Kinetochore-spindle microtubule interactions during mitosis

Current Opinion in Cell Biology

Volumn 17, Issue 1, 2005, Pages 35-46

  Kline Smith, Susan L ^a   Sandall, Sharsti ^a   Desai, Arshad ^a  

^a UNIVERSITY OF CALIFORNIA   (United States)

Author keywords

[No Author keywords available]

Indexed keywords

CENTROMERE PROTEIN A; DNA; HISTONE; ISOPROTEIN; NUCLEAR PROTEIN; UNCLASSIFIED DRUG;

CENTROMERE; CHROMOSOME SEGREGATION; DEPOLYMERIZATION; DNA SEQUENCE; GENETIC ANALYSIS; HUMAN; MICROTUBULE; MITOSIS; MOLECULAR INTERACTION; NONHUMAN; PRIORITY JOURNAL; PROTEIN ASSEMBLY; PROTEOMICS; REVIEW; SACCHAROMYCES CEREVISIAE; SISTER CHROMATID; YEAST;

ANIMALS; CHROMATIDS; DNA; HISTONES; HUMANS; KINETOCHORES; MICROTUBULES; MITOSIS; MITOTIC SPINDLE APPARATUS; MODELS, BIOLOGICAL; NUCLEAR PROTEINS; POLYMERS; SACCHAROMYCES CEREVISIAE; SACCHAROMYCES CEREVISIAE PROTEINS; SISTER CHROMATID EXCHANGE;

SACCHAROMYCES; SACCHAROMYCES CEREVISIAE;

EID: 12344289259     PISSN: 09550674     EISSN: None     Source Type: Journal    
DOI: 10.1016/j.ceb.2004.12.009     Document Type: Review

References (110)

