Cytokinesis in animal cells

Annual Review of Cell and Developmental Biology

Volumn 28, Issue , 2012, Pages 29-58

  Green, Rebecca A ^a   Paluch, Ewa ^b,c   Oegema, Karen ^a  

^a UNIVERSITY OF CALIFORNIA   (United States)

^b MAX PLANCK INSTITUTE OF MOLECULAR CELL BIOLOGY AND GENETICS   (Germany)

^c INTERNATIONAL INSTITUTE OF MOLECULAR AND CELL BIOLOGY   (Poland)

Author keywords

abscission; actin cortex; cell division; cell mechanics; central spindle; contractile ring

Indexed keywords

ANAPHASE PROMOTING COMPLEX; CYCLIN DEPENDENT KINASE 1; RHOA GUANINE NUCLEOTIDE BINDING PROTEIN;

ABSCISSION ZONE; ANAPHASE; ANIMAL CELL; CELL CYCLE; CELL MATURATION; CELL STRUCTURE; CYTOKINESIS; MICROFILAMENT; MICROTUBULE ASSEMBLY; MOLECULAR DYNAMICS; NONHUMAN; PRIORITY JOURNAL; PROTEIN TARGETING; REVIEW; SCHIZOSACCHAROMYCES POMBE; SIGNAL TRANSDUCTION; SPINDLE CELL;

ANAPHASE; ANIMALS; CELL CYCLE PROTEINS; CYTOKINESIS; CYTOSKELETAL PROTEINS; HUMANS; MITOTIC SPINDLE APPARATUS; MODELS, BIOLOGICAL; SIGNAL TRANSDUCTION;

ANIMALIA;

EID: 84866344301     PISSN: 10810706     EISSN: 15308995     Source Type: Book Series    
DOI: 10.1146/annurev-cellbio-101011-155718     Document Type: Review

References (214)

