544:
383:
151:
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559:. The axis of cell division is determined by the orientation of the spindle apparatus. Cells divide along the line connecting two centrosomes of the spindle apparatus. After formation, the spindle apparatus undergoes rotation inside the cell. The astral microtubules originating from centrosomes reach the cell membrane where they are pulled towards specific cortical clues.
38:
410:
near the pole to which it is attached until a microtubule from the opposite pole binds the sister kinetochore. This second attachment further stabilizes kinetochore attachment to the mitotic spindle. Gradually, the bi-oriented chromosome is pulled towards the center of the cell until microtubule tension is balanced on both sides of the
396:, the spindle is predominantly organized by the poleward separation of centrosomal microtubule organizing centers (MTOCs). Spindle microtubules emanate from centrosomes and 'seek' out kinetochores; when they bind a kinetochore they become stabilized and exert tension on the chromosomes. In an alternative
494:
are bound together in an amorphous mass of tangled DNA and protein. Mitotic entry triggers a dramatic reorganization of the duplicated genome, resulting in sister chromatids that are disentangled and separated from one another. Chromosomes also shorten in length, up to 10,000-fold in animal cells, in
386:
In the centrosome-mediated "search and capture" model (left), microtubules nucleated from centrosomes contact chromosomes by chance and become stabilized at kinetochores to form the spindle. In the chromatin-mediated "self-organization" model (right), microtubules are nucleated around the vicinity of
465:
Spindle assembly is largely regulated by phosphorylation events catalyzed by mitotic kinases. Cyclin dependent kinase complexes (CDKs) are activated by mitotic cyclins, whose translation increases during mitosis. CDK1 (also called CDC2) is considered the main mitotic kinase in mammalian cells and is
435:
egg extracts and that bipolar arrays of microtubules are formed in the absence of centrosomes and kinetochores. Indeed, it has also been shown that laser ablation of centrosomes in vertebrate cells inhibits neither spindle assembly nor chromosome segregation. Under this scheme, the shape and size of
510:
While these dynamic rearrangements are vitally important to ensure accurate and high-fidelity segregation of the genome, our understanding of mitotic chromosome structure remains largely incomplete. A few specific molecular players have been identified, however: Topoisomerase II uses ATP hydrolysis
430:
In contrast to the search-and-capture mechanism in which centrosomes largely dictate the organization of the mitotic spindle, this model proposes that microtubules are nucleated acentrosomally near chromosomes and spontaneously assemble into anti-parallel bundles and adopt a spindle-like structure.
456:
egg extracts, revealing that the Ran GTP gradient alone is sufficient for spindle assembly. The gradient triggers release of spindle assembly factors (SAFs) from inhibitory interactions via the transport proteins importin Ξ²/Ξ±. The unbound SAFs then promote microtubule nucleation and stabilization
409:
In this model, microtubules are nucleated at microtubule organizing centers and undergo rapid growth and catastrophe to 'search' the cytoplasm for kinetochores. Once they bind a kinetochore, they are stabilized and their dynamics are reduced. The newly mono-oriented chromosome oscillates in space
391:
In a properly formed mitotic spindle, bi-oriented chromosomes are aligned along the equator of the cell with spindle microtubules oriented roughly perpendicular to the chromosomes, their plus-ends embedded in kinetochores and their minus-ends anchored at the cell poles. The precise orientation of
421:
In this model, microtubule organizing centers are localized to the cell poles, their separation driven by microtubule polymerization and 'sliding' of antiparallel spindle microtubules with respect to one another at the spindle midzone mediated by bipolar, plus-end-directed kinesins. Such sliding
333:
are required for proper spindle assembly; in mammals, CLASP1 and CLASP2 both contribute to proper spindle assembly and microtubule dynamics in anaphase. Plus-end polymerization may be further moderated by the EB1 protein, which directly binds the growing ends of microtubules and coordinates the
313:. Although how CLIP170 recognizes plus-ends remains unclear, it has been shown that its homologues protect against catastrophe and promote rescue, suggesting a role for CLIP170 in stabilizing plus-ends and possibly mediating their direct attachment to kinetochores. CLIP-associated proteins like