1. J.R. McIntosh, E.L. Grishchuk, and R.R. West Chromosome-microtubule interactions during mitosis Annu Rev Cell Dev Biol 18 2002 193 219
2. D.W. Cleveland, Y. Mao, and K.F. Sullivan Centromeres and kinetochores: from epigenetics to mitotic checkpoint signaling Cell 112 2003 407 421
3. D.J. Lew, and D.J. Burke The spindle assembly and spindle position checkpoints Annu Rev Genet 37 2003 251 282
4. D.J. Amor, P. Kalitsis, H. Sumer, and K.H. Choo Building the centromere: from foundation proteins to 3D organization Trends Cell Biol 14 2004 359 368
5. A.D. McAinsh, J.D. Tytell, and P.K. Sorger Structure, function, and regulation of budding yeast kinetochores Annu Rev Cell Dev Biol 19 2003 519 539
6. S. Biggins, and C.E. Walczak Captivating capture: how microtubules attach to kinetochores Curr Biol 13 2003 R449 R460
7. T. Fukagawa Centromere DNA, proteins and kinetochore assembly in vertebrate cells Chromosome Res 12 2004 557 567
8. Y. Chen, D.J. Riley, P.L. Chen, and W.H. Lee Hec, a novel nuclear protein rich in leucine heptad repeats specifically involved in mitosis Mol Cell Biol 17 1997 6049 6056
9. P.A. Wigge, O.N. Jensen, S. Holmes, S. Soues, M. Mann, and J.V. Kilmartin Analysis of the Saccharomyces spindle pole by matrix-assisted laser desorption/ionization (MALDI) mass spectrometry J Cell Biol 141 1998 967 977
10. L. Zheng, Y. Chen, and W.H. Lee Hec1p, an evolutionarily conserved coiled-coil protein, modulates chromosome segregation through interaction with SMC proteins Mol Cell Biol 19 1999 5417 5428
11. M. Howe, K.L. McDonald, D.G. Albertson, and B.J. Meyer HIM-10 is required for kinetochore structure and function on Caenorhabditis elegans holocentric chromosomes J Cell Biol 153 2001 1227 1238
12. C. Janke, J. Ortiz, J. Lechner, A. Shevchenko, M.M. Magiera, C. Schramm, and E. Schiebel The budding yeast proteins Spc24p and Spc25p interact with Ndc80p and Nuf2p at the kinetochore and are important for kinetochore clustering and checkpoint control EMBO J 20 2001 777 791
13. A. Nabetani, T. Koujin, C. Tsutsumi, T. Haraguchi, and Y. Hiraoka A conserved protein, Nuf2, is implicated in connecting the centromere to the spindle during chromosome segregation: a link between the kinetochore function and the spindle checkpoint Chromosoma 110 2001 322 334
14. P.A. Wigge, and J.V. Kilmartin The Ndc80p complex from Saccharomyces cerevisiae contains conserved centromere components and has a function in chromosome segregation J Cell Biol 152 2001 349 360
15. J.G. DeLuca, B. Moree, J.M. Hickey, J.V. Kilmartin, and E.D. Salmon hNuf2 inhibition blocks stable kinetochore-microtubule attachment and induces mitotic cell death in HeLa cells J Cell Biol 159 2002 549 555
16. I. Le Masson, C. Saveanu, A. Chevalier, A. Namane, R. Gobin, M. Fromont-Racine, A. Jacquier, and C. Mann Spc24 interacts with Mps2 and is required for chromosome segregation, but is not implicated in spindle pole body duplication Mol Microbiol 43 2002 1431 1443
17. H. Appelgren, B. Kniola, and K. Ekwall Distinct centromere domain structures with separate functions demonstrated in live fission yeast cells J Cell Sci 116 2003 4035 4042
18. ••].
19. T. Hori, T. Haraguchi, Y. Hiraoka, H. Kimura, and T. Fukagawa Dynamic behavior of Nuf2-Hec1 complex that localizes to the centrosome and centromere and is essential for mitotic progression in vertebrate cells J Cell Sci 116 2003 3347 3362
20. ••].
21. •].
22. •].
23. ••].
24. C.L. Rieder The formation, structure, and composition of the mammalian kinetochore and kinetochore fibre Int Rev Cytol 79 1982 1 58
