296:
20:
239:. Cdc14 is activated by its release into the nucleus, from sequestration in the nucleolus, and subsequent export into the cytoplasm. The Cdc-14 Early Anaphase Release pathway, which stabilizes the spindle, also releases cdc14 from the nucleolus but restricts it to the nucleus. Complete release and maintained activation of cdc14 is achieved by the separate Mitotic Exit Network (MEN) pathway to a sufficient degree (to trigger the spindle disassembly and nuclear envelope assembly) only after late anaphase.
1523:
28:
354:(yeast Kip3), an ATP-dependent depolymerase, accelerate microtubule depolymerization at the plus end. It was shown the concurrent disruption of these mechanisms, but not of any one, results in dramatic spindle hyperstability during telophase, suggesting functional overlap despite the diversity of the mechanisms.
393:
during early mitosis. Ran-GTP localizes near chromosomes throughout mitosis, but does not trigger the dissociation of nuclear envelope proteins from importin β until M-Cdk targets are dephosphorylated in telophase. These envelope components include several nuclear pore components, the most studied of
481:
is reestablished. If lamin transport through nuclear pores is prevented, chromosomes remain condensed following cytokinesis, and cells fail to reenter the next S phase. In mammals, DNA licensing for S phase (the association of chromatin to the multiple protein factors necessary for its replication)
423:
In cells where the nuclear membrane is absorbed into the endoplasmic reticulum during mitosis, reassembly involves the lateral expansion around the chromatin with stabilization of the expanding membrane over the surface of the chromatin. Studies claiming this mechanism is a prerequisite to nuclear
337:
of M-Cdk substrates by the MEN is broadly held to be responsible for spindle disassembly. The phosphorylation states of microtubule stabilizing and destabilizing factors, as well as microtubule nucleators are key regulators of their activities. For example, NuMA is a minus-end crosslinking protein
246:
allows the APC/C to bind CDH1. APC/C targets CDC20 for proteolysis, resulting in a cellular switch from APC/C to APC/C activity. The ubiquitination of mitotic cyclins continues along with that of APC/C-specific targets such as the yeast mitotic spindle component, Ase1, and cdc5, the degradation of
160:
of the protein targets of M-Cdks (Mitotic Cyclin-dependent
Kinases) drives spindle assembly, chromosome condensation and nuclear envelope breakdown in early mitosis. The dephosphorylation of these same substrates drives spindle disassembly, chromosome decondensation and the reformation of daughter
473:
Chromosome decondensation (also known as relaxation or decompaction) into expanded chromatin is necessary for the cell's resumption of interphase processes, and occurs in parallel to nuclear envelope assembly during telophase in many eukaryotes. MEN-mediated Cdk dephosphorylation is necessary for
318:
Spindle depolymerization during telophase occurs from the plus end and is, in this way, a reversal of spindle assembly. Subsequent microtubule array assembly is, unlike that of the polarized spindle, interpolar. This is especially apparent in animal cells which must immediately, following mitotic
341:
A general model for spindle disassembly in yeast is that the three functionally overlapping subprocesses of spindle disengagement, destabilization, and depolymerization are primarily effected by APC/C, microtubule-stabilizer-specific kinases, and plus-end directed microtubule depolymerases,
436:
egg extracts failed to smoothen when nuclear import of lamin was inhibited, remaining wrinkled and closely bound to condensed chromosomes. However, in the case of ER lateral expansion, nuclear import is initiated before completion of the nuclear envelope reassembly, leading to a temporary
406:
dimers and nucleosomes. After binding to chromatin, ELYS recruits other components of the nuclear pore scaffold and nuclear pore trans-membrane proteins. The nuclear pore complex is assembled and integrated in the nuclear envelope in an organized manner, consecutively adding Nup107-160,
370:
internal to the inner nuclear membrane. These components are dismantled during prophase and prometaphase and reconstructed during telophase, when the nuclear envelope reforms on the surface of separated sister chromatids. The nuclear membrane is fragmented and partly absorbed by the
188:. Experimental addition of non-degradable M-cyclin to cells induces cell cycle arrest in a post-anaphase/pre-telophase-like state with condensed chromosomes segregated to cell poles, an intact mitotic spindle, and no reformation of the nuclear envelope. This has been shown in frog (
282:
activity to alter target protein conformation. Cdc48 is necessary for spindle disassembly, nuclear envelope assembly, and chromosome decondensation. Cdc48 modifies proteins structurally involved in these processes and also some ubiquitinated proteins which are thus targeted to the
414:
It is debated whether the mechanism of nuclear membrane reassembly involves initial nuclear pore assembly and subsequent recruitment of membrane vesicles around the pores or if the nuclear envelope forms primarily from extended ER cisternae, preceding nuclear pore assembly:
482:
also occurs coincidentally with the maturation of the nuclear envelope during late telophase. This can be attributed to and provides evidence for the nuclear import machinery's reestablishment of interphase nuclear and cytoplasmic protein localizations during telophase.