1. Barr FA, Gruneberg U. 2007. Cytokinesis: placing and making the final cut. Cell 131:847-60
2. Bastos RN, Barr FA. 2010. Plk1 negatively regulates Cep55 recruitment to the midbody to ensure orderly abscission. J. Cell Biol. 191:751-60
3. Bathe M, Chang F. 2010. Cytokinesis and the contractile ring in fission yeast: towards a systems-level understanding. Trends Microbiol. 18:38-45
4. Bement WM,BeninkHA, vonDassowG. 2005.Amicrotubule-dependent zone of activeRhoA during cleavage plane specification. J. Cell Biol. 170:91-101
5. Bement WM, Miller AL, Von Dassow G. 2006. Rho GTPase activity zones and transient contractile arrays. BioEssays 28:983-93
6. Bieling P, Telley IA, Surrey T. 2010. A minimal midzone protein module controls formation and length of antiparallel microtubule overlaps. Cell 142:420-32
7. Biron D, Alvarez-Lacalle E, Tlusty T, Moses E. 2005. Molecular model of the contractile ring. Phys. Rev. Lett. 95:098102
8. Brennan IM, Peters U, Kapoor TM, Straight AF. 2007. Polo-like kinase controls vertebrate spindle elongation and cytokinesis. PLoS One 2:e409
9. Bringmann H,Hyman AA. 2005.Acytokinesis furrow is positioned by twoconsecutive signals. Nature 436:731-34
10. Burkard ME, Maciejowski J, Rodriguez-Bravo V, Repka M, Lowery DM, et al. 2009. Plk1 self-organization and priming phosphorylation ofHsCYK-4 at the spindle midzone regulate the onset of division in human cells. PLoS Biol. 7:e1000111
11. Burkard ME, Randall CL, Larochelle S, Zhang C, Shokat KM, et al. 2007. Chemical genetics reveals the requirement for Polo-like kinase 1 activity in positioning RhoA and triggering cytokinesis in human cells. Proc. Natl. Acad. Sci. USA 104:4383-88
12. Caballe A, Martin-Serrano J. 2011. ESCRT machinery and cytokinesis: the road to daughter cell separation. Traffic 12:1318-26
13. Cabernard C, Prehoda KE, Doe CQ. 2010. A spindle-independent cleavage furrow positioning pathway. Nature 467:91-94
14. Calvert ME, Wright GD, Leong FY, Chiam KH, Chen Y, et al. 2011. Myosin concentration underlies cell size-dependent scalability of actomyosin ring constriction. J. Cell Biol. 195:799-813
15. Canman JC, Cameron LA, Maddox PS, Straight A, Tirnauer JS, et al. 2003. Determining the position of the cell division plane. Nature 424:1074-78
16. Canman JC, Hoffman DB, Salmon ED. 2000. The role of pre-and post-anaphase microtubules in the cytokinesis phase of the cell cycle. Curr. Biol. 10:611-14
17. Canman JC, Lewellyn L, Laband K, Smerdon SJ, Desai A, et al. 2008. Inhibition of Rac by the GAP activity of centralspindlin is essential for cytokinesis. Science 322:1543-46
18. Cao LG, Wang YL. 1990. Mechanism of the formation of contractile ring in dividing cultured animal cells. II. Cortical movement of microinjected actin filaments. J. Cell Biol. 111:1905-11
19. Cao LG, Wang YL. 1996. Signals from the spindle midzone are required for the stimulation of cytokinesis in cultured epithelial cells. Mol. Biol. Cell 7:225-32
20. Carlsson AE. 2006. Contractile stress generation by actomyosin gels. Phys. Rev. E 74:051912
21. Carlton JG, Martin-Serrano J. 2007. Parallels between cytokinesis and retroviral budding: a role for the ESCRT machinery. Science 316:1908-12
22. Carlton JG, Caballe A, Agromayor M, Kloc M, Martin-Serrano J. 2012. ESCRT-III governs the Aurora B-mediated abscission checkpoint through CHMP4C. Science 336:220-25
23. Carmena M, Ruchaud S, Earnshaw WC. 2009. Making the Auroras glow: regulation of Aurora A and B kinase function by interacting proteins. Curr. Opin. Cell Biol. 21:796-805
24. Capalbo L, Montembault E, Takeda T, Bassi ZI, Glover DM, et al. 2012. The chromosomal passenger complex controls the function of endosomal sorting complex required for transport-III Snf7 proteins during cytokinesis. Open Biol. 2:120070
25. Carvalho A, Desai A, Oegema K. 2009. Structural memory in the contractile ring makes the duration of cytokinesis independent of cell size. Cell 137:926-37
26. Castro A, Arlot-Bonnemains Y, Vigneron S, Labbe JC, Prigent C, Lorca T. 2002. APC/Fizzy-Related targets Aurora-A kinase for proteolysis. EMBO Rep. 3:457-62
27. Cesario JM, Jang JK, Redding B, Shah N, Rahman T, McKim KS. 2006. Kinesin 6 family member Subito participates in mitotic spindle assembly and interacts with mitotic regulators. J. Cell Sci. 119:4770-80
28. Chalamalasetty RB, Hummer S, Nigg EA, Sillje HH. 2006. Influence of human Ect2 depletion and overexpression on cleavage furrow formation and abscission. J. Cell Sci. 119:3008-19
29. ChangDC, Xu N, Luo KQ. 2003. Degradation of cyclin B is required for the onset of anaphase in mammalian cells. J. Biol. Chem. 278:37865-73
30. Charras G, Paluch E. 2008. Blebs lead the way: how to migrate without lamellipodia. Nat. Rev. Mol. Cell Biol. 9:730-36
31. Chen Q, PollardTD. 2011. Actin filament severing by cofilin is more important for assembly than constriction of the cytokinetic contractile ring. J. Cell Biol. 195:485-98
32. Chen W, Foss M, Tseng KF, Zhang D. 2008. Redundant mechanisms recruit actin into the contractile ring in silkworm spermatocytes. PLoS Biol. 6:e209