25. DeLuca JG, Dong Y, Hergert P, Strauss J, Hickey JM, Salmon ED, McEwen BF: Hec1 and Nuf2 are core components of the kinetochore outer plate essential for organizing microtubule attachment sites. Mol Biol Cell 2004, in press. The authors probe the ultrastructural consequences of Nuf2 siRNA in HeLa cells to reveal a severely disrupted outer kinetochore with little microtubule binding. In addition, they confirm localization of Nuf2 at the outer kinetochore plate, but not the corona, by immuno-electron microscopy. On the basis of these findings and others, a model is put forth involving the Ndc80 complex in the formation of structurally stable binding sites for the plus ends of microtubules. Upon depletion of Ndc80 complex components from kinetochores, other microtubule binding factors, such as MAPs and motors, may facilitate a few lateral or end-on kinetochore-microtubule attachments, although these are insufficient to support poleward forces at kinetochores.
26. B.F. McEwen, A.B. Heagle, G.O. Cassels, K.F. Buttle, and C.L. Rieder Kinetochore fiber maturation in PtK1 cells and its implications for the mechanisms of chromosome congression and anaphase onset J Cell Biol 137 1997 1567 1580
27. S. Martin-Lluesma, V.M. Stucke, and E.A. Nigg Role of Hec1 in spindle checkpoint signaling and kinetochore recruitment of Mad1/Mad2 Science 297 2002 2267 2270
28. X. He, D.R. Rines, C.W. Espelin, and P.K. Sorger Molecular analysis of kinetochore-microtubule attachment in budding yeast Cell 106 2001 195 206
29. •].
30. •].
31. •].
32. E.S. Gillett, C.W. Espelin, and P.K. Sorger Spindle checkpoint proteins and chromosome-microtubule attachment in budding yeast J Cell Biol 164 2004 535 546
33. C. Janke, J. Ortiz, T.U. Tanaka, J. Lechner, and E. Schiebel Four new subunits of the Dam1-Duo1 complex reveal novel functions in sister kinetochore biorientation EMBO J 21 2002 181 193
34. Y. Li, J. Bachant, A.A. Alcasabas, Y. Wang, J. Qin, and S.J. Elledge The mitotic spindle is required for loading of the DASH complex onto the kinetochore Genes Dev 16 2002 183 197
35. •].
36. ••], the authors state that nocodazole treatment does not restore Mad 1/2 localization to kinetochores upon complete disruption of the Ndc80 complex.
37. K. Oegema, A. Desai, S. Rybina, M. Kirkham, and A.A. Hyman Functional analysis of kinetochore assembly in Caenorhabditis elegans J Cell Biol 153 2001 1209 1226
38. C.L. Rieder, and E.D. Salmon The vertebrate cell kinetochore and its roles during mitosis Trends Cell Biol 8 1998 310 318
39. L.A. Ligon, S.S. Shelly, M. Tokito, and E.L. Holzbaur The microtubule plus-end proteins EB1 and dynactin have differential effects on microtubule polymerization Mol Biol Cell 14 2003 1405 1417
40. N.E. Faulkner, D.L. Dujardin, C.Y. Tai, K.T. Vaughan, C.B. O'Connell, Y. Wang, and R.B. Vallee A role for the lissencephaly gene LIS1 in mitosis and cytoplasmic dynein function Nat Cell Biol 2 2000 784 791
41. K.B. Kaplan, A.A. Burds, J.R. Swedlow, S.S. Bekir, P.K. Sorger, and I.S. Nathke A role for the adenomatous polyposis coli protein in chromosome segregation Nat Cell Biol 3 2001 429 432
42. M. Nakamura, X.Z. Zhou, and K.P. Lu Critical role for the EB1 and APC interaction in the regulation of microtubule polymerization Curr Biol 11 2001 1062 1067
43. S.L. Rogers, G.C. Rogers, D.J. Sharp, and R.D. Vale Drosophila EB1 is important for proper assembly, dynamics, and positioning of the mitotic spindle J Cell Biol 158 2002 873 884
44. C.Y. Tai, D.L. Dujardin, N.E. Faulkner, and R.B. Vallee Role of dynein, dynactin, and CLIP-170 interactions in LIS1 kinetochore function J Cell Biol 156 2002 959 968
45. F.M. Coquelle, M. Caspi, F.P. Cordelieres, J.P. Dompierre, D.L. Dujardin, C. Koifman, P. Martin, C.C. Hoogenraad, A. Akhmanova, and N. Galjart LIS1, CLIP-170's key to the dynein/dynactin pathway Mol Cell Biol 22 2002 3089 3102
46. R.A. Green, and K.B. Kaplan Chromosome instability in colorectal tumor cells is associated with defects in microtubule plus-end attachments caused by a dominant mutation in APC J Cell Biol 163 2003 949 961
