel together. Cell. 131:271–85.PubMedCrossRefGoogle Scholar
  39. Jones, M.H., X. He, T.H. Giddings, and M. Winey. 2001. Yeast Dam1p has a role at the kinetochore in assembly of the mitotic spindle. Proc Natl Acad Sci USA. 98:13675–80.PubMedCrossRefGoogle Scholar
  40. Kaiser, B.K., Z.A. Zimmerman, H. Charbonneau, and P.K. Jackson. 2002. Disruption of centrosome structure, chromosome segregation, and cytokinesis by misexpression of human Cdc14A phosphatase. Mol Biol Cell. 13:2289–300.PubMedCrossRefGoogle Scholar
  41. Klein, U.R., E.A. Nigg, and U. Gruneberg. 2006. Centromere targeting of the chromosomal passenger complex requires a ternary subcomplex of Borealin, Survivin, and the N-terminal domain of INCENP. Mol Biol Cell. 17:2547–58.PubMedCrossRefGoogle Scholar
  42. Lavoie, B.D., E. Hogan, and D. Koshland. 2004. In vivo requirements for rDNA chromosome condensation reveal two cell-cycle-regulated pathways for mitotic chromosome folding. Genes Dev. 18:76–87.PubMedCrossRefGoogle Scholar
  43. Li, D., R. Liu, L. Song, H. Zhou, J. Chen, and X. Huang. 2007. The special location of p-H3 and p-CENP-A on heterochromatin during mitosis in MCF-7. Mol Biol Rep. 10.1007/s11033-007-9136-9Google Scholar
  44. Li, J., W.L. Lee, and J.A. Cooper. 2005. NudEL targets dynein to microtubule ends through LIS1. Nat Cell Biol. 7:686–90.PubMedCrossRefGoogle Scholar
  45. Lingelbach, L.B., and K.B. Kaplan. 2004. The Interaction between Sgt1p and Skp1p Is Regulated by HSP90 Chaperones and Is Required for Proper CBF3 Assembly. Mol Cell Biol. 24:8938–50.PubMedCrossRefGoogle Scholar
  46. Lipp, J.J., T. Hirota, I. Poser, and J.M. Peters. 2007. Aurora B controls the association of condensin I but not condensin II with mitotic chromosomes. J Cell Sci. 120:1245–55.PubMedCrossRefGoogle Scholar
  47. Longtine, M.S., and E. Bi. 2003. Regulation of septin organization and function in yeast. Trends Cell Biol. 13:403–9.PubMedCrossRefGoogle Scholar
  48. Machin, F., J. Torres-Rosell, A. Jarmuz, and L. Aragon. 2005. Spindle-independent condensation-mediated segregation of yeast ribosomal DNA in late anaphase. J Cell Biol. 168:209–19.PubMedCrossRefGoogle Scholar
  49. Mackay, A.M., A.M. Ainsztein, D.M. Eckley, and W.C. Earnshaw. 1998. A dominant mutant of inner centromere protein (INCENP), a chromosomal protein, disrupts prometaphase congression and cytokinesis. J Cell Biol. 140:991–1002.PubMedCrossRefGoogle Scholar
  50. Mailand, N., C. Lukas, B.K. Kaiser, P.K. Jackson, J. Bartek, and J. Lukas. 2002. Deregulated human Cdc14A phosphatase disrupts centrosome separation and chromosome segregation. Nat Cell Biol. 4:317–22.PubMedCrossRefGoogle Scholar
  51. McAinsh, A.D., J.D. Tytell, and P.K. Sorger. 2003. Structure, function, and regulation of budding yeast kinetochores. Annu Rev Cell Dev Biol. 19:519–39.PubMedCrossRefGoogle Scholar
  52. Minoshima, Y., T. Kawashima, K. Hirose, Y. Tonozuka, A. Kawajiri, Y.C. Bao, X. Deng, M. Tatsuka, S. Narumiya, W.S. May, Jr., T. Nosaka, K. Semba, T. Inoue, T. Satoh, M. Inagaki, and T. Kitamura. 2003. Phosphorylation by aurora B converts MgcRacGAP to a RhoGAP during cytokinesis. Dev Cell. 4:549–60.PubMedCrossRefGoogle Scholar
  53. Miranda, J.J., P. De Wulf, P.K. Sorger, and S.C. Harrison. 2005. The yeast DASH complex forms closed rings on microtubules. Nat Struct Mol Biol. 12:138–43.PubMedCrossRefGoogle Scholar
  54. Miranda, J.J., D.S. King, and S.C. Harrison. 2007. Protein arms in the kinetochore-microtubule interface of the yeast DASH complex. Mol Biol Cell. 18:2503–10.PubMedCrossRefGoogle Scholar
