Mitotic Prometaphase (Homo sapiens)

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3, 10, 2120, 274, 18, 25, 27271-3, 5, 10...6, 8, 17, 19, 294, 12, 14, 15, 18...5, 215, 21193, 10, 133, 7, 9, 13, 16...32cytosolRANBP2 p-S454-RAD21 Ac-Cohesin:PDS5:CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesKNTC1 ADPPDS5B PLK1 PDS5A PDS5A TAOK1 NDEL1 CENPN CDC20 SMC1A 2xAcK-SMC3 CENPI NSL1 ZWINT WAPAL AURKB NCAPD2 MLF1IP CENPP SisterChromosomalArms:p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPALSPC25 ATPRCC2 APITD1 microtubuleKIF18A CENPO CENPP B9D2 p-T1339,T1384,T1389-NCAPD2 MAD2L1 KIF18A APITD1 SMC1A CoA-SHSPC24 DSN1 CENPM NDC80 H2ONUP133 CENPN CASC5 BUB1 CENPI NDEL1 MAD2L1 CASC5 CENPK 2xAcK-SMC3 ERCC6L PAFAH1B1 STAG1 SPC25 RPS27 SPC25 CSNK2B ADPSTAG1 BIRC5 KIF2C CENPP CENPL CASC5 SEC13 MAPRE1 p-S21,S75,T159-CDCA5CASC5 NUP37 WAPAL CLASP1 SGOL1 KIF18A SMC1A p-T161-CDK1 RCC2 CLIP1 p-STAG2 KIF2A NUP160 ITGB3BP SGOL2 CENPO KIF2C NDE1 PiCENPF NUP85 BUB3 ZWILCH CENPA PMF1 RANGAP1 2xAcK-SMC3 Condensed prophasechromosomesCENPH ATPRPS27 SKA1 STAG2 BUB3 NUP43 CENPK CENPA CENPQ XPO1 WAPAL NUP85 ZWILCH CENPE NSL1 WAPAL RANGAP1 SEH1L-1 PPP1CC 2xAcK-SMC3 NUP43 CLIP1 SGOL2 SMC1A DSN1 SGOL2 ATPAHCTF1 CENPI NDC80 NUP85 HDAC8MAD2L1 STAG1 MAPRE1 KIF2B SPC25 SGOL2 SKA1 MIS12 PDS5B PDS5A ZWINT CDCA8 p-RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesSKA2 KNTC1 WAPAL SGOL1 SKA1 ZW10 CASC5 CLASP2 NUP98-5 CENPT CENPK CCNB:p-T161-CDK1INCENP p-Cohesin:PDS5:WAPALSEH1L-1 PPP1CC SPC25 MLF1IP B9D2 NUP85 SGOL2 SMC2 PPP1CC SGOL1 SKA1 RCC2 CDC20 2xAcK-SMC3 CENPE NDE1 MIS12 p-S21,S75,T159-CDCA5p-S454-RAD21 AURKB