47. J.S. Tirnauer, J.C. Canman, E.D. Salmon, and T.J. Mitchison EB1 targets to kinetochores with attached, polymerizing microtubules Mol Biol Cell 13 2002 4308 4316
48. J.S. Tirnauer, S. Grego, E.D. Salmon, and T.J. Mitchison EB1-microtubule interactions in Xenopus egg extracts: role of EB1 in microtubule stabilization and mechanisms of targeting to microtubules Mol Biol Cell 13 2002 3614 3626
49. A. Akhmanova, C.C. Hoogenraad, K. Drabek, T. Stepanova, B. Dortland, T. Verkerk, W. Vermeulen, B.M. Burgering, C.I. De Zeeuw, and F. Grosveld Clasps are CLIP-115 and -170 associating proteins involved in the regional regulation of microtubule dynamics in motile fibroblasts Cell 104 2001 923 935
50. H. Maiato, E.A. Fairley, C.L. Rieder, J.R. Swedlow, C.E. Sunkel, and W.C. Earnshaw Human CLASP1 is an outer kinetochore component that regulates spindle microtubule dynamics Cell 113 2003 891 904 The authors report localization and functional studies of human CLASP1, which has a more peripheral localization (i.e. is closer to microtubules) than previously studied kinetochore components. CLASP1 inhibition leads to monopolar spindles with centrally localized chromosomes, which exhibit reduced oscillations. Measurements of kinetochore fiber lengths in live cells reveal an overall decrease in kinetochore- microtubule dynamics. The authors suggest that CLASP1 is required to promote polymerization of microtubules at kinetochores.
51. Y.H. Inoue, M. do Carmo Avides, M. Shiraki, P. Deak, M. Yamaguchi, Y. Nishimoto, A. Matsukage, and D.M. Glover Orbit, a novel microtubule-associated protein essential for mitosis in Drosophila melanogaster J Cell Biol 149 2000 153 166
52. H. Maiato, P. Sampaio, C.L. Lemos, J. Findlay, M. Carmena, W.C. Earnshaw, and C.E. Sunkel MAST/Orbit has a role in microtubule-kinetochore attachment and is essential for chromosome alignment and maintenance of spindle bipolarity J Cell Biol 157 2002 749 760
53. P. Maddox, A. Straight, P. Coughlin, T.J. Mitchison, and E.D. Salmon Direct observation of microtubule dynamics at kinetochores in Xenopus extract spindles: implications for spindle mechanics J Cell Biol 162 2003 377 382 Fluorescent speckle microscopy of fluorescently labeled tubulin in spindles assembled in Xenopus egg extracts enables the authors to analyze in detail the dynamics of kinetochore fibers versus non-kinetochore microtubules during metaphase and anaphase. Although anaphase is dominated by depolymerization of microtubules at the pole, kinetochore-localized depolymerization occasionally occurs, and suggests that tension across the centromere governs kinetochore-mediated alterations in microtubule dynamics.
54. ••].
55. ••].
56. C.J. Lawrence, R.K. Dawe, K.R. Christie, D.W. Cleveland, S.C. Dawson, S.A. Endow, L.S. Goldstein, H.V. Goodson, N. Hirokawa, and J. Howard A standardized kinesin nomenclature J Cell Biol 167 2004 19 22
57. A. Desai, S. Verma, T.J. Mitchison, and C.E. Walczak Kin I kinesins are microtubule-destabilizing enzymes Cell 96 1999 69 78
58. K.M. Hertzer, S.C. Ems-McClung, and C.E. Walczak Kin I kinesins: insights into the mechanism of depolymerization Crit Rev Biochem Mol Biol 38 2003 453 469
59. Y. Ovechkina, and L. Wordeman Unconventional motoring: an overview of the Kin C and Kin I kinesins Traffic 4 2003 367 375