33. Connell JW, Lindon C, Luzio JP, Reid E. 2009. Spastin couples microtubule severing to membrane traffic in completion of cytokinesis and secretion. Traffic 10:42-56
34. D'Avino PP. 2009. How to scaffold the contractile ring for a safe cytokinesis-lessons from Anillin-related proteins. J. Cell Sci. 122:1071-79
35. D'Avino PP, Savoian MS, Glover DM. 2004. Mutations in sticky lead to defective organization of the contractile ring during cytokinesis and are enhanced by Rho and suppressed by Rac. J. Cell Biol. 166:61-71
36. D'Avino PP, Savoian MS, Glover DM. 2005. Cleavage furrow formation and ingression during animal cytokinesis: a microtubule legacy. J. Cell Sci. 118:1549-58
37. D'Avino PP, Takeda T, Capalbo L, Zhang W, Lilley KS, et al. 2008. Interaction between Anillin and Rac-GAP50C connects the actomyosin contractile ring with spindle microtubules at the cell division site. J. Cell Sci. 121:1151-58
38. Dean SO, Rogers SL, Stuurman N, Vale RD, Spudich JA. 2005. Distinct pathways control recruitment and maintenance ofmyosin II at the cleavage furrow during cytokinesis. Proc. Natl. Acad. Sci. USA 102:13473-78
39. DeBiasio RL, LaRocca GM, Post PL, Taylor DL. 1996. Myosin II transport, organization, and phosphorylation: evidence for cortical flow/solation- contraction coupling during cytokinesis and cell locomotion. Mol. Biol. Cell 7:1259-82
40. Dechant R, Glotzer M. 2003. Centrosome separation and central spindle assembly act in redundant pathways that regulate microtubule density and trigger cleavage furrow formation. Dev. Cell 4:333-44
41. Dickinson RB, Caro L, Purich DL. 2004. Force generation by cytoskeletal filament end-tracking proteins. Biophys. J. 87:2838-54
42. Douglas ME, Davies T, Joseph N, Mishima M. 2010. Aurora B and 14-3-3 coordinately regulate clustering of centralspindlin during cytokinesis. Curr. Biol. 20:927-33
43. Echard A, Hickson GR, Foley E, O'Farrell PH. 2004. Terminal cytokinesis events uncovered after an RNAi screen. Curr. Biol. 14:1685-93
44. Echard A, O'Farrell PH. 2003. The degradation of twomitotic cyclins contributes to the timing of cytokinesis. Curr. Biol. 13:373-83
45. Effler JC, Kee YS, Berk JM, Tran MN, Iglesias PA, Robinson DN. 2006. Mitosis-specific mechanosensing and contractile-protein redistribution control cell shape. Curr. Biol. 16:1962-67
46. Eggert US, Mitchison TJ, Field CM. 2006. Animal cytokinesis: from parts list to mechanisms. Annu. Rev. Biochem. 75:543-66
47. Elia N, Sougrat R, Spurlin TA, Hurley JH, Lippincott-Schwartz J. 2011. Dynamics of endosomal sorting complex required for transport (ESCRT) machinery during cytokinesis and its role in abscission. Proc. Natl. Acad. Sci. USA 108:4846-51
48. Estey MP, Di Ciano-Oliveira C, Froese CD, Bejide MT, Trimble WS. 2010. Distinct roles of septins in cytokinesis: SEPT9 mediates midbody abscission. J. Cell Biol. 191:741-49
49. Euteneuer U, McIntosh JR. 1980. Polarity ofmidbody and phragmoplastmicrotubules. J. Cell Biol. 87:509-15
50. FabbroM, Zhou BB, TakahashiM, Sarcevic B, Lal P, et al. 2005. Cdk1/Erk2-and Plk1-dependent phosphorylation of a centrosome protein, Cep55, is required for its recruitment to midbody and cytokinesis. Dev. Cell 9:477-88
51. Fishkind D, Wang Y. 1993. Orientation and three-dimensional organization of actin filaments in dividing cultured cells. J. Cell Biol. 123:837-48
52. Floyd S, Pines J, Lindon C. 2008. APC/C Cdh1 targets aurora kinase to control reorganization of the mitotic spindle at anaphase. Curr. Biol. 18:1649-58
53. Foe VE, Von Dassow G. 2008. Stable and dynamic microtubules coordinately shape the myosin activation zone during cytokinetic furrow formation. J. Cell Biol. 183:457-70
54. Gai M, Camera P, Dema A, Bianchi F, Berto G, et al. 2011. Citron kinase controls abscission through RhoA and anillin. Mol. Biol. Cell 22:3768-78
55. Gaillard J, Neumann E, Van Damme D, Stoppin-Mellet V, Ebel C, et al. 2008. Two microtubule-associated proteins of Arabidopsis MAP65s promote antiparallel microtubule bundling. Mol. Biol. Cell 19:4534-44
56. Gallant P,Nigg EA. 1992. Cyclin B2 undergoes cell cycle-dependent nuclear translocation and,when expressed as a non-destructible mutant, causes mitotic arrest in HeLa cells. J. Cell Biol. 117:213-24
57. Garrus JE, Von Schwedler UK, Pornillos OW, Morham SG, Zavitz KH, et al. 2001. Tsg101 and the vacuolar protein sorting pathway are essential for HIV-1 budding. Cell 107:55-65
58. Giet R, GloverDM.2001. Drosophila aurora B kinase is required for histoneH3 phosphorylation and condensin recruitment during chromosome condensation and to organize the central spindle during cytokinesis. J. Cell Biol. 152:669-82
59. Girard KD, Chaney C, Delannoy M, Kuo SC, Robinson DN. 2004. Dynacortin contributes to cortical viscoelasticity and helps define the shape changes of cytokinesis. EMBO J. 23:1536-46
60. Glotzer M. 2005. The molecular requirements for cytokinesis. Science 307:1735-39
61. Glotzer M. 2009. The 3Ms of central spindle assembly: microtubules, motors and MAPs. Nat. Rev. Mol. Cell Biol. 10:9-20
62. Goldbach P, Wong R, Beise N, Sarpal R, Trimble WS, Brill JA. 2010. Stabilization of the actomyosin ring enables spermatocyte cytokinesis in Drosophila. Mol. Biol. Cell 21:1482-93