193:
onset. Microtubule polymerization and depolymerization dynamic drive chromosome congression. Depolymerization of microtubules generates tension at kinetochores; bipolar attachment of sister kinetochores to microtubules emanating from opposite cell poles couples opposing tension forces, aligning
400:
model, microtubules undergo acentrosomal nucleation among the condensed chromosomes. Constrained by cellular dimensions, lateral associations with antiparallel microtubules via motor proteins, and end-on attachments to kinetochores, microtubules naturally adopt a spindle-like structure with
547:
Cartoon of the dividing epithelium cell surrounded by epithelium tissue. Spindle apparatus rotates inside the cell. The rotation is a result of astral microtubules pulling towards tri-cellular-junctions (TCJ), signaling centers localized at the regions where three cells
495:
a process called condensation. Condensation begins in prophase and chromosomes are maximally compacted into rod-shaped structures by the time they are aligned in the middle of the spindle at metaphase. This gives mitotic chromosomes the classic "X" shape seen in
392:
this complex is required to ensure accurate chromosome segregation and to specify the cell division plane. However, it remains unclear how the spindle becomes organized. Two models predominate the field, which are synergistic and not mutually exclusive. In the
354:
to induce destabilizing conformational changes in protofilament structure that cause kinesin release and microtubule depolymerization. Loss of their activity results in numerous mitotic defects. Additional microtubule destabilizing proteins include
253:
spindles lack centrosomes or asters at the spindle poles, respectively, and occur for example during female meiosis in most animals. In this instance, a Ran GTP gradient is the main regulator of spindle microtubule organization and assembly. In
452:) is attached to nucleosomes via core histones H2A and H2B. Thus, a gradient of GTP-bound Ran is generated around the vicinity of mitotic chromatin. Glass beads coated with RCC1 induce microtubule nucleation and bipolar spindle formation in
337:
Opposing the action of these microtubule-stabilizing proteins are a number of microtubule-depolymerizing factors which permit the dynamic remodeling of the mitotic spindle to promote chromosome congression and attainment of
511:
to catalyze decatenation of DNA entanglements, promoting sister chromatid resolution. Condensins are 5-subunit complexes that also use ATP-hydrolysis to promote chromosome condensation. Experiments in
350:
XKCM1. MCAK localizes to the growing tips of microtubules at kinetochores where it can trigger catastrophe in direct competition with stabilizing +TIP activity. These proteins harness the energy of
571:
localized at cell vertices. The spatial distribution of cortical clues leads to the force field that determine final spindle apparatus orientation and the subsequent orientation of cell division.
531:. If chromosomes are not properly attached to the mitotic spindle by the time of this checkpoint, the onset of anaphase will be delayed. Failure of this spindle assembly checkpoint can result in
305:
The growing ends of microtubules are protected against catastrophe by the action of plus-end microtubule tracking proteins (+TIPs) to promote their association with kinetochores at the midzone.
317:
in humans have also been shown to localize to plus-ends and the outer kinetochore as well as to modulate the dynamics of kinetochore microtubules (Maiato 2003). CLASP homologues in
346:-13 superfamily of MAPs contains a class of plus-end-directed motor proteins with associated microtubule depolymerization activity including the well-studied mammalian MCAK and
474:
is a member of the chromosomal passenger complex and mediates chromosome-microtubule attachment and sister chromatid cohesion. Polo-like kinase, also known as PLK, especially
568:
1629:
R. Heald; R. Tournebize; et al. (1996). "Self-organization of microtubules into bipolar spindles around artificial chromosomes in
Xenopus egg extracts".
366:
The activities of these MAPs are carefully regulated to maintain proper microtubule dynamics during spindle assembly, with many of these proteins serving as
2323:
Bosveld F, Markova O, Guirao B, Martin C, Wang Z, Pierre A, Balakireva M, Gaugue I, Ainslie A, Christophorou N, Lubensky DK, Minc N, BellaΓ―che Y (2016).
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Marko, JF. The mitotic chromosome: structure and
Mechanics. 2012. Genome Organization and Function in the Cell Nucleus. Wiley-VCH, Ch. 18, 449-485.