303:
The breaking of the mitotic spindle, common to the completion of mitosis in all eukaryotes, is the event most often used to define the anaphase-B to telophase transition, although the initiation of nuclear reassembly tends to precede that of spindle disassembly.
332:
While spindle assembly has been well studied and characterized as a process where tentative structures are edified by the SAC, the molecular basis of spindle disassembly is not understood in comparable detail. The late-mitotic
419:
In cells where the nuclear membrane fragments into non-ER vesicles during mitosis, a Ran-GTP–dependent pathway can direct these discrete vesicle populations to chromatin where they fuse to reform the nuclear
342:
respectively. These effectors are known to be highly conserved between yeast and higher eukaryotes. The APC/C targets crosslinking microtubule-associated proteins (NuMA, Ase1, Cin1 and more).
1336:
Wiese C, Goldberg MW, Allen TD, Wilson KL (July 1997). "Nuclear envelope assembly in
Xenopus extracts visualized by scanning EM reveals a transport-dependent 'envelope smoothing' event".
307:
Spindle disassembly is an irreversible process which must effect not the ultimate degradation, but the reorganization of constituent microtubules; microtubules are detached from
440:
Lamin subunits disassembled in prophase are inactivated and sequestered during mitosis. Lamina reassembly is triggered by lamin dephosphorylation (and additionally by methyl-
869:
Hetzer M, Meyer HH, Walther TC, Bilbao-Cortes D, Warren G, Mattaj IW (December 2001). "Distinct AAA-ATPase p97 complexes function in discrete steps of nuclear assembly".
235:-anaphase transition. However, the existence of differential phases to cdc14 activity between anaphase and telophase is suggestive of additional, unexplored late-
575:
Juang YL, Huang J, Peters JM, McLaughlin ME, Tai CY, Pellman D (February 1997). "APC-mediated proteolysis of Ase1 and the morphogenesis of the mitotic spindle".
216:
The requirement for phosphatase activation can be seen in budding yeast, which do not have redundant phosphatases for mitotic exit and rely on the phosphatase
379:
to the chromatin occurs during telophase in a reversal of this process. Membrane-forming vesicles aggregate directly to the surface of chromatin, where they
161:
nuclei in telophase. Establishing a degree of dephosphorylation permissive to telophase events requires both the inactivation of Cdks and the activation of
428:
The envelope smoothens and expands following its enclosure of the whole chromatid set. This probably occurs due to the nuclear pores' import of
1555:
424:
pore formation have found that bare-chromatin-associated Nup107–160 complexes are present in single units instead of as assembled pre-pores.
511:
Reece, Jane; Urry, Lisa; Cain, Michael; Wasserman, Steven; Minorsky, Peter; Jackson, Robert (2011). Campbell
Biology (10th ed.). Pearson.
398:, which can recognize DNA regions rich in A:T base pairs (in vitro), and may therefore bind directly to the DNA. However, experiments in
456:
enters the reforming nucleus but continues to slowly assemble into the peripheral lamina over several hours in throughout the G1 phase.