63. Goldstein AY, Jan YN, Luo L. 2005. Function and regulation of Tumbleweed (RacGAP50C) in neuroblast proliferation and neuronal morphogenesis. Proc. Natl. Acad. Sci. USA 102:3834-39
64. Goode BL, Eck MJ. 2007. Mechanism and function of formins in the control of actin assembly. Annu. Rev. Biochem. 76:593-627
65. Goss JW, Toomre DK. 2008. Both daughter cells traffic and exocytose membrane at the cleavage furrow during mammalian cytokinesis. J. Cell Biol. 181:1047-54
66. Gregory SL, Ebrahimi S, Milverton J, Jones WM, Bejsovec A, Saint R. 2008. Cell division requires a direct link between microtubule-bound RacGAP and Anillin in the contractile ring. Curr. Biol. 18:25-29
67. Guha M, Zhou M, Wang YL. 2005. Cortical actin turnover during cytokinesis requires myosin II. Curr. Biol. 15:732-36
68. Guizetti J, Gerlich DW. 2012. ESCRT-III polymers in membrane neck constriction. Trends Cell Biol. 22:133-40
69. Guizetti J, Schermelleh L, Mantler J, Maar S, Poser I, et al. 2011. Cortical constriction during abscission involves helices of ESCRT-III-dependent filaments. Science 331:1616-20
70. Guse A, MishimaM,Glotzer M. 2005. Phosphorylation of ZEN-4/MKLP1 by aurora B regulates completion of cytokinesis. Curr. Biol. 15:778-86
71. Hagting A, Den Elzen N, Vodermaier HC, Waizenegger IC, Peters JM, Pines J. 2002. Human securin proteolysis is controlled by the spindle checkpoint and reveals when the APC/C switches from activation by Cdc20 to Cdh1. J. Cell Biol. 157:1125-37
72. Hauf S, Cole RW, LaTerra S, Zimmer C, Schnapp G, et al. 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. 161:281-94
73. Haviv L, Gillo D, Backouche F, Bernheim-Groswasser A. 2008. A cytoskeletal demolition worker: myosin II acts as an actin depolymerization agent. J. Mol. Biol. 375:325-30
74. Henne WM, Buchkovich NJ, Emr SD. 2011. The ESCRT pathway. Dev. Cell 21:77-91
75. Hu CK, Coughlin M, Field CM, Mitchison TJ. 2008. Cell polarization during monopolar cytokinesis. J. Cell Biol. 181:195-202
76. Hu CK, Coughlin M, Field CM, Mitchison TJ. 2011. KIF4 regulates midzone length during cytokinesis. Curr. Biol. 21:815-24
77. Hu CK, Coughlin M, Mitchison TJ. 2012a. Midbody assembly and its regulation during cytokinesis. Mol. Biol. Cell 23:1024-34
78. Hu CK, Ozlu N, Coughlin M, Steen JJ, Mitchison TJ. 2012b. Plk1 negatively regulates PRC1 to prevent premature midzone formation before cytokinesis. Mol. Biol. Cell In press
79. Hummer S, Mayer TU. 2009. Cdk1 negatively regulates midzone localization of the mitotic kinesin Mklp2 and the chromosomal passenger complex. Curr. Biol. 19:607-12
80. Hurley JH, Hanson PI. 2010. Membrane budding and scission by the ESCRT machinery: It's all in the neck. Nat. Rev. Mol. Cell Biol. 11:556-66
81. Hutterer A, Glotzer M, Mishima M. 2009. Clustering of centralspindlin is essential for its accumulation to the central spindle and the midbody. Curr. Biol. 19:2043-49
82. Inoue YH, Savoian MS, Suzuki T, Mathe E, Yamamoto MT, Glover DM. 2004. Mutations in orbit/mast reveal that the central spindle is comprised of two microtubule populations, those that initiate cleavage and those that propagate furrow ingression. J. Cell Biol. 166:49-60
83. Janson LW, Sellers JR, Taylor DL. 1992. Actin-binding proteins regulate the work performed by myosin II motors on single actin filaments. Cell Motil. Cytoskeleton 22:274-80
84. Janson ME, Loughlin R, Loiodice I, Fu C, Brunner D, et al. 2007. Crosslinkers and motors organize dynamic microtubules to form stable bipolar arrays in fission yeast. Cell 128:357-68
85. Jantsch-Plunger V, Gonczy P, Romano A, Schnabel H, Hamill D, et al. 2000. CYK-4: a Rho family GTPase activating protein (GAP) required for central spindle formation and cytokinesis. J. Cell Biol. 149:1391-404
86. JiangW, Jimenez G, Wells NJ, Hope TJ,Wahl GM, et al. 1998. PRC1: a human mitotic spindle-associated CDK substrate protein required for cytokinesis. Mol. Cell 2:877-85
87. Kaitna S, MendozaM, Jantsch-Plunger V, GlotzerM. 2000. Incenp and an Aurora-like kinase form a complex essential for chromosome segregation and efficient completion of cytokinesis. Curr. Biol. 10:1172-81
88. Kamasaki T, Osumi M, Mabuchi I. 2007. Three-dimensional arrangement of F-actin in the contractile ring of fission yeast. J. Cell Biol. 178:765-71
89. Kamijo K, Ohara N, Abe M, Uchimura T, Hosoya H, et al. 2006. Dissecting the role of Rho-mediated signaling in contractile ring formation. Mol. Biol. Cell 17:43-55
90. Kawashima T, Hirose K, Satoh T, Kaneko A, Ikeda Y, et al. 2000. MgcRacGAP is involved in the control of growth and differentiation of hematopoietic cells. Blood 96:2116-24
91. Kechad A, Jananji S, Ruella Y, Hickson GR. 2012. Anillin acts as a bifunctional linker coordinating midbody ring biogenesis during cytokinesis. Curr. Biol. 22:197-203
92. Kondo T, Hamao K, Kamijo K, Kimura, H,MoritaM, et al. 2011. Enhancement of myosin II/actin turnover at the contractile ring induces slower furrowing in dividing HeLa cells. Biochem. J. 435:569-76