555:
is of major importance for tissue architecture, cell fates and morphogenesis. Cells tend to divide along their long axis according to the so-called
1981:
194:
chromosomes at the cell equator and poising them for segregation to daughter cells. Once every chromosome is bi-oriented, anaphase commences and
161:
via a multiprotein complex called the kinetochore. Polar microtubules interdigitate at the spindle midzone and push the spindle poles apart via
1231:"EB1-microtubule interactions in Xenopus egg extracts: Role of EB1 in microtubule stabilization and mechanisms of targeting to microtubules"
2272:
Thery M, Jimenez-Dalmaroni A, Racine V, Bornens M, Julicher F (2007). "Experimental and theoretical study of mitotic spindle orientation".
1938:
Peters, U., J. Cherian; et al. (2006). "Probing cell-division phenotype space and Polo-like kinase function using small molecules".
302:. The microtubule-associated protein Augmin acts in conjunction with Ξ³-TURC to nucleate new microtubules off of existing microtubules.
278:
determines to a large extent the shape of the mitotic spindle and promotes the proper alignment of chromosomes at the spindle midzone.
1525:"The bipolar kinesin, KLP61F, cross-links microtubules within interpolar microtubule bundles of Drosophila embryonic mitotic spindles"
431:
Classic experiments by Heald and
Karsenti show that functional mitotic spindles and nuclei form around DNA-coated beads incubated in
298:
into microtubules. Recruitment of Ξ³-TuRC to the pericentrosomal region stabilizes microtubule minus-ends and anchors them near the
1186:
A. Akhmanova; M.O. Steinmetz (April 2008). "Tracking the ends: a dynamic protein network controls the fate of microtubule tips".
282:(MAPs) associate with microtubules at the midzone and the spindle poles to regulate their dynamics. Ξ³-tubulin is a specialized
2002:
422:
forces may account not only for spindle pole separation early in mitosis, but also spindle elongation during late anaphase.
2389:
1372:
H. Maiato; P Sampaio; C.E. Sunkel (2004). "Microtubule-associated proteins and their essential roles during mitosis".
154:
This diagram depicts the organization of a typical mitotic spindle found in animal cells. Chromosomes are attached to
2221:
1403:
768:
678:
626:
2237:
Baker DJ, Chen J, van
Deursen JM (2005). "The mitotic checkpoint in cancer and aging: what have mice taught us?".
1139:"Mammalian CLASP1 and CLASP2 Cooperate to Ensure Mitotic Fidelity by Regulating Spindle and Kinetochore Function"
363:
which have roles in remodeling the mitotic spindle as well as promoting chromosome segregation during anaphase.
279:
17:
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Morgan DO: The Cell Cycle: Principles of
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339:
1429:"Control of microtubule dynamics by the antagonistic activities of XMAP215 and XKCM1 in Xenopus egg extracts"
299:
174:
2165:"Histone H1 is essential for mitotic chromosome architecture and segregation in Xenopus laevis egg extracts"
528:
234:
186:
309:
was shown to localize near microtubule plus-ends in HeLa cells and to accumulate in kinetochores during
651:
436:
the mitotic spindle are a function of the biophysical properties of the cross-linking motor proteins.
229:
that may be present at the spindle poles depending on the cell type. The spindle apparatus is vaguely
552:
543:
233:
in cross section and tapers at the ends. In the wide middle portion, known as the spindle midzone,
31:
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activated by Cyclin B1. Aurora kinases are required for proper spindle assembly and separation.
319:
527:
The completion of spindle formation is a crucial transition point in the cell cycle called the
291:
2405:
1975:
1080:"Tip1/CLIP-170 Protein Is Required for Correct Chromosome Poleward Movement in Fission Yeast"
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around mitotic chromatin, and spindle bipolarity is organized by microtubule motor proteins.
275:
287:
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1783:
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The dynamic lengthening and shortening of spindle microtubules, through a process known as
226:
884:"Identification of a novel nucleotide-sensitive microtubule-binding protein in HeLa cells"
8:
2325:"Epithelial tricellular junctions act as interphase cell shape sensors to orient mitosis"
166:
2340:
2285:
2016:"Large-scale chromatin organization: The good, the surprising, and the still perplexing"
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1642:
1589:
1095:
933:"Evidence for a role of CLIP-170 in the establishment of metaphase chromosome alignment"
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C. E. Walczak; R. Heald (2008). "Mechanisms of
Mitotic Spindle Assembly and Function".