SKA2 PiKinetochoreCENPQ CLIP1 CLASP2 NUP98-5 ATPCENPL AURKB CENPO p-S454-RAD21 B9D2 CENPM SPC24 MAPRE1 PDS5B KIF2A CondensedprometaphasechromosomesSMC4 STAG1 NUDC MIS12 SPDL1 KIF2A NUP98-5 SGOL1 NUP37 SPDL1 MAD2L1 INCENP PDS5A SisterChromosomalArms:Ac-Cohesin:PDS5:CDCA5:WAPALCENPE PDS5A NUF2 XPO1 NUP107 NUP107 CENPQ CASC5 CENPK KIF18A SPC25 BIRC5 S PhasePMF1 CENPC1 CENPO CENPC1 ITGB3BP RANBP2 PPP1CC MAD1L1 ATPCENPA CENPH SPDL1 NUP98-5 NCAPH KIF2B BIRC5 RANGAP1 NUP107 CENPN RCC2 NUDC CENPA RAD21 TAOK1 NUF2 MIS12 INCENP NSL1 AURKB PAFAH1B1 SKA2 NDE1 APITD1 NUP160 CKAP5 PPP1CC CLASP2 KIF2B STAG1 CLIP1 XPO1 RCC2 CK2 PhosphorylatedCondensin IZW10 PDS5B CENPE PDS5B STAG1 ERCC6L SGOL1 SMC2 SKA2 NUP160 CENPT SPC24 PMF1 PDS5B 2xAcK-SMC3 NUP133 CKAP5 MLF1IP BUB1B KIF2A p-S-NCAPG NUP133 NDC80 STAG1 SKA2 NUP160 NUDC 2xAcK-SMC3 CENPI KIF2C STAG2 XPO1 CENPL TAOK1 SMC4 NDE1 p-S454-RAD21 APITD1 CDK1 PhosphorylatedCondensin IWAPAL NDC80 SMC1A Condensin IRANGAP1 NUDC CENPP ERCC6L NUP37 BUB3 CDCA8 NUP85 PAFAH1B1 SPDL1 CENPM PLK1 PDS5B PDS5A NUP133 CLASP1 MLF1IP CENPL ZWINT CLIP1 SMC1A ITGB3BP STAG1 CENPH NUP133 SPC24 CDCA5 ZWILCH BUB1B CENPC1 CENPQ CENPO NUP107 RAD21 BIRC5 B9D2 CDCA8 CDCA5 SEH1L-1 ERCC6L CENPF CLASP1 ITGB3BP SMC1A CLASP1 RAD21 STAG1 CENPF CENPM CLASP1 AURKB CENPA PDS5B MAD2L1 NUF2 SPDL1 CENPM SEH1L-1 B9D2 KIF18A CENPF NUDC MAD1L1 DSN1 BUB1B KNTC1 SEC13 PDS5A CENPE ZW10 SEC13 2xAcK-SMC3 NDEL1 RPS27 NDC80 CDC20 CENPP CDCA8 KIF2B KIF18A NUP107 SPC24 DSN1 AHCTF1 MAD2L1 STAG2 KNTC1 PMF1 SisterChromosomalArms:Ac-Cohesin:PDS5:p-CDCA5:WAPALCLIP1 RANBP2 NUP160 NUP43 SGOL2 CENPF KNTC1 NUP85 MLF1IP CDCA5 PLK1 