93. Kurasawa Y, Earnshaw WC, Mochizuki Y, Dohmae N, Todokoro K. 2004. Essential roles of KIF4 and its binding partner PRC1 in organized central spindle midzone formation. EMBO J. 23:3237-48
94. Lee HH, Elia N, Ghirlando R, Lippincott-Schwartz J, Hurley JH. 2008. Midbody targeting of the ESCRT machinery by a noncanonical coiled coil in CEP55. Science 322:576-80
95. Lewellyn L, Carvalho A, Desai A, Maddox AS, Oegema K. 2011. The chromosomal passenger complex and centralspindlin independently contribute to contractile ring assembly. J. Cell Biol. 193:155-69
96. Lewellyn L,Dumont J,Desai A, Oegema K. 2010. Analyzing the effects of delaying aster separation on furrow formation during cytokinesis in the Caenorhabditis elegans embryo. Mol. Biol. Cell 21:50-62
97. Lindas AC, Karlsson EA, Lindgren MT, Ettema TJ, Bernander R. 2008. A unique cell division machinery in the Archaea. Proc. Natl. Acad. Sci. USA 105:18942-46
98. Lindon C, Pines J. 2004. Ordered proteolysis in anaphase inactivates Plk1 to contribute to proper mitotic exit in human cells. J. Cell Biol. 164:233-41
99. Liot C, Seguin L, Siret A, Crouin C, Schmidt S, Bertoglio J. 2011. APCcdh1 mediates degradation of the oncogenic Rho-GEF Ect2 after mitosis. PLoS One 6:e23676
100. Littlepage LE, Ruderman JV. 2002. Identification of a new APC/C recognition domain, the A box, which is required for the Cdh1-dependent destruction of the kinase Aurora-A during mitotic exit. Genes Dev. 16:2274-85
101. Loiodice I, Staub J, Setty TG, Nguyen NP, Paoletti A, Tran PT. 2005. Ase1p organizes antiparallel microtubule arrays during interphase and mitosis in fission yeast. Mol. Biol. Cell 16:1756-68
102. Loria A, Longhini KM, Glotzer M. 2012. The RhoGAP domain of CYK-4 has an essential role in RhoA activation. Curr. Biol. 22:213-19
103. Lu Q, HopeLW,BraschM,Reinhard C, Cohen SN. 2003. TSG101interactionwithHRSmediates endosomal trafficking and receptor down-regulation. Proc. Natl. Acad. Sci. USA 100:7626-31
104. Ma X, Kovacs M, ContiMA, Wang A, Zhang Y, et al. 2012. Nonmuscle myosin II exerts tension but does not translocate actin in vertebrate cytokinesis. Proc. Natl. Acad. Sci. USA 109:4509-14
105. Mabuchi I. 1994. Cleavage furrow: timing of emergence of contractile ring actin filaments and establishment of the contractile ring by filament bundling in sea urchin eggs. J. Cell Sci. 107(Pt 7):1853-62
106. Mabuchi I, Tsukita S, Tsukita S, Sawai T. 1988. Cleavage furrow isolated from newt eggs: contraction, organization of the actin filaments, and protein components of the furrow. Proc. Natl. Acad. Sci. USA 85:5966-70
107. Madaule P, Eda M, Watanabe N, Fujisawa K, Matsuoka T, et al. 1998. Role of citron kinase as a target of the small GTPase Rho in cytokinesis. Nature 394:491-94
108. Martineau SN, Andreassen PR, Margolis RL. 1995. Delay of HeLa cell cleavage into interphase using dihydrocytochalasin B: retention of a postmitotic spindle and telophase disc correlates with synchronous cleavage recovery. J. Cell Biol. 131:191-205
109. Mastronarde DN, McDonald KL, Ding R,McIntosh JR. 1993. Interpolar spindle microtubules in PTK cells. J. Cell Biol. 123:1475-89
110. Matsumura F, Yamakita Y, Yamashiro S. 2011. Myosin light chain kinases and phosphatase in mitosis and cytokinesis. Arch. Biochem. Biophys. 510:76-82
111. Matzke R, Jacobson KA, Radmacher M. 2001. Direct, high-resolution measurement of furrow stiffening during division of adherent cells. Nat. Cell Biol. 3:607-10
112. Maupin P, Pollard TD. 1986. Arrangement of actin filaments andmyosin-like filaments in the contractile ring and of actin-like filaments in the mitotic spindle of dividing HeLa cells. J. Ultrastruct. Mol. Struct. Res. 94:92-103
113. Miller AL, Bement WM. 2009. Regulation of cytokinesis by Rho GTPase flux. Nat. Cell Biol. 11:71-77
114. Mishima M, Pavicic V, Gruneberg U, Nigg EA, Glotzer M. 2004. Cell cycle regulation of central spindle assembly. Nature 430:908-13
115. Mollinari C, Kleman JP, JiangW, Schoehn G, Hunter T, Margolis RL. 2002. PRC1 is a microtubule binding and bundling protein essential to maintain the mitotic spindle midzone. J. Cell Biol. 157:1175-86
116. Mollinari C, Kleman JP, Saoudi Y, Jablonski SA, Perard J, et al. 2005. Ablation of PRC1 by small interfering RNA demonstrates that cytokinetic abscission requires a central spindle bundle in mammalian cells, whereas completion of furrowing does not. Mol. Biol. Cell 16:1043-55
117. Morita E. 2012. Differential requirements of mammalian ESCRTs in multivesicular body formation, virus budding and cell division. FEBS J. 279:1399-406
118. Morita E, Sandrin V, Chung HY, Morham SG, Gygi SP, et al. 2007. Human ESCRT and ALIX proteins interact with proteins of the midbody and function in cytokinesis. EMBO J. 26:4215-27
119. Motegi F, Sugimoto A. 2006. Sequential functioning of the ECT-2RhoGEF, RHO-1 andCDC-42 establishes cell polarity in Caenorhabditis elegans embryos. Nat. Cell Biol. 8:978-85
120. Mukhina S, Wang YL, Murata-Hori M. 2007. Alpha-actinin is required for tightly regulated remodeling of the actin cortical network during cytokinesis. Dev. Cell 13:554-65
121. Mullins JM, Biesele JJ. 1977. Terminal phase of cytokinesis in D-98s cells. J. Cell Biol. 73:672-84