  55. Mishima, M., S. Kaitna, and M. Glotzer. 2002. Central spindle assembly and cytokinesis require a kinesin-like protein/RhoGAP complex with microtubule bundling activity. Dev Cell. 2:41–54.PubMedCrossRefGoogle Scholar
  56. Miyauchi, K., X. Zhu, C. Foong, H. Hosoya, and M. Murata-Hori. 2007. Aurora B kinase activity is required to prevent polar cortical ingression during cytokinesis. Cell Cycle. 6:2549–53.PubMedCrossRefGoogle Scholar
  57. Mollinari, C., C. Reynaud, S. Martineau-Thuillier, S. Monier, S. Kieffer, J. Garin, P.R. Andreassen, A. Boulet, B. Goud, J.P. Kleman, and R.L. Margolis. 2003. The mammalian passenger protein TD-60 is an RCC1 family member with an essential role in prometaphase to metaphase progression. Dev Cell. 5:295–307.PubMedCrossRefGoogle Scholar
  58. Morgan, D.O. 1999. Regulation of the APC and the exit from mitosis. Nat Cell Biol. 1:E47–53.PubMedCrossRefGoogle Scholar
  59. Murnion, M.E., R.R. Adams, D.M. Callister, C.D. Allis, W.C. Earnshaw, and J.R. Swedlow. 2001. Chromatin-associated protein phosphatase 1 regulates aurora-B and histone H3 phosphorylation. J Biol Chem. 276:26656–65.PubMedCrossRefGoogle Scholar
  60. Neef, R., U.R. Klein, R. Kopajtich, and F.A. Barr. 2006. Cooperation between mitotic kinesins controls the late stages of cytokinesis. Curr Biol. 16:301–7.PubMedCrossRefGoogle Scholar
  61. Niikura, Y., S. Ohta, K.J. Vandenbeldt, R. Abdulle, B.F. McEwen, and K. Kitagawa. 2006. 17-AAG, an Hsp90 inhibitor, causes kinetochore defects: a novel mechanism by which 17-AAG inhibits cell proliferation. Oncogene. 25:4133–46.Google Scholar
  62. Norden, C., M. Mendoza, J. Dobbelaere, C.V. Kotwaliwale, S. Biggins, and Y. Barral. 2006. The NoCut pathway links completion of cytokinesis to spindle midzone function to prevent chromosome breakage. Cell. 125:85–98.PubMedCrossRefGoogle Scholar
  63. Parry, D.H., G.R. Hickson, and P.H. O'Farrell. 2003. Cyclin B destruction triggers changes in kinetochore behavior essential for successful anaphase. Curr Biol. 13:647–53.PubMedCrossRefGoogle Scholar
  64. Parry, D.H., and P.H. O'Farrell. 2001. The schedule of destruction of three mitotic cyclins can dictate the timing of events during exit from mitosis. Curr Biol. 11:671–83.PubMedCrossRefGoogle Scholar
  65. Pavicic-Kaltenbrunner, V., M. Mishima, and M. Glotzer. 2007. Cooperative assembly of CYK-4/MgcRacGAP and ZEN-4/MKLP1 to form the centralspindlin complex. Mol Biol Cell. 18:4992–5003.PubMedCrossRefGoogle Scholar
  66. Pellman, D., M. Bagget, Y.H. Tu, G.R. Fink, and H. Tu. 1995. Two microtubule-associated proteins required for anaphase spindle movement in Saccharomyces cerevisiae. J Cell Biol. 130:1373–85.PubMedCrossRefGoogle Scholar
  67. Pereira, G., and E. Schiebel. 2001. The role of the yeast spindle pole body and the mammalian centrosome in regulating late mitotic events. Curr Opin Cell Biol. 13:762–9.PubMedCrossRefGoogle Scholar
  68. Pereira, G., and E. Schiebel. 2003. Separase regulates INCENP-Aurora B anaphase spindle function through Cdc14. Science. 302:2120–4.PubMedCrossRefGoogle Scholar
  69. Queralt, E., C. Lehane, B. Novak, and F. Uhlmann. 2006. Downregulation of PP2A(Cdc55) phosphatase by separase initiates mitotic exit in budding yeast. Cell. 125:719–32.PubMedCrossRefGoogle Scholar
  70. Rattner, J.B., A. Rao, M.J. Fritzler, D.W. Valencia, and T.J. Yen. 1993. CENP-F is a .ca 400 kDa kinetochore protein that exhibits a cell-cycle dependent localization. Cell Motil Cytoskeleton. 26:214–26.PubMedCrossRefGoogle Scholar