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1385:
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998:
981:
835:"Branching microtubule nucleation in Xenopus egg extracts mediated by augmin and TPX2"
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D. Dujardin; U.I. Wacker; A. Moreau; T.A. Schroer; J.E. Rickard; J.R. DeMey (1998).
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has important roles in the spindle maintenance by regulating microtubule dynamics.
371:
263:
203:
199:
100:
1878:
Fu J, Jiang Q, Zhang C (2010). "Coordination of Cell Cycle Events by Ran GTPase".
802:
785:
1845:
1476:"The distribution of spindle microtubules during mitosis in cultured human cells"
1104:
580:
487:
415:
241:. At the pointed ends, known as spindle poles, microtubules are nucleated by the
92:
1367:
1365:
1329:
982:"CLIP-170-like tip1p spatially organizes microtubular dynamics in fission yeast"
2067:
1078:
S. Goldstone; C. Reyes; G. Gay; T. CourthΓ©oux; M. Dubarry; et al. (2010).
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104:
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Champoux JJ (2001). "DNA TOPOISOMERASES: Structure, Function, and
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2031:
1772:"Structure of the RCC1 chromatin factor bound to the nucleosome core particle"
1747:
1730:
1597:
1283:"Regulation of localization and activity of the microtubule depolymerase MCAK"
1228:
425:
2399:
1426:
1362:
1332:"XKCM1 acts on a single protofilament and requires the C terminus of tubulin"
445:
367:
170:
162:
96:
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1829:"Mitotic spindle assembly around RCC1-coated beads in Xenopus egg extracts"
1813:
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1509:
1452:
1413:
1355:
1347:
1330:
H. Niederstrasser; H. Salehi-Had; E.C. Gan; C. Walczak; E. Nogales (2002).
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R. Tournebize; A. Popov; K. Kinoshita; A.J. Ashford; et al. (2000).
295:
210:
182:
158:
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128:
61:
53:
2348:
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2116:"Condensins: universal organizers of chromosomes with diverse functions"
1951:
1795:
563:, the distribution of cortical clues is set up by the adhesive pattern.
1915:
1898:
1394:
1229:
J.S. Tirnauer; S. Grego; E.D. Salmon; T.J. Mitchison (1 October 2002).
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A.L. Pereira; A.J. Pereira; A.R.R. Maia; et al. (1 October 2006).
1077:
727:
532:
516:
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associates with centrosomes and is believed to regulate mitotic entry.
411:
387:
mitotic chromatin and organized into a bipolar array by motor proteins.
269:
242:
222:
115:, a process that produces genetically identical daughter cells, or the
45:
41:
134:
Besides chromosomes, the spindle apparatus is composed of hundreds of
1728:
1650:
1280:
712:"Structure-function insights into the yeast Dam1 kinetochore complex"
499:, with each condensed sister chromatid linked along their lengths by
230:
69:
1331:
1199:
150:
2271:
1136:
471:
467:
356:
190:
2211:
1444:
30:
This article is about the cellular structure. For other uses, see
1682:"Centrosome-independent mitotic spindle formation in vertebrates"
1522:
500:
440:
Chromatin-mediated microtubule nucleation by the Ran GTP gradient
418:
until anaphase onset releases cohesion of the sister chromatids.
360:
343:
283:
238:
214:
195:
135:
124:
120:
112:
73:
1031:"Cytoplasmic linker proteins promote microtubule rescue in vivo"
786:"Centrosome reduction during gametogenesis and its significance"
783:
2384:
314:
255:
218:
37:
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1371:
330:
306:
65:
519:
as an important regulator of mitotic chromosome compaction.
444:
The guanine nucleotide exchange factor for the small GTPase
1523:
D.J. Sharp; K.L. McDonald; H.M. Brown; et al. (1999).
475:
449:
426:
Chromatin-mediated self-organization of the mitotic spindle
57:
49:
2322:
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1185:
535:
and may be involved in aging and the formation of cancer.
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181:
Attachment of microtubules to chromosomes is mediated by
1028:
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comprise the most abundant components of the machinery.