60. A.W. Hunter, M. Caplow, D.L. Coy, W.O. Hancock, S. Diez, L. Wordeman, and J. Howard The kinesin-related protein MCAK is a microtubule depolymerase that forms an ATP-hydrolyzing complex at microtubule ends Mol Cell 11 2003 445 457 Using stabilized microtubules polymerized with GMPCPP (a non-hydrolysable GTP analog), the authors study the mechanism by which MCAK, a member of the kinesin-13 family, depolymerizes microtubules in vitro. Unlike motile kinesins, MCAK exhibits end-stimulated and tubulin-dimer-stimulated activity. Quantitative analysis of ATPase activity suggests a single binding site for MCAK at the end of each microtubule protofilament. In addition, the authors suggest that one-dimensional diffusion of MCAK along the microtubule lattice facilitates rapid targeting of MCAK to microtubule ends. Once it is at the end, the authors propose that MCAK acts processively to release multiple tubulin subunits per ATP hydrolysis event.
61. T. Maney, A.W. Hunter, M. Wagenbach, and L. Wordeman Mitotic centromere-associated kinesin is important for anaphase chromosome segregation J Cell Biol 142 1998 787 801
62. S.L. Kline-Smith, A. Khodjakov, P. Hergert, and C.E. Walczak Depletion of centromeric MCAK leads to chromosome congression and segregation defects due to improper kinetochore attachments Mol Biol Cell 15 2004 1146 1159 By disrupting centromeric MCAK with a dominant-negative GFP fusion, the authors observe the consequences on chromosome positioning without perturbing overall spindle morphology. Despite normal rates of chromosome and chromatid movement, chromosome congression and segregation are abnormal; in fact, some chromosomes remain at spindle poles or at the spindle equator even after anaphase has taken place. This is due to the presence of improperly positioned microtubule attachments at kinetochores, which inhibit chromosome movements. These findings imply that MCAK functions to depolymerize improperly attached microtubules at the centromere.
63. C.E. Walczak, E.C. Gan, A. Desai, T.J. Mitchison, and S.L. Kline-Smith The microtubule-destabilizing kinesin XKCM1 is required for chromosome positioning during spindle assembly Curr Biol 12 2002 1885 1889
64. G.C. Rogers, S.L. Rogers, T.A. Schwimmer, S.C. Ems-McClung, C.E. Walczak, R.D. Vale, J.M. Scholey, and D.J. Sharp Two mitotic kinesins cooperate to drive sister chromatid separation during anaphase Nature 427 2004 364 370 This study explores the activity of the Drosophila kinesin-13 proteins Klp59C and Klp10A, which are shown to depolymerize microtubules in vitro. Antibody-mediated inhibition of KLP59C leads to chromosome congression defects and slowed chromosome segregation. Fluorescent speckle microscopy suggests that this slowed poleward chromatid rate results from a loss of kinetochore-mediated depolymerization of microtubules, causing the chromatids to be moved solely by the depolymerization of kinetochore microtubules at spindle poles, although the presence of improper kinetochore-microtubule attachments cannot be ruled out. The authors also suggest on the basis of similar experiments that KLP10A depolymerizes microtubules at spindle poles during anaphase.
65. G. Goshima, and R.D. Vale The roles of microtubule-based motor proteins in mitosis: comprehensive RNAi analysis in the Drosophila S2 cell line J Cell Biol 162 2003 1003 1016
66. C.G. Pearson, P.S. Maddox, T.R. Zarzar, E.D. Salmon, and K. Bloom Yeast kinetochores do not stabilize Stu2p-dependent spindle microtubule dynamics Mol Biol Cell 14 2003 4181 4195 The authors explore the relationship between kinetochores and spindle microtubule dynamics using stu2 mutants in budding yeast. The dramatic decrease in spindle (kinetochore and non-kinetochore) microtubule turnover in stu2 mutants is not affected by preventing kinetochore assembly using stringent ndc10 mutants. Surprisingly, a partial loss of function ndc10 mutant rescues the microtubule dynamics defects in the absence of Stu2 activity, suggesting that the kinetochore and Stu2 differentially regulate spindle microtubule turnover in budding yeast.