  71. Rogers, S.L., U. Wiedemann, U. Hacker, C. Turck, and R.D. Vale. 2004. Drosophila RhoGEF2 associates with microtubule plus ends in an EB1-dependent manner. Curr Biol. 14:1827–33.PubMedCrossRefGoogle Scholar
  72. Romano, A., A. Guse, I. Krascenicova, H. Schnabel, R. Schnabel, and M. Glotzer. 2003. CSC-1: a subunit of the Aurora B kinase complex that binds to the survivin-like protein BIR-1 and the incenp-like protein ICP-1. J Cell Biol. 161:229–36.PubMedCrossRefGoogle Scholar
  73. Rosa, J., P. Canovas, A. Islam, D.C. Altieri, and S.J. Doxsey. 2006. Survivin modulates microtubule dynamics and nucleation throughout the cell cycle. Mol Biol Cell. 17:1483–93.PubMedCrossRefGoogle Scholar
  74. Rosasco-Nitcher, S.E., W. Lan, S. Khorasanizadeh, and P.T. Stukenberg. 2008. Centromeric Aurora-B activation requires TD-60, microtubules, and substrate priming phosphorylation. Science. 319:469–72.PubMedCrossRefGoogle Scholar
  75. Ruchaud, S., M. Carmena, and W.C. Earnshaw. 2007. Chromosomal passengers: conducting cell division. Nat Rev Mol Cell Biol. 8:798–812.PubMedCrossRefGoogle Scholar
  76. Sandall, S., F. Severin, I.X. McLeod, J.R. Yates, 3rd, K. Oegema, A. Hyman, and A. Desai. 2006. A Bir1-Sli15 complex connects centromeres to microtubules and is required to sense kinetochore tension. Cell. 127:1179–91.PubMedCrossRefGoogle Scholar
  77. Scholey, J.M., I. Brust-Mascher, and A. Mogilner. 2003. Cell division. Nature. 422:746–52.PubMedCrossRefGoogle Scholar
  78. Schuyler, S.C., J.Y. Liu, and D. Pellman. 2003. The molecular function of Ase1p: evidence for a MAP-dependent midzone-specific spindle matrix. Microtubule-associated proteins. J Cell Biol. 160:517–28.PubMedCrossRefGoogle Scholar
  79. Severson, A.F., D.R. Hamill, J.C. Carter, J. Schumacher, and B. Bowerman. 2000. The aurora-related kinase AIR-2 recruits ZEN-4/CeMKLP1 to the mitotic spindle at metaphase and is required for cytokinesis. Curr Biol. 10:1162–71.PubMedCrossRefGoogle Scholar
  80. Shannon, K.B., and E.D. Salmon. 2002. Chromosome dynamics: new light on Aurora B kinase function. Curr Biol. 12:R458–60.PubMedCrossRefGoogle Scholar
  81. Shirayama, M., A. Toth, M. Galova, and K. Nasmyth. 1999. APC(Cdc20) promotes exit from mitosis by destroying the anaphase inhibitor Pds1 and cyclin Clb5 [see comments]. Nature. 402:203–7.PubMedCrossRefGoogle Scholar
  82. Steensgaard, P., M. Garre, I. Muradore, P. Transidico, E.A. Nigg, K. Kitagawa, W.C. Earnshaw, M. Faretta, and A. Musacchio. 2004. Sgt1 is required for human kinetochore assembly. EMBO Rep. 5:626–31.PubMedCrossRefGoogle Scholar
  83. Stehman, S.A., Y. Chen, R.J. McKenney, and R.B. Vallee. 2007. NudE and NudEL are required for mitotic progression and are involved in dynein recruitment to kinetochores. J Cell Biol. 178:583–94.PubMedCrossRefGoogle Scholar
  84. Stemmann, O., A. Neidig, T. Kocher, M. Wilm, and J. Lechner. 2002. Hsp90 enables Ctf13p/Skp1p to nucleate the budding yeast kinetochore. Proc Natl Acad Sci USA. 99:8585–90.PubMedCrossRefGoogle Scholar
  85. Stoepel, J., M.A. Ottey, C. Kurischko, P. Hieter, and F.C. Luca. 2005. The mitotic exit network Mob1p-Dbf2p kinase complex localizes to the nucleus and regulates passenger protein localization. Mol Biol Cell. 16:5465–79.PubMedCrossRefGoogle Scholar
  86. Strickland, L.I., Y. Wen, G.G. Gundersen, and D.R. Burgess. 2005. Interaction between EB1 and p150glued is required for anaphase astral microtubule elongation and stimulation of cytokinesis. Curr Biol. 15:2249–55.PubMedCrossRefGoogle Scholar
  87. Strunnikov, A.V. 2005. A case of selfish nucleolar segregation. Cell Cycle. 4:113–7.PubMedCrossRefGoogle Scholar