122. Murata-Hori M, Fumoto K, Fukuta Y, Iwasaki T, Kikuchi A, et al. 2000. Myosin II regulatory light chain as a novel substrate for AIM-1, an Aurora/Ipl1p-related kinase from rat. J. Biochem. 128:903-7
123. Murthy K, Wadsworth P. 2005. Myosin-II-dependent localization and dynamics of F-actin during cytokinesis. Curr. Biol. 15:724-31
124. Neef R, Gruneberg U, Kopajtich R, Li X, Nigg EA, et al. 2007. Choice of Plk1 docking partners during mitosis and cytokinesis is controlled by the activation state of Cdk1. Nat. Cell Biol. 9:436-44
125. Neef R, Preisinger C, Sutcliffe J, Kopajtich R, Nigg EA, et al. 2003. Phosphorylation of mitotic kinesin-like protein 2 by polo-like kinase 1 is required for cytokinesis. J. Cell Biol. 162:863-75
126. Neto H, Collins LL, GouldGW. 2011. Vesicle trafficking andmembrane remodelling in cytokinesis. Biochem. J. 437:13-24
127. Nguyen HG, Chinnappan D, Urano T, Ravid K. 2005. Mechanism of Aurora-B degradation and its dependency on intact KEN and A-boxes: identification of an aneuploidy-promoting property. Mol. Cell. Biol. 25:4977-92
128. Niiya F, Tatsumoto T, Lee KS, Miki T. 2006. Phosphorylation of the cytokinesis regulator ECT2 at at G2/M phase stimulates association of the mitotic kinase Plk1 and accumulation of GTP-bound RhoA. Oncogene 25:827-37
129. Niiya F, Xie X, Lee KS, Inoue H, Miki T. 2005. Inhibition of cyclin-dependent kinase 1 induces cytokinesis without chromosome segregation in an ECT2 and MgcRacGAP-dependent manner. J. Biol. Chem. 280:36502-9
130. Nishimura Y, Yonemura S. 2006. Centralspindlin regulates ECT2 and RhoA accumulation at the equatorial cortex during cytokinesis. J. Cell Sci. 119:104-14
131. Norden C, Mendoza M, Dobbelaere J, Kotwaliwale CV, Biggins S, et al. 2006. The NoCut pathway links completion of cytokinesis to spindle midzone function to prevent chromosome breakage. Cell 125:85-98
132. Oegema K, Mitchison TJ. 1997. Rappaport rules: cleavage furrow induction in animal cells. Proc. Natl. Acad. Sci. USA 94:4817-20
133. Ojkic N, Vavylonis D. 2010. Kinetics of myosin node aggregation into a contractile ring. Phys. Rev. Lett. 105:048102
134. Ou G, Stuurman N, D'Ambrosio M, Vale RD. 2010. Polarized myosin produces unequal-size daughters during asymmetric cell division. Science 330:677-80
135. Pavicic-Kaltenbrunner V, Mishima M, Glotzer M. 2007. Cooperative assembly of CYK-4/MgcRacGAP and ZEN-4/MKLP1 to form the centralspindlin complex. Mol. Biol. Cell 18:4992-5003
136. Pelham RJ, Chang F. 2002. Actin dynamics in the contractile ring during cytokinesis in fission yeast. Nature 419:82-86
137. PetronczkiM,Glotzer M, KrautN, Peters JM. 2007. Polo-like kinase 1 triggers the initiation of cytokinesis in human cells by promoting recruitment of the RhoGEF Ect2 to the central spindle. Dev. Cell 12:713-25
138. PetronczkiM, Lenart P, Peters J-M. 2008. Polo on the rise-from mitotic entry to cytokinesis with Plk1. Dev. Cell 14:646-59
139. Piekny A, Werner M, Glotzer M. 2005. Cytokinesis: Welcome to the Rho zone. Trends Cell Biol. 15:651-58
140. Piekny AJ, GlotzerM. 2008. Anillin is a scaffold protein that links RhoA, actin, and myosin during cytokinesis. Curr. Biol. 18:30-36
141. Piekny AJ, Maddox AS. 2010. The myriad roles of Anillin during cytokinesis. Semin. Cell Dev. Biol. 21:881-91
142. Pines J. 2011. Cubism and the cell cycle: the many faces of the APC/C. Nat. Rev. Mol. Cell Biol. 12:427-38
143. Pollard TD. 2010. Mechanics of cytokinesis in eukaryotes. Curr. Opin. Cell Biol. 22:50-56
144. Pollard TD, Wu JQ. 2010. Understanding cytokinesis: lessons from fission yeast. Nat. Rev. Mol. Cell Biol. 11:149-55
145. Potapova TA, Daum JR, Pittman BD, Hudson JR, Jones TN, et al. 2006. The reversibility of mitotic exit in vertebrate cells. Nature 440:954-58
146. Powers J, Bossinger O, Rose D, Strome S, Saxton W. 1998. A nematode kinesin required for cleavage furrow advancement. Curr. Biol. 8:1133-36
147. Prekeris R, GouldGW. 2008. Breaking up is hard to do-membrane traffic in cytokinesis. J. Cell Sci. 121:1569-76
148. Prokopenko SN, Brumby A, O'Keefe L, Prior L, He Y, et al. 1999. A putative exchange factor for Rho1 GTPase is required for initiation of cytokinesis in Drosophila. Genes Dev. 13:2301-14
149. Raich WB, Moran AN, Rothman JH, Hardin J. 1998. Cytokinesis and midzone microtubule organization in Caenorhabditis elegans require the kinesin-like protein ZEN-4. Mol. Biol. Cell 9:2037-49
150. Rappaport R. 1985. Repeated furrow formation from a single mitotic apparatus in cylindrical sand dollar eggs. J. Exp. Zool. 234:167-71
151. Rappaport R. 1996. Cytokinesis in Animal Cells. Cambridge, UK
152. Reichl EM, Ren Y,Morphew MK, DelannoyM, Effler JC, et al. 2008. Interactions between myosin and actin crosslinkers control cytokinesis contractility dynamics and mechanics. Curr. Biol. 18:471-80
153. Reid E, Connell J, Edwards TL, Duley S, Brown SE, Sanderson CM. 2005. The hereditary spastic paraplegia protein spastin interacts with the ESCRT-III complex-associated endosomal protein CHMP1B. Hum. Mol. Genet. 14:19-38