1769:
1937:
1473:
881:
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661:. International Review of Cell and Molecular Biology.
522:
1628:
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758:
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652:"Interplay between spindle architecture and function"
604:
2236:
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270:
Microtubule-associated proteins and spindle dynamics
209:
The cellular spindle apparatus includes the spindle
1896:
1827:Halpin D, Kalab P, Wang J, Weis K, Heald R (2011).
1826:
503:proteins and joined, often near the center, at the
414:; the congressed chromosome then oscillates at the
377:
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286:variant that assembles into a ring complex called
1574:"Anastral spindle assembly: a mathematical model"
649:
460:
221:molecular motors, condensed chromosomes, and any
173:. Microtubule polymerization is nucleated at the
2397:
2316:
2265:
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567:polarity cues are determined by localization of
538:
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1993:
1991:
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2230:
1980:: CS1 maint: multiple names: authors list (
1877:
401:chromosomes aligned along the cell equator.
266:, which does not break down during mitosis.
202:, is severed, permitting the transit of the
777:
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448:(Regulator of chromosome condensation 1 or
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2139:
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801:
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735:
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14:
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2205:
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213:, associated proteins, which include
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169:anchor the spindle poles to the
2156:
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2072:
2056:
2007:
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1022:
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461:Regulation of spindle assembly
294:polymerization of Ξ±/Ξ² tubulin
13:
1:
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300:microtubule-organizing center
175:microtubule organizing center
1846:10.1371/journal.pbio.1001225
1105:10.1371/journal.pone.0010634
482:Mitotic chromosome structure
237:microtubules are bundled by
7:
574:
529:spindle assembly checkpoint
107:. It is referred to as the
10:
2427:
2068:10.1002/9783527639991.ch18
851:10.1016/j.cell.2012.12.044
123:, a process that produces
29:
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185:, which actively monitor
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394:search-and-capture model
334:binding of other +TIPs.
258:, spindles form between
127:with half the number of
32:Spindle (disambiguation)
1155:10.1091/mbc.E06-07-0579
2132:10.1101/gad.194746.112
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1247:10.1091/mbc.02-04-0210
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189:and prevent premature
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276:dynamic instability
260:spindle pole bodies
206:to opposite poles.
167:Astral microtubules
1916:10.1242/jcs.013136
728:10.1242/jcs.004689
550:
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179:
95:that forms during
77:
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2390:Spindle apparatus
2388:Media related to
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1637:(6590): 420β425.
1241:(10): 3614β3626.
1149:(10): 4526β4542.
722:(21): 3831β3836.
492:sister chromatids
204:sister chromatids
200:sister chromatids
187:spindle formation
146:Spindle structure
103:between daughter
101:sister chromatids
85:spindle apparatus
16:(Redirected from
2418:
2387:
2371:
2370:
2360:
2320:
2314:
2313:
2269:
2263:
2262:
2234:
2228:
2227:
2209:
2203:
2202:
2192:
2160:
2154:
2153:
2143:
2111:
2105:
2104:
2081:Annu Rev Biochem
2076:
2070:
2060:
2054:
2053:
2043:
2011:
2005:
1995:
1986:
1985:
1979:
1971:
1935:
1929:
1928:
1918:
1894:
1888:
1887:
1880:Nature Education
1875:
1869:
1868:
1858:
1848:
1839:(12): e1001225.
1824:
1818:
1817:
1807:
1767:
1761:
1760:
1750:
1726:
1720:
1719:
1701:
1677:
1671:
1670:
1651:10.1038/382420a0
1626:
1620:
1619:
1609:
1584:(8): 2191β2201.
1569:
1563:
1562:
1552:
1520:
1514:
1513:
1503:
1471:
1465:
1464:
1424:
1418:
1417:
1397:
1369:
1360:
1359:
1327:
1321:
1320:
1310:
1278:
1269:
1268:
1258:
1226:
1220:
1219:
1183:
1177:
1176:
1166:
1134:
1128:
1127:
1117:
1107:
1075:
1069:
1068:
1058:
1026:
1020:
1019:
1001:
977:
971:
970:
960:
928:
922:
921:
911:
894:(5): 1623β1633.