NUF2 2xAcK-SMC3 CENPQ CLASP1 SMC3 XPO1 SEC13 ZWILCH ZWINT RCC2 NDE1 SPC24 TAOK1 BUB1B B9D2 STAG2 NUP107 Microtubule-boundkinetochoreMIS12 TAOK1 ADPMAD1L1 AHCTF1 SMC1A H2OCENPA NDEL1 p-T308,T332-NCAPG NUP98-5 CENPT CCNB1 BUB3 PLK1 PMF1 PLK1 NUP98-5 CENPP SEC13 BUB1 CDCA8 RPS27 SPDL1 KIF2B MAD1L1 NDEL1 PLK1CENPH NSL1 CENPC1 p-S21,S75,T159-CDCA5SEH1L-1 BUB3 NUP43 AHCTF1 CLASP2 Ac-CoAMAD1L1 XPO1 NDC80 CENPT p-S21,S75,T159-CDCA5KIF2A Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubulesp-S-NCAPD2 RPS27 CENPN PAFAH1B1 DSN1 PAFAH1B1 BUB3 p-S570-NCAPH ADPBUB1 CENPK ZW10 CENPM SKA1 NUF2 NCAPG ERCC6L BUB1 SKA2 MAPRE1 INCENP CENPO ZW10 p-S21,S75,T159-CDCA5MIS12 p-S454-RAD21 PDS5A Sister ChromosomalArmITGB3BP ZWINT CENPI NUF2 CENPF NUP37 PPP1CC CENPE KIF2C BIRC5 RAD21 CENPQ BUB1 SEH1L-1 RAD21 CKAP5 SisterCentromeres:Ac-Cohesin:PDS5:CDCA5:WAPALSMC1A CKAP5 INCENP WAPAL PAFAH1B1 KIF2C DSN1 PDS5A STAG2 CENPI NSL1 AHCTF1 PDS5A ADPNUP160 KIF2B ZWILCH CENPN CENPC1 CDC20 SKA1 CENPL PLK1 CENPT AURKB SGOL1 RANBP2 CDC20 WAPAL CDC20 RANBP2 CENPK CSNK2A1 p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesCDCA8 NUDC p-S21,S75,T159-CDCA5p-STAG2 ATPNUP43 KNTC1 PDS5B CENPC1 p-Ac-Cohesin:PDS5:WAPALCENPL SEC13 ITGB3BP KIF2A BUB1B INCENP WAPAL BIRC5 NSL1 BUB1B SMC1A NUP43 MAD1L1 p-T-NCAPH PDS5B CENPH p-S-SMC4 CENPT CLASP2 MAPRE1 ZWILCH CENPN ADPp-STAG2 CKAP5 NUP37 KIF2C BUB1 Unknown PhosphataseNUP37 MLF1IP SMC2 Mitotic Metaphaseand AnaphaseSTAG1 RANBP2 APITD1 AHCTF1 TAOK1 RANGAP1 WAPAL STAG2 RPS27 RANGAP1 PMF1 NDE1 NUP133 NDEL1 p-STAG2 CCNB2 CENPH ZWINT ZW10 CSNK2A2 CKAP5 CLASP2 Casein kinase IIAPITD1 ERCC6L MAPRE1 44