979:"Mitotic spindle disassembly occurs via distinct subprocesses driven by the anaphase-promoting complex, Aurora B kinase, and kinesin-8"
242:
Cdc14-mediated dephosphorylation activates downstream regulatory processes unique to telophase. For example, the dephosphorylation of
31:
Fluorescence micrograph of a human cell in telophase showing chromosomes (DNA) in blue, microtubules in green and kinetochores in pink
1847:
1741:
389:
is required for early nuclear envelope assembly at the surface of the chromosomes: it releases envelope components sequestered by
915:
Aist JR (2002-01-01). "Mitosis and motor proteins in the filamentous ascomycete, Nectria haematococca, and some related fungi".
325:
in order to regulate cytokinesis. The ATPase p97 is required for the establishment of the relatively stable and long interphase
1100:
1035:
666:
630:
559:
516:
465:
Yeast lack lamins; their nuclear envelope remains intact throughout mitosis and nuclear division happens during cytokinesis.
267:
The anaphase-mediated distancing of chromosomes from the metaphase plate may trigger spatial cues for the onset of telophase.
350:(yeast Bim1), which then dissociates from microtubules, and the destabilizer She1, which then associates with microtubules.
1787:
220:. Blocking cdc14 activation in these cells results in the same phenotypic arrest as does blocking M-cyclin degradation.
1548:
1191:
936:
748:
462:
egg extracts and human cancer cell lines have been the primary models used for studying nuclear envelope reassembly.
263:
profile is only the broadest of many regulatory mechanisms contributing to the onset of individual telophase events.
452:). Lamin-B can target chromatin as early as mid-anaphase. During telophase, when nuclear import is reestablished,
376:
338:
and Cdk substrate whose dissociation from the microtubule is effected by its dephosphorylation during telophase.
1065:
Lodish, Harvey; Berk, Arnold; Zipursky, S. Lawrence; Matsudaira, Paul; Baltimore, David; Darnell, James (2000).
326:
1541:
1051:
1471:"GAK is phosphorylated by c-Src and translocated from the centrosome to chromatin at the end of telophase"
766:"Cdc14 Early Anaphase Release, FEAR, Is Limited to the Nucleus and Dispensable for Efficient Mitotic Exit"
1857:
1289:"Formation of the postmitotic nuclear envelope from extended ER cisternae precedes nuclear pore assembly"
1998:
1701:
1527:
271:
177:
1215:
Clarke PR, Zhang C (2004). "Spatial and temporal control of nuclear envelope assembly by Ran GTPase".
1696:
1691:
1686:
1681:
1676:
1671:
1666:
1661:
1656:
209:
1118:"Nucleosomal regulation of chromatin composition and nuclear assembly revealed by histone depletion"
402:
egg extracts have concluded that ELYS fails to associate with bare DNA and will only directly bind
203:
1170:
Gay S, Foiani M (2015-01-01). "Nuclear envelope and chromatin, lock and key of genome integrity".
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334:
197:
145:
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652:
58:
45:
39:
295:
1980:
1731:
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Fukushima K, Wang M, Naito Y, Uchihashi T, Kato Y, Mukai S, Yabuta N, Nojima H (March 2017).
478:
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intra-nuclear protein gradient between the distal and medial aspects of the forming nucleus.
372:
236:
777:
625:(6th ed.). New York, NY: Garland Science, Taylor and Francis Group. pp. 995–996.
8:
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1993:
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228:
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typically begins before late telophase and, when complete, segregates the two daughter
928:
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during prometaphase and the targeting of inner nuclear membrane protein-containing ER
1988:
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Mechanisms of
Mitotic Spindle Disassembly and Positioning in Saccharomyces cerevisiae
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spindle disassembly, establish the antiparallel bundle of microtubules known as the
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96:
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494: – Network of filamentous proteins that forms the internal framework of cells
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343:
321:
157:
1388:
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441:
432:, which can be retained within a continuous membrane. The nuclear envelopes of
380:
367:
329:
following disassembly of the highly dynamic and relatively short mitotic ones.
278:, both structurally and functionally), a protein that mechanically employs its
260:
76:
19:
16:
Final stage of a cell division for eukaryotic cells both in mitosis and meiosis
830:"The AAA-ATPase Cdc48/p97 regulates spindle disassembly at the end of mitosis"
168:
Cdk inactivation is primarily the result of the destruction of its associated
2047:
1927:
1420:
Dimitrova DS, Prokhorova TA, Blow JJ, Todorov IT, Gilbert DM (January 2002).