  88. Sullivan, M., and D.O. Morgan. 2007. Finishing mitosis, one step at a time. Nat Rev Mol Cell Biol. 8:894–903.PubMedCrossRefGoogle Scholar
  89. Sumara, I., M. Quadroni, C. Frei, M.H. Olma, G. Sumara, R. Ricci, and M. Peter. 2007. A Cul3-based E3 ligase removes Aurora B from mitotic chromosomes, regulating mitotic progression and completion of cytokinesis in human cells. Dev Cell. 12:887–900.PubMedCrossRefGoogle Scholar
  90. Tang, X., and Y. Wang. 2006. Pds1/Esp1-dependent and -independent sister chromatid separation in mutants defective for protein phosphatase 2A. Proc Natl Acad Sci USA. 103:16290–5.PubMedCrossRefGoogle Scholar
  91. Thomas, S., and K.B. Kaplan. 2007. A Bir1p Sli15p kinetochore passenger complex regulates septin organization during anaphase. Mol Biol Cell. 18:3820–34.PubMedCrossRefGoogle Scholar
  92. Tinker-Kulberg, R.L., and D.O. Morgan. 1999. Pds1 and Esp1 control both anaphase and mitotic exit in normal cells and after DNA damage. Genes Dev. 13:1936–49.PubMedCrossRefGoogle Scholar
  93. Torres-Rosell, J., F. Machin, and L. Aragon. 2005. Cdc14 and the temporal coordination between mitotic exit and chromosome segregation. Cell Cycle. 4:109–12.PubMedCrossRefGoogle Scholar
  94. Trinkle-Mulcahy, L., and A.I. Lamond. 2006. Mitotic phosphatases: no longer silent partners. Curr Opin Cell Biol. 18:623–31.PubMedCrossRefGoogle Scholar
  95. Vagnarelli, P., and W.C. Earnshaw. 2004. Chromosomal passengers: the four-dimensional regulation of mitotic events. Chromosoma. 113:211–22.PubMedCrossRefGoogle Scholar
  96. Vagnarelli, P., D.F. Hudson, S.A. Ribeiro, L. Trinkle-Mulcahy, J.M. Spence, F. Lai, C.J. Farr, A.I. Lamond, and W.C. Earnshaw. 2006. Condensin and Repo-Man-PP1 co-operate in the regulation of chromosome architecture during mitosis. Nat Cell Biol. 8:1133–42.PubMedCrossRefGoogle Scholar
  97. Westermann, S., H.W. Wang, A. Avila-Sakar, D.G. Drubin, E. Nogales, and G. Barnes. 2006. The Dam1 kinetochore ring complex moves processively on depolymerizing microtubule ends. Nature. 440:565–9.PubMedCrossRefGoogle Scholar
  98. Widlund, P.O., J.S. Lyssand, S. Anderson, S. Niessen, J.R. Yates, 3rd, and T.N. Davis. 2006. Phosphorylation of the chromosomal passenger protein Bir1 is required for localization of Ndc10 to the spindle during anaphase and full spindle elongation. Mol Biol Cell. 17:1065–74.PubMedCrossRefGoogle Scholar
  99. Wolyniak, M.J., K. Blake-Hodek, K. Kosco, E. Hwang, L. You, and T.C. Huffaker. 2006. The regulation of microtubule dynamics in Saccharomyces cerevisiae by three interacting plus-end tracking proteins. Mol Biol Cell. 17:2789–98.PubMedCrossRefGoogle Scholar
  100. Yan, X., F. Li, Y. Liang, Y. Shen, X. Zhao, Q. Huang, and X. Zhu. 2003. Human Nudel and NudE as regulators of cytoplasmic dynein in poleward protein transport along the mitotic spindle. Mol Cell Biol. 23:1239–50.PubMedCrossRefGoogle Scholar
  101. Yang, D., A. Welm, and J.M. Bishop. 2004. Cell division and cell survival in the absence of survivin. Proc Natl Acad Sci USA. 101:15100–5.PubMedCrossRefGoogle Scholar
  102. Zeitlin, S.G., R.D. Shelby, and K.F. Sullivan. 2001. CENP-A is phosphorylated by Aurora B kinase and plays an unexpected role in completion of cytokinesis. J Cell Biol. 155:1147–57.PubMedCrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC 2009

Authors and Affiliations

  1. 1.Department of Molecular and Cellular BiologyUniversity of California, Davis, One Shields Ave.DavisU.S.A

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