Description

The dissolution of the nuclear membrane marks the beginning of the prometaphase. Kinetochores are created when proteins attach to the centromeres. Microtubules then attach at the kinetochores, and the chromosomes begin to move to the metaphase plate. Source:Reactome.

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Bibliography

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  28. Liu ST, Rattner JB, Jablonski SA, Yen TJ.; ''Mapping the assembly pathways that specify formation of the trilaminar kinetochore plates in human cells.''; PubMed Europe PMC Scholia
  29. Kitajima TS, Hauf S, Ohsugi M, Yamamoto T, Watanabe Y.; ''Human Bub1 defines the persistent cohesion site along the mitotic chromosome by affecting Shugoshin localization.''; PubMed Europe PMC Scholia
  30. Takemoto A, Kimura K, Yokoyama S, Hanaoka F.; ''Cell cycle-dependent phosphorylation, nuclear localization, and activation of human condensin.''; PubMed Europe PMC Scholia
  31. Deardorff MA, Bando M, Nakato R, Watrin E, Itoh T, Minamino M, Saitoh K, Komata M, Katou Y, Clark D, Cole KE, De Baere E, Decroos C, Di Donato N, Ernst S, Francey LJ, Gyftodimou Y, Hirashima K, Hullings M, Ishikawa Y, Jaulin C, Kaur M, Kiyono T, Lombardi PM, Magnaghi-Jaulin L, Mortier GR, Nozaki N, Petersen MB, Seimiya H, Siu VM, Suzuki Y, Takagaki K, Wilde JJ, Willems PJ, Prigent C, Gillessen-Kaesbach G, Christianson DW, Kaiser FJ, Jackson LG, Hirota T, Krantz ID, Shirahige K.; ''HDAC8 mutations in Cornelia de Lange syndrome affect the cohesin acetylation cycle.''; PubMed Europe PMC Scholia
  32. Kawashima SA, Yamagishi Y, Honda T, Ishiguro K, Watanabe Y.; ''Phosphorylation of H2A by Bub1 prevents chromosomal instability through localizing shugoshin.''; PubMed Europe PMC Scholia
  33. Haren L, Merdes A.; ''Direct binding of NuMA to tubulin is mediated by a novel sequence motif in the tail domain that bundles and stabilizes microtubules.''; PubMed Europe PMC Scholia
  34. Gandhi R, Gillespie PJ, Hirano T.; ''Human Wapl is a cohesin-binding protein that promotes sister-chromatid resolution in mitotic prophase.''; PubMed Europe PMC Scholia
  35. Foltz DR, Jansen LE, Black BE, Bailey AO, Yates JR, Cleveland DW.; ''The human CENP-A centromeric nucleosome-associated complex.''; PubMed Europe PMC Scholia
  36. Cheeseman IM, Chappie JS, Wilson-Kubalek EM, Desai A.; ''The conserved KMN network constitutes the core microtubule-binding site of the kinetochore.''; PubMed Europe PMC Scholia
  37. Murphy LA, Sarge KD.; ''Phosphorylation of CAP-G is required for its chromosomal DNA localization during mitosis.''; PubMed Europe PMC Scholia
  38. Kitajima TS, Kawashima SA, Watanabe Y.; ''The conserved kinetochore protein shugoshin protects centromeric cohesion during meiosis.''; PubMed Europe PMC Scholia
  39. Adib R, Montgomery JM, Atherton J, O'Regan L, Richards MW, Straatman KR, Roth D, Straube A, Bayliss R, Moores CA, Fry AM.; ''Mitotic phosphorylation by NEK6 and NEK7 reduces the microtubule affinity of EML4 to promote chromosome congression.''; PubMed Europe PMC Scholia
  40. Pollmann M, Parwaresch R, Adam-Klages S, Kruse ML, Buck F, Heidebrecht HJ.; ''Human EML4, a novel member of the EMAP family, is essential for microtubule formation.''; PubMed Europe PMC Scholia
  41. Yamagishi Y, Sakuno T, Shimura M, Watanabe Y.; ''Heterochromatin links to centromeric protection by recruiting shugoshin.''; PubMed Europe PMC Scholia
  42. Hauf S, Roitinger E, Koch B, Dittrich CM, Mechtler K, Peters JM.; ''Dissociation of cohesin from chromosome arms and loss of arm cohesion during early mitosis depends on phosphorylation of SA2.''; PubMed Europe PMC Scholia
  43. Hirota T, Gerlich D, Koch B, Ellenberg J, Peters JM.; ''Distinct functions of condensin I and II in mitotic chromosome assembly.''; PubMed Europe PMC Scholia
  44. Aumais JP, Williams SN, Luo W, Nishino M, Caldwell KA, Caldwell GA, Lin SH, Yu-Lee LY.; ''Role for NudC, a dynein-associated nuclear movement protein, in mitosis and cytokinesis.''; PubMed Europe PMC Scholia
  45. Dreier MR, Bekier ME, Taylor WR.; ''Regulation of sororin by Cdk1-mediated phosphorylation.''; PubMed Europe PMC Scholia