67. K.A. Kosco, C.G. Pearson, P.S. Maddox, P.J. Wang, I.R. Adams, E.D. Salmon, K. Bloom, and T.C. Huffaker Control of microtubule dynamics by Stu2p is essential for spindle orientation and metaphase chromosome alignment in yeast Mol Biol Cell 12 2001 2870 2880
68. •].
69. K. Kinoshita, B. Habermann, and A.A. Hyman XMAP215: a key component of the dynamic microtubule cytoskeleton Trends Cell Biol 12 2002 267 273
70. •].
71. A.V. Popov, and E. Karsenti Stu2p and XMAP215: turncoat microtubule-associated proteins? Trends Cell Biol 13 2003 547 550
72. C.E. Walczak, T.J. Mitchison, and A. Desai XKCM1: a Xenopus kinesin-related protein that regulates microtubule dynamics during mitotic spindle assembly Cell 84 1996 37 47
73. R. Tournebize, A. Popov, K. Kinoshita, A.J. Ashford, S. Rybina, A. Pozniakovsky, T.U. Mayer, C.E. Walczak, E. Karsenti, and A.A. Hyman Control of microtubule dynamics by the antagonistic activities of XMAP215 and XKCM1 in Xenopus egg extracts Nat Cell Biol 2 2000 13 19
74. K. Kinoshita, I. Arnal, A. Desai, D.N. Drechsel, and A.A. Hyman Reconstitution of physiological microtubule dynamics using purified components Science 294 2001 1340 1343
75. T. Maney, M. Wagenbach, and L. Wordeman Molecular dissection of the microtubule depolymerizing activity of mitotic centromere-associated kinesin J Biol Chem 276 2001 34753 34758
76. S.L. Kline-Smith, and C.E. Walczak The microtubule-destabilizing kinesin XKCM1 regulates microtubule dynamic instability in cells Mol Biol Cell 13 2002 2718 2731
77. F. Gergely, V.M. Draviam, and J.W. Raff The ch-TOG/XMAP215 protein is essential for spindle pole organization in human somatic cells Genes Dev 17 2003 336 341
78. P. Holmfeldt, S. Stenmark, and M. Gullberg Differential functional interplay of TOGp/XMAP215 and the KinI kinesin MCAK during interphase and mitosis EMBO J 23 2004 627 637
79. L. Cassimeris, and J. Morabito TOGp, the human homolog of XMAP215/Dis1, is required for centrosome integrity, spindle pole organization, and bipolar spindle assembly Mol Biol Cell 15 2004 1580 1590
80. D. Cimini, B. Moree, J.C. Canman, and E.D. Salmon Merotelic kinetochore orientation occurs frequently during early mitosis in mammalian tissue cells and error correction is achieved by two different mechanisms J Cell Sci 116 2003 4213 4225 The authors have previously shown that cells recovering from nocodazole depolymerization of microtubules exhibit an increased frequency of improper microtubule attachments at kinetochores. Here, they find that improper attachments occur frequently in early mitosis in untreated cells, although virtually all of them are corrected and do not lead to mis-segregation of chromosomes. High-resolution fluorescence imaging suggests that error detection and correction mechanisms resolve these attachments both before and during anaphase.
81. T.U. Tanaka Bi-orienting chromosomes on the mitotic spindle Curr Opin Cell Biol 14 2002 365 371
82. S. Biggins, and A.W. Murray The budding yeast protein kinase Ipl1/Aurora allows the absence of tension to activate the spindle checkpoint Genes Dev 15 2001 3118 3129
83. •].
84. •].
85. T.U. Tanaka, N. Rachidi, C. Janke, G. Pereira, M. Galova, E. Schiebel, M.J. Stark, and K. Nasmyth Evidence that the Ipl1-Sli15 (Aurora kinase-INCENP) complex promotes chromosome bi-orientation by altering kinetochore-spindle pole connections Cell 108 2002 317 329
86. •].
87. •].
88. K.B. Shannon, and E.D. Salmon Chromosome dynamics: new light on Aurora B kinase function Curr Biol 12 2002 R458 R460
89. M.J. Kallio, M.L. McCleland, P.T. Stukenberg, and G.J. Gorbsky Inhibition of aurora B kinase blocks chromosome segregation, overrides the spindle checkpoint, and perturbs microtubule dynamics in mitosis Curr Biol 12 2002 900 905
90. •].
91. X. Li, and R.B. Nicklas Tension-sensitive kinetochore phosphorylation and the chromosome distribution checkpoint in praying mantid spermatocytes J Cell Sci 110 1997 537 545