1030:. Rice University, Houston, Texas 77005: OpenStax College. pp. 281–283.
621:
Alberts B, Johnson A, Lewis J, Morgan D, Raff M, Roberts K, Walter P (2015).
1422:"Mammalian nuclei become licensed for DNA replication during late telophase"
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84:
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1970:
1939:
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243:
173:
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130:
120:
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and telophase are events that occur passively after satisfaction of the
1891:
1564:
346:(yeast IpI1) phosphorylates the spindle-associated stabilizing protein
312:
284:
184:-bound APC (APC/C) targets mitotic cyclins for degradation starting in
124:
112:
92:
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1133:
50:
2014:
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232:
108:
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reappear, and chromosomes begin to decondense back into the expanded
100:
88:
73:
180:(APC), also known as the cyclosome, a ubiquitin-ligase. The active,
2019:
1959:
1944:
1913:
1899:
1629:
1611:
1597:
1583:
1091:
Pollard TD, Earnshaw WC, Lippincott-Schwartz J, Johnson GT (2017).
449:
362:
The main components of the nuclear envelope are a double membrane,
248:
224:
185:
104:
80:
477:
In vertebrates, chromosome decondensation is initiated only after
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are depolymerized. Telophase accounts for approximately 2% of the
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69:
65:
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Yael A, Choi J, DeSaix J, Jurukovski V, Wisem R, Rye C (2013).
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408:
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1095:(3rd ed.). Philadelphia, PA: Elsevier. pp. 770–771.
27:
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217:
181:
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23:
This image describes the final stage in mitosis, telophase.
574:
254:
1852:
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1806:
1779:
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620:
275:
1335:
1116:
Zierhut C, Jenness C, Kimura H, Funabiki H (July 2014).
1025:
1231:
554:. London, UK: New Science Press Ltd. pp. 154–155.
1058:
689:"Spatial control of the anaphase-telophase transition"
151:
91:
and nuclear membrane disintegrating) are reversed. As
1286:
828:
Cao K, Nakajima R, Meyer HH, Zheng Y (October 2003).
827:
976:
270:
An important regulator and effector of telophase is
1370:
1172:International Review of Cell and Molecular Biology
977:Woodruff JB, Drubin DG, Barnes G (November 2010).
738:
686:
657:. Oxford, UK: Blackwell Publishing Ltd. pp.
247:which is required for the return of cells to the
2045:
1287:Lu L, Ladinsky MS, Kirchhausen T (August 2011).
1217:Symposia of the Society for Experimental Biology
64: 'appearance') is the final stage in both
1371:Taddei A, Schober H, Gasser SM (August 2010).
357:
290:
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763:
1462:
743:. New York, NY: Humana Press. pp. 3–8.
468:
55: 'end, result, completion' and
1214:
394:which is the nuclear pore scaffold protein
299:Stages of late M phase in a vertebrate cell
213:) yeast, and in multiple human cell lines.
137:between a pair of separate daughter cells.
1556:
1542:
1377:Cold Spring Harbor Perspectives in Biology
1244:Cold Spring Harbor Perspectives in Biology
1848:Cellular apoptosis susceptibility protein
1494:
1445:
1396:
1312:
1263:
1169:
1141:
1122:Nature Structural & Molecular Biology
1002:
845:
799:
789:
712:
959:
654:Cell Cycle Control and Plant Development
294:
26:
18:
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315:and return to their interphase states.
255:Additional mechanisms driving telophase
223:Historically, it has been thought that
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1237:
549:
383:laterally into a continuous membrane.
119:is disassembled and remaining spindle
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1071:. W H Freeman. pp. Section 13.4.
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140:Telophase is primarily driven by the
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687:Afonso O, Matos I, Maiato H (2014).
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274:(homologous to yeast cdc48 is human
1068:Molecular Cell Biology. 4th edition
152:Dephosphorylation of Cdk substrates
99:is re-assembled around each set of
79:. During telophase, the effects of
13:
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1075:
905:
14:
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1515:
816:
739:Monje-Casas F, Queralt E (2017).
729:
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917:International Review of Cytology
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764:Yellman CM, Roeder GS (2015).