879:
873:
872:
862:
830:
824:
823:
805:
781:
775:
774:
756:
750:
749:
739:
707:
701:
700:
656:
647:
641:
640:
602:
372:Polo-like kinase
264:nuclear envelope
262:embedded in the
198:, which couples
93:eukaryotic cells
21:
2426:
2425:
2421:
2420:
2419:
2417:
2416:
2415:
2396:
2395:
2380:
2375:
2374:
2335:(7591): 496β8.
2321:
2317:
2280:(7143): 493β6.
2270:
2266:
2235:
2231:
2224:
2210:
2206:
2161:
2157:
2112:
2108:
2077:
2073:
2061:
2057:
2012:
2008:
1996:
1989:
1973:
1972:
1936:
1932:
1895:
1891:
1876:
1872:
1825:
1821:
1768:
1764:
1727:
1723:
1678:
1674:
1627:
1623:
1570:
1566:
1521:
1517:
1472:
1468:
1425:
1421:
1406:
1370:
1363:
1328:
1324:
1287:Bioarchitecture
1279:
1272:
1227:
1223:
1200:10.1038/nrm2369
1184:
1180:
1135:
1131:
1076:
1072:
1027:
1023:
978:
974:
929:
925:
880:
876:
831:
827:
782:
778:
771:
757:
753:
708:
704:
681:
654:
648:
644:
629:
603:
599:
594:
581:Centralspindlin
577:
541:
525:
488:DNA replication
484:
463:
442:
428:
416:metaphase plate
407:
380:
272:
148:
117:meiotic spindle
109:mitotic spindle
35:
28:
23:
22:
15:
12:
11:
5:
2424:
2414:
2413:
2408:
2394:
2393:
2379:
2378:External links
2376:
2373:
2372:
2315:
2264:
2229:
2222:
2204:
2155:
2106:
2087:(1): 369β413.
2071:
2055:
2006:
1987:
1946:(11): 618β26.
1930:
1889:
1870:
1819:
1762:
1721:
1672:
1621:
1564:
1535:(1): 125β138.
1515:
1486:(2): 468β497.
1466:
1419:
1404:
1361:
1342:(3): 817β828.
1322:
1270:
1221:
1194:(4): 309β322.
1178:
1129:
1070:
1041:(4): 589β599.
1021:
992:(5): 695β704.
972:
943:(4): 849β862.
923:
874:
845:(4): 768β777.
825:
776:
769:
751:
702:
679:
642:
627:
596:
595:
593:
590:
589:
588:
586:Spindle poison
583:
576:
573:
540:
537:
524:
521:
486:By the end of
483:
480:
462:
459:
441:
438:
427:
424:
406:
403:
379:
376:
352:ATP hydrolysis
271:
268:
163:motor proteins
147:
144:
26:
18:Spindle fibers
9:
6:
4:
3:
2:
2423:
2412:
2409:
2407:
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2391:
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2225:
2223:0-7167-1007-2
2219:
2215:
2208:
2200:
2196:
2191:
2186:
2182:
2178:
2175:(6): 859β69.
2174:
2170:
2166:
2159:
2151:
2147:
2142:
2137:
2133:
2129:
2125:
2121:
2117:
2110:
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1994:
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1983:
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1969:
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1949:
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1940:Nat Chem Biol
1934:
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1493:
1489:
1485:
1481:
1477:
1470:
1462:
1458:
1454:
1450:
1446:
1445:10.1038/71330
1442:
1438:
1434:
1433:Nat Cell Biol
1430:
1423:
1415:
1411:
1407:
1405:9780123646453
1401:
1396:
1391:
1387:
1383:
1379:
1375:
1374:Int Rev Cytol
1368:
1366:
1357:
1353:
1349:
1345:
1341:
1337:
1333:
1326:
1318:
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1277:
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1248:
1244:
1240:
1236:
1235:Mol Biol Cell
1232:
1225:
1217:
1213:
1209:
1205:
1201:
1197:
1193:
1189:
1182:
1174:
1170:
1165:
1160:
1156:
1152:
1148:
1144:
1143:Mol Biol Cell
1140:
1133:
1125:
1121:
1116:
1111:
1106:
1101:
1097:
1093:
1090:(5): e10634.