92. R.B. Nicklas, S.C. Ward, and G.J. Gorbsky Kinetochore chemistry is sensitive to tension and may link mitotic forces to a cell cycle checkpoint J Cell Biol 130 1995 929 939
93. I.M. Cheeseman, S. Anderson, M. Jwa, E.M. Green, J. Kang, J.R. Yates III, C.S. Chan, D.G. Drubin, and G. Barnes Phospho-regulation of kinetochore-microtubule attachments by the Aurora kinase Ipl1p Cell 111 2002 163 172
94. T. Tanaka, J. Fuchs, J. Loidl, and K. Nasmyth Cohesin ensures bipolar attachment of microtubules to sister centromeres and resists their precocious separation Nat Cell Biol 2 2000 492 499
95. E. Sonoda, T. Matsusaka, C. Morrison, P. Vagnarelli, O. Hoshi, T. Ushiki, K. Nojima, T. Fukagawa, I.C. Waizenegger, and J.M. Peters Scc1/Rad21/Mcd1 is required for sister chromatid cohesion and kinetochore function in vertebrate cells Dev Cell 1 2001 759 770
96. ••].
97. A. Salic, J.C. Waters, and T.J. Mitchison Vertebrate shugoshin links sister centromere cohesion and kinetochore microtubule stability in mitosis Cell 118 2004 567 578 The authors identify a new microtubule-associated protein in vertebrates with similarities to Drosophila MEI-S332 and fungal Sgo1 (Shugoshin), proteins that previously have only been implicated in centromere cohesion during meiosis. Sgo binds and stabilizes microtubules in Xenopus egg extracts. In addition, Sgo is present at kinetochores in HeLa cells, where it appears to stabilize cohesive forces between sister chromatids and is also required for kinetochore-microtubule stability. Thus, Sgo analysis reveals an exciting potential link between chromatid cohesion and kinetochore-microtubule stability.
98. P.D. Andrews, E. Knatko, W.J. Moore, and J.R. Swedlow Mitotic mechanics: the auroras come into view Curr Opin Cell Biol 15 2003 672 683
99. A. Romano, A. Guse, I. Krascenicova, H. Schnabel, R. Schnabel, and M. Glotzer 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 2003 229 236
100. •].
101. ••].
102. S.L. Kline-Smith, and C.E. Walczak Mitotic spindle assembly and chromosome segregation: refocusing on microtubule dynamics Mol Cell 15 2004 317 327
103. ••].
104. ••].
105. •].
106. R. Ohi, M.L. Coughlin, W.S. Lane, and T.J. Mitchison An inner centromere protein that stimulates the microtubule depolymerizing activity of a KinI kinesin Dev Cell 5 2003 309 321 In an effort to identify novel microtubule binding proteins from Xenopus egg extracts, the authors discover ICIS, an F-box domain protein with a novel role in regulating microtubule dynamics. ICIS increases the depolymerization activity of purified MCAK in vitro, immunoprecipitates with MCAK, INCENP and Aurora B from egg extracts, and is required for bipolar spindle formation. By immuno-electron microscopy, ICIS is localized to the inner centromere and on microtubules coming into contact with this region. On the basis of these findings, the authors propose that ICIS may be used at the centromere to enhance the depolymerase activity of MCAK during microtubule attachment correction.
107. C.L. Rieder, and A. Khodjakov Mitosis through the microscope: advances in seeing inside live dividing cells Science 300 2003 91 96
108. C. Obuse, O. Iwasaki, T. Kiyomitsu, G. Goshima, Y. Toyoda, and M. Yanagida A conserved Mis12 centromere complex is linked to heterochromatic HP1 and outer kinetochore protein Zwint-1 Nat Cell Biol 6 2004 1135 1141
109. Maiato H, Rieder CL and Khodjakov A: Kinetochore-driven formation of kinetochore fibers contributes to spindle assembly during animal mitosis. J Cell Biol 2004, in press.
110. Maiato H, Khodjakov A and Rieder CL: Drosophila CLASP is required for microtubule subunit incorporation into fluxing kinetochore fibers. Nat Cell Biol 2004, in press.