154. Robinson DN,Cavet G, WarrickHM,Spudich JA. 2002. Quantitation of the distribution and flux ofmyosin-II during cytokinesis. BMC Cell Biol. 3:4
155. Rossman KL, Der CJ, Sondek J. 2005. GEF means go: turning on RHO GTPases with guanine nucleotideexchange factors. Nat. Rev. Mol. Cell Biol. 6:167-80
156. Ruchaud S, Carmena M, Earnshaw WC. 2007. Chromosomal passengers: conducting cell division. Nat. Rev. Mol. Cell Biol. 8:798-812
157. Salbreux G, Prost J, Joanny JF. 2009. Hydrodynamics of cellular cortical flows and the formation of contractile rings. Phys. Rev. Lett. 103:058102
158. Salmon ED, Wolniak SM. 1990. Role of microtubules in stimulating cytokinesis in animal cells. Ann. N.Y. Acad. Sci. 582:88-98
159. Samson RY, Obita T, Freund SM, Williams RL, Bell SD. 2008. A role for the ESCRT system in cell division in archaea. Science 322:1710-13
160. Sanger JM, Sanger JW. 1980. Banding and polarity of actin filaments in interphase and cleaving cells. J. Cell Biol. 86:568-75
161. Schiel JA, Prekeris R. 2010. Making the final cut-mechanisms mediating the abscission step of cytokinesis. Sci. World J. 10:1424-34
162. Schiel JA, Prekeris R. 2011. ESCRT or endosomes? Tales of the separation of two daughter cells. Commun. Integr. Biol. 4:606-8
163. Schroeder TE. 1972. The contractile ring. II. Determining its brief existence, volumetric changes, and vital role in cleaving Arbacia eggs. J. Cell Biol. 53:419-34
164. Schroeder TE. 1990. The contractile ring and furrowing in dividing cells. Ann. N.Y. Acad. Sci. 582:78-87
165. Schumacher JM, Golden A, Donovan PJ. 1998. AIR-2: An Aurora/Ipl1-related protein kinase associated with chromosomes and midbody microtubules is required for polar body extrusion and cytokinesis in Caenorhabditis elegans embryos. J. Cell Biol. 143:1635-46
166. Sedzinski J, Biro M, Oswald A, Tinevez J-Y, Salbreux G, Paluch E. 2011. Polar acto-myosin contractility destabilises the position of the cytokinetic furrow. Nature 476:462-66
167. Severson AF, Baillie DL, Bowerman B. 2002. A Formin Homology protein and a profilin are required for cytokinesis and Arp2/3-independent assembly of cortical microfilaments in C. elegans. Curr. Biol. 12:2066-75
168. Severson AF, Hamill DR, Carter JC, Schumacher J, Bowerman B. 2000. The Aurora-related kinase AIR-2 recruits ZEN-4/CeMKLP1 to themitotic spindle atmetaphase and is required for cytokinesis. Curr. Biol. 10:1162-71
169. Shelton CA, Carter JC, Ellis GC, Bowerman B. 1999. The nonmuscle myosin regulatory light chain gene mlc-4 is required for cytokinesis, anterior-posterior polarity, and body morphology during Caenorhabditis elegans embryogenesis. J. Cell Biol. 146:439-51
170. Smith TC, Fang Z, Luna EJ. 2010. Novel interactors and a role for supervillin in early cytokinesis. Cytoskeleton 67:346-64
171. Somers WG, Saint R. 2003. A RhoGEF and Rho family GTPase-activating protein complex links the contractile ring to cortical microtubules at the onset of cytokinesis. Dev. Cell 4:29-39
172. Somma MP, Fasulo B, Cenci G, Cundari E, Gatti M. 2002. Molecular dissection of cytokinesis by RNA interference in Drosophila cultured cells. Mol. Biol. Cell 13:2448-60
173. Speliotes EK, Uren A, Vaux D, Horvitz HR. 2000. The survivin-like C. elegans BIR-1 protein acts with the Aurora-like kinase AIR-2 to affect chromosomes and the spindle midzone. Mol. Cell 6:211-23
174. Steigemann P, Wurzenberger C, SchmitzMH, Held M, Guizetti J, et al. 2009. Aurora B-mediated abscission checkpoint protects against tetraploidization. Cell 136:473-84
175. Stewart S, Fang G. 2005. Destruction box-dependent degradation of Aurora B is mediated by the anaphasepromoting complex/cyclosome and Cdh1. Cancer Res. 65:8730-35
176. Straight AF, Cheung A, Limouze J, Chen I,Westwood NJ, et al. 2003. Dissecting temporal and spatial control of cytokinesis with a myosin II inhibitor. Science 299:1743-47
177. Straight AF, Field CM, Mitchison TJ. 2005. Anillin binds nonmuscle myosin II and regulates the contractile ring. Mol. Biol. Cell 16:193-201
178. Su KC, Takaki T, Petronczki M. 2011. Targeting of the RhoGEF Ect2 to the equatorial membrane controls cleavage furrow formation during cytokinesis. Dev. Cell 21:1104-15
179. Surcel A, Kee YS, Luo T, Robinson DN. 2010. Cytokinesis through biochemical-mechanical feedback loops. Semin. Cell Dev. Biol. 21:866-73
180. Szafer-Glusman E, FullerMT, GiansantiMG. 2011. Role of Survivin in cytokinesis revealed by a separationof-function allele. Mol. Biol. Cell 22:3779-90
181. Taguchi S, Honda K, Sugiura K, Yamaguchi A, Furukawa K, UranoT. 2002. Degradation of human Aurora-A protein kinase is mediated by hCdh1. FEBS Lett. 519:59-65
182. Tcherkezian J, Lamarche-Vane N. 2007. Current knowledge of the large RhoGAP family of proteins. Biol. Cell 99:67-86
183. Toure A, Dorseuil O, Morin L, Timmons P, Jegou B, et al. 1998. MgcRacGAP, a new human GTPaseactivating protein for Rac and Cdc42 similar to Drosophila rotundRacGAP gene product, is expressed in male germ cells. J. Biol. Chem. 273:6019-23
184. Toure A, Mzali R, Liot C, Seguin L, Morin L, et al. 2008. Phosphoregulation of MgcRacGAP in mitosis involves Aurora B and Cdk1 protein kinases and the PP2A phosphatase. FEBS Lett. 582:1182-88