History

View all...
CompareRevisionActionTimeUserComment
114617view16:07, 25 January 2021ReactomeTeamReactome version 75
113066view11:12, 2 November 2020ReactomeTeamReactome version 74
112301view15:21, 9 October 2020ReactomeTeamReactome version 73
101199view11:09, 1 November 2018ReactomeTeamreactome version 66
100737view20:33, 31 October 2018ReactomeTeamreactome version 65
100281view19:11, 31 October 2018ReactomeTeamreactome version 64
99827view15:54, 31 October 2018ReactomeTeamreactome version 63
99384view14:32, 31 October 2018ReactomeTeamreactome version 62 (2nd attempt)
99083view12:39, 31 October 2018ReactomeTeamreactome version 62
93970view13:48, 16 August 2017ReactomeTeamreactome version 61
93570view11:27, 9 August 2017ReactomeTeamreactome version 61
87985view13:22, 25 July 2016RyanmillerOntology Term : 'cell cycle pathway, mitotic' added !
87971view13:17, 25 July 2016RyanmillerOntology Term : 'regulatory pathway' added !
86672view09:23, 11 July 2016ReactomeTeamreactome version 56
83244view10:29, 18 November 2015ReactomeTeamVersion54
81349view12:52, 21 August 2015ReactomeTeamVersion53
76819view08:04, 17 July 2014ReactomeTeamFixed remaining interactions
76523view11:45, 16 July 2014ReactomeTeamFixed remaining interactions
75856view09:50, 11 June 2014ReactomeTeamRe-fixing comment source
75556view10:35, 10 June 2014ReactomeTeamReactome 48 Update
74911view13:44, 8 May 2014AnweshaFixing comment source for displaying WikiPathways description
74555view08:35, 30 April 2014ReactomeTeamNew pathway

External references

DataNodes

View all...
NameTypeDatabase referenceComment
2xAcK-SMC3 ProteinQ9UQE7 (Uniprot-TrEMBL)
ADPMetaboliteCHEBI:16761 (ChEBI)
AHCTF1 ProteinQ8WYP5 (Uniprot-TrEMBL)
APITD1 ProteinQ8N2Z9 (Uniprot-TrEMBL)
ATPMetaboliteCHEBI:15422 (ChEBI)
AURKB ProteinQ96GD4 (Uniprot-TrEMBL)
Ac-CoAMetaboliteCHEBI:15351 (ChEBI)
Ac-Cohesin:PDS5:CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesComplexR-HSA-2484819 (Reactome)
Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesComplexR-HSA-2468282 (Reactome)
B9D2 ProteinQ9BPU9 (Uniprot-TrEMBL)
BIRC5 ProteinO15392 (Uniprot-TrEMBL)
BUB1 ProteinO43683 (Uniprot-TrEMBL)
BUB1B ProteinO60566 (Uniprot-TrEMBL)
BUB3 ProteinO43684 (Uniprot-TrEMBL)
CASC5 ProteinQ8NG31 (Uniprot-TrEMBL)
CCNB1 ProteinP14635 (Uniprot-TrEMBL)
CCNB2 ProteinO95067 (Uniprot-TrEMBL)
CCNB:p-T161-CDK1ComplexR-HSA-2311324 (Reactome)
CDC20 ProteinQ12834 (Uniprot-TrEMBL)
CDCA5 ProteinQ96FF9 (Uniprot-TrEMBL)
CDCA8 ProteinQ53HL2 (Uniprot-TrEMBL)
CDK1 Phosphorylated Condensin IComplexR-HSA-2520845 (Reactome)
CENPA ProteinP49450 (Uniprot-TrEMBL)
CENPC1 ProteinQ03188 (Uniprot-TrEMBL)
CENPE ProteinQ02224 (Uniprot-TrEMBL)
CENPF ProteinP49454 (Uniprot-TrEMBL)
CENPH ProteinQ9H3R5 (Uniprot-TrEMBL)
CENPI ProteinQ92674 (Uniprot-TrEMBL)
CENPK ProteinQ9BS16 (Uniprot-TrEMBL)
CENPL ProteinQ8N0S6 (Uniprot-TrEMBL)
CENPM ProteinQ9NSP4 (Uniprot-TrEMBL)
CENPN ProteinQ96H22 (Uniprot-TrEMBL)
CENPO ProteinQ9BU64 (Uniprot-TrEMBL)
CENPP ProteinQ6IPU0 (Uniprot-TrEMBL)
CENPQ ProteinQ7L2Z9 (Uniprot-TrEMBL)
CENPT ProteinQ96BT3 (Uniprot-TrEMBL)
CK2 Phosphorylated Condensin IComplexR-HSA-2529014 (Reactome)
CKAP5 ProteinQ14008 (Uniprot-TrEMBL)
CLASP1 ProteinQ7Z460 (Uniprot-TrEMBL)
CLASP2 ProteinO75122 (Uniprot-TrEMBL)
CLIP1 ProteinP30622 (Uniprot-TrEMBL)
CSNK2A1 ProteinP68400 (Uniprot-TrEMBL)
CSNK2A2 ProteinP19784 (Uniprot-TrEMBL)
CSNK2B ProteinP67870 (Uniprot-TrEMBL)
Casein kinase IIComplexR-HSA-201711 (Reactome)
CoA-SHMetaboliteCHEBI:15346 (ChEBI)
Condensed