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1:
1487:10.1080/15384101.2016.1241916
929:10.1016/S0074-7696(01)12007-3
847:10.1016/S0092-8674(03)00815-8
623:Molecular Biology of the Cell
589:10.1126/science.275.5304.1311
498:
1184:10.1016/bs.ircmb.2015.03.001
791:10.1371/journal.pone.0128604
705:10.4161/15384101.2014.959853
7:
1858:Maturation promoting factor
1389:10.1101/cshperspect.a000612
1373:"The budding yeast nucleus"
1293:The Journal of Cell Biology
1256:10.1101/cshperspect.a000539
983:The Journal of Cell Biology
485:
474:chromosome decondensation.
358:Nuclear envelope reassembly
291:Mitotic spindle disassembly
229:spindle-assembly checkpoint
172:. Cyclins are targeted for
10:
2075:
1999:Postreplication checkpoint
259:A shift in the whole-cell
178:anaphase promoting complex
2007:
1979:
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1881:
1825:
1797:
1740:
1647:
1574:
469:Chromosome decondensation
335:dephosphorylation cascade
210:Schizosaccharomyces pombe
1238:Hetzer MW (March 2010).
1050:: CS1 maint: location (
741:The Mitotic Exit Network
204:Saccharomyces cerevisiae
198:Drosophilla melanogaster
95:reach the cell poles, a
1426:Journal of Cell Science
1350:10.1242/jcs.110.13.1489
1338:Journal of Cell Science
231:(SAC) that defines the
174:proteolytic degradation
146:cyclin-dependent kinase
111:that is present during
1981:Cell cycle checkpoints
1240:"The nuclear envelope"
966:(Thesis). UC Berkeley.
364:nuclear pore complexes
300:
32:
24:
2008:Other cellular phases
1732:CDK-activating kinase
1305:10.1083/jcb.201012063
995:10.1083/jcb.201006028
373:endoplasmic reticulum
298:
30:
22:
1530:at Wikimedia Commons
1438:10.1242/jcs.115.1.51
960:Woodruff JB (2011).
883:10.1038/ncb1201-1086
871:Nature Cell Biology
782:2015PLoSO..1028604Y
313:spindle pole bodies
237:mitotic checkpoints
195:eggs, fruit flies (
1994:Spindle checkpoint
1799:P53 p63 p73 family
327:microtubule arrays
301:
148:(Cdk) substrates.
33:
25:
2041:
2040:
2037:
2036:
1989:Restriction point
1526:Media related to
1134:10.1038/nsmb.2845
1102:978-0-323-34126-4
1037:978-1-938168-09-3
668:978-1-4051-5043-9
632:978-0-8153-4432-2
561:978-0-9539181-2-6
550:Morgan D (2007).
517:978-0-321-77565-8
142:dephosphorylation
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1344:(13): 1489–502.
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583:(5304): 1311–4.
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97:nuclear envelope
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1219:(56): 193–204.
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877:(12): 1086–91.
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863:
826:
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776:(6): e0128604.
762:
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751:
737:
730:
685:
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669:
651:Inzé D (2007).
649:
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569:
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523:
510:
506:
501:
488:
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411:, and FG Nups.
360:
322:central spindle
293:
257:
207:) and fission (
158:phosphorylation
154:
117:mitotic spindle
17:
12:
11:
5:
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1516:External links
1514:
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1481:(5): 415–427.
1461:
1432:(Pt 1): 51–9.
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1383:(8): a000612.
1363:
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1250:(3): a000539.
1230:
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989:(4): 795–808.
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552:The Cell Cycle
521:
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40:Ancient Greek
37:
29:
21:
1964:
1950:Prometaphase
1478:
1474:
1464:
1429:
1425:
1415:
1380:
1376:
1366:
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1093:Cell Biology
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551:
507:
492:Cytoskeleton
476:
472:
464:
459:
458:
448:residues on
439:
433:
427:
413:
399:
385:
361:
340:
331:
320:
317:
309:kinetochores
306:
302:
258:
241:
222:
215:
208:
202:
201:), budding (
196:
189:
167:
163:phosphatases
155:
139:
129:
121:microtubules
85:prometaphase
61:
56:
48:
43:
35:
34:
1971:Cytokinesis
1940:Preprophase
1884:checkpoints
1178:: 267–330.