1089:
1085:
1081:
1074:
1066:
1062:
1057:
1052:
1048:
1044:
1040:
1036:
1032:
1025:
1017:
1013:
1009:
1005:
1000:
995:
991:
987:
983:
976:
968:
964:
959:
954:
950:
946:
942:
938:
934:
927:
919:
915:
910:
905:
901:
897:
893:
889:
885:
878:
870:
866:
861:
856:
852:
848:
844:
840:
836:
829:
821:
817:
813:
809:
804:
799:
795:
791:
787:
780:
772:
770:0-8053-7171-0
766:
762:
755:
747:
743:
738:
733:
729:
725:
721:
717:
713:
706:
698:
694:
690:
686:
682:
680:9780124076945
676:
672:
668:
664:
660:
653:
646:
638:
634:
630:
628:9780123743329
624:
620:
616:
612:
608:
601:
597:
587:
584:
582:
579:
578:
572:
570:
566:
562:
558:
554:
545:
536:
534:
530:
520:
518:
514:
508:
506:
502:
498:
493:
489:
479:
477:
473:
469:
458:
455:
451:
447:
437:
434:
423:
419:
417:
413:
402:
399:
398:self assembly
395:
384:
375:
373:
369:
364:
362:
358:
353:
349:
345:
341:
335:
332:
328:
327:
322:
321:
316:
312:
308:
303:
301:
297:
293:
289:
285:
281:
277:
267:
265:
261:
257:
252:
248:
244:
240:
236:
232:
228:
224:
220:
216:
212:
207:
205:
201:
197:
192:
188:
184:
176:
172:
171:cell membrane
168:
164:
160:
157:
152:
143:
141:
137:
132:
130:
126:
122:
118:
114:
110:
106:
102:
98:
97:cell division
94:
91:structure of
90:
86:
82:
75:
71:
67:
63:
59:
55:
51:
47:
43:
39:
33:
19:
2406:Cell anatomy
2332:
2328:
2318:
2277:
2273:
2267:
2245:(6): 583β9.
2242:
2238:
2232:
2213:
2207:
2172:
2169:J. Cell Biol
2168:
2158:
2123:
2119:
2109:
2084:
2080:
2074:
2058:
2023:
2019:
2009:
1976:cite journal
1943:
1939:
1933:
1906:
1902:
1892:
1883:
1879:
1873:
1836:
1832:
1822:
1779:
1775:
1765:
1738:
1734:
1724:
1692:(2): 59β67.
1689:
1685:
1675:
1634:
1630:
1624:
1581:
1577:
1567:
1532:
1528:
1518:
1483:
1479:
1469:
1439:(1): 13β19.
1436:
1432:
1422:
1377:
1373:
1339:
1335:
1325:
1293:(2): 80β87.
1290:
1286:
1238:
1234:
1224:
1191:
1187:
1181:
1146:
1142:
1132:
1087:
1083:
1073:
1038:
1034:
1024:
989:
985:
975:
940:
936:
926:
891:
887:
877:
842:
838:
828:
793:
790:Biol. Reprod
789:
779:
760:
754:
719:
715:
705:
662:
658:
645:
610:
606:
600:
564:
560:
557:Hertwig rule
551:
526:
512:
509:
485:
464:
453:
443:
432:
429:
420:
408:
397:
393:
390:
374:substrates.
365:
347:
336:
324:
318:
311:prometaphase
304:
296:heterodimers
273:
250:
247:Acentrosomal
246:
235:antiparallel
211:microtubules
208:
183:kinetochores
180:
159:microtubules
140:Microtubules
133:
116:
108:
99:to separate
89:cytoskeletal
84:
81:cell biology
78:
62:microtubules
54:kinetochores
1529:J Cell Biol
1480:J Cell Biol
1395:10216/53621
1035:J Cell Biol
937:J Cell Biol
888:J Cell Biol
796:(1): 2β13.
613:: 111β158.
243:centrosomes
223:centrosomes
156:kinetochore
129:chromosomes
46:chromosomes
2400:Categories
1903:J Cell Sci
1380:: 53β153.
1336:J Mol Biol
716:J Cell Sci
665:: 83β125.