prometaphase

chromosomes		R-NUL-2520884 (Reactome)
Condensed prophase chromosomesR-NUL-2520882 (Reactome)
Condensin IComplexR-HSA-1638143 (Reactome)
DSN1 ProteinQ9H410 (Uniprot-TrEMBL)
ERCC6L ProteinQ2NKX8 (Uniprot-TrEMBL)
H2OMetaboliteCHEBI:15377 (ChEBI)
HDAC8ProteinR-HSA-2545205 (Reactome)
INCENP ProteinQ9NQS7 (Uniprot-TrEMBL)
ITGB3BP ProteinQ13352 (Uniprot-TrEMBL)
KIF18A ProteinQ8NI77 (Uniprot-TrEMBL)
KIF2A ProteinO00139 (Uniprot-TrEMBL)
KIF2B ProteinQ8N4N8 (Uniprot-TrEMBL)
KIF2C ProteinQ99661 (Uniprot-TrEMBL)
KNTC1 ProteinP50748 (Uniprot-TrEMBL)
KinetochoreComplexR-HSA-375305 (Reactome)
MAD1L1 ProteinQ9Y6D9 (Uniprot-TrEMBL)
MAD2L1 ProteinQ13257 (Uniprot-TrEMBL)
MAPRE1 ProteinQ15691 (Uniprot-TrEMBL)
MIS12 ProteinQ9H081 (Uniprot-TrEMBL)
MLF1IP ProteinQ71F23 (Uniprot-TrEMBL)
Microtubule-bound kinetochoreComplexR-HSA-375303 (Reactome)
Mitotic Metaphase and AnaphasePathwayR-HSA-2555396 (Reactome) Metaphase is marked by the formation of the metaphase plate. The metaphase plate is formed when the spindle fibers align the chromosomes along the middle of the cell. Such an organization helps to ensure that later, when the chromosomes are separated, each new nucleus that is formed receives one copy of each chromosome. This pathway has not yet been annotated in Reactome.

The metaphase to anaphase transition during mitosis is triggered by the destruction of mitotic cyclins.