144:of mitotic
131:Cytokinesis
93:chromosomes
2054:Cell cycle
2048:Categories
1892:Interphase
1882:Phases and
1565:Cell cycle
1475:Cell Cycle
923:: 239–63.
693:Cell Cycle
499:References
391:importin β
285:proteasome
125:cell cycle
113:interphase
101:chromatids
74:eukaryotic
38:(from
2015:Apoptosis
1965:Telophase
1955:Metaphase
1750:INK4a/ARF
1528:Telophase
1046:cite book
420:envelope.
233:metaphase
109:chromatin
89:nucleolus
36:Telophase
1960:Anaphase
1945:Prophase
1568:proteins
1505:28135906
1456:11801723
1407:20554704
1323:21825076
1274:20300205
1225:15565882
1202:26008788
1152:24952593
1013:21079246
947:11804038
899:19261807
891:11781570
856:14636562
810:26090959
770:PLOS ONE
723:25486554
605:12265554
486:See also
377:vesicles
366:, and a
352:Kinesin8
249:G1 phase
225:anaphase
186:anaphase
105:nucleoli
81:prophase
62:(phásis)
2059:Mitosis
2029:Meiosis
1936:Mitosis
1928:M phase
1909:S phase
1776:cip/kip
1496:5351929
1447:1255924
1398:2908769
1358:9224766
1314:3153650
1265:2829960
1143:4082469
1028:Biology
1004:2983061
801:4474866
778:Bibcode
714:4614036
597:9036857
577:Science
460:Xenopus
454:lamin-A
450:lamin-B
434:Xenopus
404:histone
400:Xenopus
387:Ran-GTP
344:AuroraB
191:Xenopus
176:by the
70:mitosis
66:meiosis
1868:Cullin
1754:p14arf
1576:Cyclin
1503:
1493:
1454:
1444:
1405:
1395:
1356:
1321:
1311:
1272:
1262:
1223:
1200:
1190:
1150:
1140:
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1034:
1011:
1001:
945:
935:
897:
889:
854:
808:
798:
747:
721:
711:
665:
659:99–103
629:
603:
595:
558:
515:
409:POM121
280:ATPase
170:cyclin
135:nuclei
115:. The
103:, the
2024:phase
1918:phase
1904:phase
1843:Cdc42
1838:Cdc25
1826:Other
1608:, B3)
895:S2CID
601:S2CID
430:lamin
272:cdc48
218:cdc14
182:CDC20
87:(the
72:in a
59:φάσις
51:télos
46:τέλος
42:
1872:CUL7
1833:Cdc2
1501:PMID
1452:PMID
1403:PMID
1354:PMID
1319:PMID
1270:PMID
1221:PMID
1198:PMID
1188:ISBN
1148:PMID
1097:ISBN
1052:link
1032:ISBN
1009:PMID
943:PMID
933:ISBN
887:PMID
852:PMID
834:Cell
806:PMID
745:ISBN
719:PMID
663:ISBN
627:ISBN
593:PMID
556:ISBN
513:ISBN
446:COOH
396:ELYS
381:fuse
311:and
244:CDH1
156:The
83:and
77:cell
68:and
1863:Wee
1853:E2F
1817:p73
1812:p63
1807:p53
1788:p57
1784:p27
1780:p21
1770:p19
1766:p18
1762:p15
1758:p16
1712:11B
1707:11A
1649:CDK
1491:PMC
1483:doi
1442:PMC
1434:doi
1430:115
1393:PMC
1385:doi
1346:doi
1342:110
1309:PMC
1301:doi
1297:194
1260:PMC
1252:doi
1180:doi
1176:317
1138:PMC
1130:doi
999:PMC
991:doi
987:191
925:doi
921:212
879:doi
842:doi
838:115
796:PMC
786:doi
709:PMC
701:doi
585:doi
581:275
444:of
348:EB1
276:p97
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1782:,
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1636:,
1634:E1
1624:D3
1622:,
1620:D2
1618:,
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Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.