185. Uehara R, Goshima G, Mabuchi I, Vale RD, Spudich JA, et al. 2010. Determinants of myosin II cortical localization during cytokinesis. Curr. Biol. 20:1080-85
186. Vale RD, Spudich JA, Griffis ER. 2009. Dynamics of myosin,microtubules, and Kinesin-6 at the cortex during cytokinesis in Drosophila S2 cells. J. Cell Biol. 186:727-38
187. Vavylonis D, Wu JQ, Hao S, O'Shaughnessy B, Pollard TD. 2008. Assembly mechanism of the contractile ring for cytokinesis by fission yeast. Science 319:97-100
188. Verbrugghe KJ, White JG. 2004. SPD-1 is required for the formation of the spindle midzone but is not essential for the completion of cytokinesis in C. elegans embryos. Curr. Biol. 14:1755-60
189. Verbrugghe KJ, White JG. 2007. Cortical centralspindlin and G alpha have parallel roles in furrow initiation in early C. elegans embryos. J. Cell Sci. 120:1772-78
190. Von Dassow G. 2009. Concurrent cues for cytokinetic furrow induction in animal cells. Trends Cell Biol. 19:165-73
191. Von Dassow G, Verbrugghe KJ, Miller AL, Sider JR, Bement WM. 2009. Action at a distance during cytokinesis. J. Cell Biol. 187:831-45
192. Werner M, Munro E, Glotzer M. 2007. Astral signals spatially bias cortical myosin recruitment to break symmetry and promote cytokinesis. Curr. Biol. 17:1286-97
193. Wheatley SP,Hinchcliffe EH,Glotzer M, Hyman AA, SluderG,Wang Y. 1997. CDK1 inactivation regulates anaphase spindle dynamics and cytokinesis in vivo. J. Cell Biol. 138:385-93
194. Wilson CA, Tsuchida MA, Allen GM, Barnhart EL, Applegate KT, et al. 2010. Myosin II contributes to cell-scale actin network treadmilling through network disassembly. Nature 465:373-77
195. Wolf F, Sigl R, Geley S. 2007. '. . .The end of the beginning': Cdk1 thresholds and exit from mitosis. Cell Cycle 6:1408-11
196. Wolf F, Wandke C, Isenberg N, Geley S. 2006. Dose-dependent effects of stable cyclin B1 on progression through mitosis in human cells. EMBO J. 25:2802-13
197. Wolfe BA, Takaki T, Petronczki M, Glotzer M. 2009. Polo-like kinase 1 directs assembly of the HsCyk-4 RhoGAP/Ect2 RhoGEF complex to initiate cleavage furrow formation. PLoS Biol. 7:e1000110
198. Wu JQ, Pollard TD. 2005. Counting cytokinesis proteins globally and locally in fission yeast. Science 310:310-14
199. Yamada T, Hikida M, Kurosaki T. 2006. Regulation of cytokinesis by mgcRacGAP in B lymphocytes is independent of GAP activity. Exp. Cell Res. 312:3517-25
200. Yang D, Rismanchi N, Renvoise B, Lippincott-Schwartz J, Blackstone C, Hurley JH. 2008. Structural basis for midbody targeting of spastin by the ESCRT-III protein CHMP1B. Nat. Struct. Mol. Biol. 15:1278-86
201. Yoneda M, Dan K. 1972. Tension at the surface of the dividing sea-urchin egg. J. Exp. Biol. 57:575-87
202. Yuce O, Piekny A, GlotzerM. 2005. An ECT2-centralspindlin complex regulates the localization and function of RhoA. J. Cell Biol. 170:571-82
203. Zavortink M, Contreras N, Addy T, Bejsovec A, Saint R. 2005. Tum/RacGAP50C provides a critical link between anaphase microtubules and the assembly of the contractile ring in Drosophila melanogaster. J. Cell Sci. 118:5381-92
204. ZhangW, Robinson DN. 2005. Balance of actively generated contractile and resistive forces controls cytokinesis dynamics. Proc. Natl. Acad. Sci. USA 102:7186-91
205. Zhao WM, Fang G. 2005a. Anillin is a substrate of anaphase-promoting complex/cyclosome (APC/C) that controls spatial contractility of myosin during late cytokinesis. J. Biol. Chem. 280:33516-24
206. Zhao WM, Fang G. 2005b. MgcRacGAP controls the assembly of the contractile ring and the initiation of cytokinesis. Proc. Natl. Acad. Sci. USA 102:13158-63
207. Zhao WM, Seki A, Fang G. 2006. Cep55, a microtubule-bundling protein, associates with centralspindlin to control the midbody integrity and cell abscission during cytokinesis. Mol. Biol. Cell 17:3881-96
208. Zhou M, Wang YL. 2008. Distinct pathways for the early recruitment of myosin II and actin to the cytokinetic furrow. Mol. Biol. Cell 19:318-26
209. Zhou Q, Kee YS, Poirier CC, Jelinek C, Osborne J, et al. 2010. 14-3-3 coordinates microtubules, Rac, and myosin II to control cell mechanics and cytokinesis. Curr. Biol. 20:1881-89
210. ZhuC,Bossy-WetzelE, JiangW. 2005. Recruitment ofMKLP1to the spindle midzone/midbody byINCENP is essential for midbody formation and completion of cytokinesis in human cells. Biochem. J. 389:373-81
211. Zhu C, Jiang W. 2005. Cell cycle-dependent translocation of PRC1 on the spindle by Kif4 is essential for midzone formation and cytokinesis. Proc. Natl. Acad. Sci. USA 102:343-48
212. Zhu C, Lau E, Schwarzenbacher R, Bossy-Wetzel E, Jiang W. 2006. Spatiotemporal control of spindle midzone formation by PRC1 in human cells. Proc. Natl. Acad. Sci. USA 103:6196-201
213. Zumdieck A, Cosentino-Lagomarsino M, Tanase C, Kruse K, Mulder B, et al. 2005. Continuum description of the cytoskeleton: ring formation in the cell cortex. Phys. Rev. Lett. 95:258103
214. Zumdieck A, Kruse K, Bringmann H, Hyman AA, Julicher F. 2007. Stress generation and filament turnover during actin ring constriction. PLoS One 2:e696