In anaphase, the paired chromosomes separate at the centromeres, and move to the opposite sides of the cell. The movement of the chromosomes is facilitated by a combination of kinetochore movement along the spindle microtubules and through the physical interaction of polar microtubules.
NCAPD2 ProteinQ15021 (Uniprot-TrEMBL)
NCAPG ProteinQ9BPX3 (Uniprot-TrEMBL)
NCAPH ProteinQ15003 (Uniprot-TrEMBL)
NDC80 ProteinO14777 (Uniprot-TrEMBL)
NDE1 ProteinQ9NXR1 (Uniprot-TrEMBL)
NDEL1 ProteinQ9GZM8 (Uniprot-TrEMBL)
NSL1 ProteinQ96IY1 (Uniprot-TrEMBL)
NUDC ProteinQ9Y266 (Uniprot-TrEMBL)
NUF2 ProteinQ9BZD4 (Uniprot-TrEMBL)
NUP107 ProteinP57740 (Uniprot-TrEMBL)
NUP133 ProteinQ8WUM0 (Uniprot-TrEMBL)
NUP160 ProteinQ12769 (Uniprot-TrEMBL)
NUP37 ProteinQ8NFH4 (Uniprot-TrEMBL)
NUP43 ProteinQ8NFH3 (Uniprot-TrEMBL)
NUP85 ProteinQ9BW27 (Uniprot-TrEMBL)
NUP98-5 ProteinP52948-5 (Uniprot-TrEMBL)
PAFAH1B1 ProteinP43034 (Uniprot-TrEMBL)
PDS5A ProteinQ29RF7 (Uniprot-TrEMBL)
PDS5B ProteinQ9NTI5 (Uniprot-TrEMBL)
PLK1 ProteinP53350 (Uniprot-TrEMBL)
PLK1ProteinP53350 (Uniprot-TrEMBL)
PMF1 ProteinQ6P1K2 (Uniprot-TrEMBL)
PPP1CC ProteinP36873 (Uniprot-TrEMBL)
PiMetaboliteCHEBI:18367 (ChEBI)
RAD21 ProteinO60216 (Uniprot-TrEMBL)
RANBP2 ProteinP49792 (Uniprot-TrEMBL)
RANGAP1 ProteinP46060 (Uniprot-TrEMBL)
RCC2 ProteinQ9P258 (Uniprot-TrEMBL)
RPS27 ProteinP42677 (Uniprot-TrEMBL)
S PhasePathwayR-HSA-69242 (Reactome) DNA synthesis occurs in the S phase, or the synthesis phase, of the cell cycle. The cell duplicates its hereditary material, and two copies of the chromosome are formed. As DNA replication continues, the E type cyclins shared by the G1 and S phases, are destroyed and the levels of the mitotic cyclins rise.
SEC13 ProteinP55735 (Uniprot-TrEMBL)
SEH1L-1 ProteinQ96EE3-1 (Uniprot-TrEMBL)
SGOL1 ProteinQ5FBB7 (Uniprot-TrEMBL)
SGOL2 ProteinQ562F6 (Uniprot-TrEMBL)
SKA1 ProteinQ96BD8 (Uniprot-TrEMBL)
SKA2 ProteinQ8WVK7 (Uniprot-TrEMBL)
SMC1A ProteinQ14683 (Uniprot-TrEMBL)
SMC2 ProteinO95347 (Uniprot-TrEMBL)
SMC3 ProteinQ9UQE7 (Uniprot-TrEMBL)
SMC4 ProteinQ9NTJ3 (Uniprot-TrEMBL)
SPC24 ProteinQ8NBT2 (Uniprot-TrEMBL)
SPC25 ProteinQ9HBM1 (Uniprot-TrEMBL)
SPDL1 ProteinQ96EA4 (Uniprot-TrEMBL)
STAG1 ProteinQ8WVM7 (Uniprot-TrEMBL)
STAG2 ProteinQ8N3U4 (Uniprot-TrEMBL)
Sister Centromeres:Ac-Cohesin:PDS5:CDCA5:WAPALComplexR-HSA-1638799 (Reactome)
Sister