592:References
533:aneuploidy
517:Histone H1
505:centromere
497:karyotypes
412:centromere
340:bipolarity
320:Drosophila
42:Micrograph
2120:Genes Dev
2026:: 69β78.
1833:PLOS Biol
1735:Curr Biol
1686:Curr Biol
1578:Biophys J
292:nucleates
231:ellipsoid
70:metaphase
2367:26886796
2302:17495931
2259:16226453
2199:15967810
2150:22855829
2101:11395412
2050:24529248
1968:22213611
1960:17028580
1925:17715155
1886:(9): 32.
1865:22215983
1814:20739938
1757:17702580
1708:10662665
1616:19843451
1510:19866774
1461:10732643
1453:10620801
1414:15548419
1356:11866534
1317:21866268
1265:12388761
1216:24977579
1208:18322465
1173:16914514
1124:20498706
1084:PLOS ONE
1065:12446741
1016:11948950
1008:11007487
869:23415226
820:37305534
812:15385423
746:19889968
689:24016524
637:18275887
575:See also
561:In vitro
472:Aurora B
468:Aurora A
357:stathmin
251:anastral
239:kinesins
191:anaphase
136:proteins
2411:Mitosis
2358:5450930
2337:Bibcode
2310:4391685
2282:Bibcode
2190:2171634
2141:3418584
2041:3927141
1856:3246454
1805:3168546
1784:Bibcode
1716:9976687
1667:4238425
1659:8684481
1639:Bibcode
1607:2764103
1586:Bibcode
1559:9885249
1550:2148119
1501:2108320
1308:3158623
1164:1635371
1115:2869355
1092:Bibcode
1056:2173097
967:9585405
958:2132766
918:1970824
909:2200191
860:3680348
737:2773187
697:8145444
565:In vivo
513:Xenopus
501:cohesin
454:Xenopus
433:Xenopus
361:katanin
348:Xenopus
344:kinesin
326:Xenopus
307:CLIP170
284:tubulin
215:kinesin
196:cohesin
125:gametes
121:meiosis
119:during
113:mitosis
111:during
87:is the
74:mitosis
68:during
2365:
2355:
2329:Nature
2308:
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2274:Nature
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1966:
1958:
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1776:Nature
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1631:Nature
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368:Aurora
342:. The
329:, and
315:CLASP1
290:which
288:Ξ³-TuRC
227:asters
219:dynein
83:, the
60:, and
2306:S2CID
1964:S2CID
1712:S2CID
1663:S2CID
1457:S2CID
1212:S2CID
1012:S2CID
816:S2CID
693:S2CID
655:(PDF)
548:meet.
355:Op18/
331:yeast
256:fungi
105:cells
66:green
2363:PMID
2298:PMID
2255:PMID
2218:ISBN
2195:PMID
2146:PMID
2097:PMID
2046:PMID
1999:ISBN
1982:link
1956:PMID
1921:PMID
1861:PMID
1810:PMID
1753:PMID
1704:PMID
1655:PMID
1612:PMID
1555:PMID
1506:PMID
1449:PMID
1410:PMID
1400:ISBN
1352:PMID
1313:PMID
1261:PMID
1204:PMID
1169:PMID
1120:PMID
1061:PMID
1004:PMID
986:Cell
963:PMID
914:PMID
865:PMID
839:Cell
808:PMID
765:ISBN
742:PMID
685:PMID
675:ISBN
633:PMID
623:ISBN
476:PLK1
450:RCC1
370:and
359:and
217:and
58:pink
50:blue
2353:PMC
2345:doi
2333:530
2290:doi
2278:447
2247:doi
2185:PMC
2177:doi
2173:169
2136:PMC
2128:doi
2089:doi
2064:doi
2036:PMC
2028:doi
1948:doi
1911:doi
1907:120
1851:PMC
1841:doi
1800:PMC
1792:doi
1780:467
1743:doi
1694:doi
1647:doi
1635:382
1602:PMC
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1545:PMC
1537:doi
1533:144
1496:PMC
1488:doi
1441:doi
1390:hdl
1382:doi
1378:241
1344:doi
1340:316
1303:PMC
1295:doi
1251:PMC
1243:doi
1196:doi
1159:PMC
1151:doi
1110:PMC
1100:doi
1051:PMC
1043:doi
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732:PMC
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