Chromosomal

Arms:Ac-Cohesin:PDS5:CDCA5:WAPAL		ComplexR-HSA-1638802 (Reactome)
Sister

Chromosomal

Arms:Ac-Cohesin:PDS5:p-CDCA5:WAPAL		ComplexR-HSA-2468274 (Reactome)
Sister

Chromosomal

Arms:p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL		ComplexR-HSA-1638805 (Reactome)
Sister Chromosomal ArmR-NUL-1638790 (Reactome)
TAOK1 ProteinQ7L7X3 (Uniprot-TrEMBL)
Unknown PhosphataseR-NUL-2529006 (Reactome)
WAPAL ProteinQ7Z5K2 (Uniprot-TrEMBL)
XPO1 ProteinO14980 (Uniprot-TrEMBL)
ZW10 ProteinO43264 (Uniprot-TrEMBL)
ZWILCH ProteinQ9H900 (Uniprot-TrEMBL)
ZWINT ProteinO95229 (Uniprot-TrEMBL)
microtubuleR-HSA-190599 (Reactome)
p-Ac-Cohesin:PDS5:WAPALComplexR-HSA-2484812 (Reactome)
p-Cohesin:PDS5:WAPALComplexR-HSA-2545249 (Reactome)
p-RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesComplexR-HSA-2500242 (Reactome)
p-S-NCAPD2 ProteinQ15021 (Uniprot-TrEMBL)
p-S-NCAPG ProteinQ9BPX3 (Uniprot-TrEMBL)
p-S-SMC4 ProteinQ9NTJ3 (Uniprot-TrEMBL)
p-S21,S75,T159-CDCA5 ProteinQ96FF9 (Uniprot-TrEMBL)
p-S21,S75,T159-CDCA5ProteinQ96FF9 (Uniprot-TrEMBL)
p-S454-RAD21 ProteinO60216 (Uniprot-TrEMBL)
p-S570-NCAPH ProteinQ15003 (Uniprot-TrEMBL)
p-STAG2 ProteinQ8N3U4 (Uniprot-TrEMBL)
p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesComplexR-HSA-1638796 (Reactome)
p-T-NCAPH ProteinQ15003 (Uniprot-TrEMBL)
p-T1339,T1384,T1389-NCAPD2 ProteinQ15021 (Uniprot-TrEMBL)
p-T161-CDK1 ProteinP06493 (Uniprot-TrEMBL)
p-T308,T332-NCAPG ProteinQ9BPX3 (Uniprot-TrEMBL)

Annotated Interactions

View all...
SourceTargetTypeDatabase referenceComment
ADPArrowR-HSA-1638803 (Reactome)
ADPArrowR-HSA-2466068 (Reactome)
ADPArrowR-HSA-2468287 (Reactome)
ADPArrowR-HSA-2468293 (Reactome)
ADPArrowR-HSA-2514854 (Reactome)
ADPArrowR-HSA-2529020 (Reactome)
ATPR-HSA-1638803 (Reactome)
ATPR-HSA-2466068 (Reactome)
ATPR-HSA-2468287 (Reactome)
ATPR-HSA-2468293 (Reactome)
ATPR-HSA-2514854 (Reactome)
ATPR-HSA-2529020 (Reactome)
Ac-CoAArrowR-HSA-2545253 (Reactome)
Ac-Cohesin:PDS5:CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesArrowR-HSA-2484822 (Reactome)
Ac-Cohesin:PDS5:CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesR-HSA-2468287 (Reactome)
Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesArrowR-HSA-2468287 (Reactome)
Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesR-HSA-1638803 (Reactome)
CCNB:p-T161-CDK1mim-catalysisR-HSA-2468287 (Reactome)
CCNB:p-T161-CDK1mim-catalysisR-HSA-2468293 (Reactome)
CCNB:p-T161-CDK1mim-catalysisR-HSA-2514854 (Reactome)
CDK1 Phosphorylated Condensin IArrowR-HSA-2514854 (Reactome)
CDK1 Phosphorylated Condensin IArrowR-HSA-2520883 (Reactome)
CK2 Phosphorylated Condensin IArrowR-HSA-2529020 (Reactome)
CK2 Phosphorylated Condensin IR-HSA-2529015 (Reactome)
Casein kinase IImim-catalysisR-HSA-2529020 (Reactome)
CoA-SHR-HSA-2545253 (Reactome)
Condensed

prometaphase

chromosomes		ArrowR-HSA-2520883 (Reactome)
Condensed prophase chromosomesR-HSA-2520883 (Reactome)
Condensin IArrowR-HSA-2529015 (Reactome)
Condensin IR-HSA-2514854 (Reactome)
Condensin IR-HSA-2529020 (Reactome)
H2OR-HSA-1638821 (Reactome)
H2OR-HSA-2529015 (Reactome)
HDAC8mim-catalysisR-HSA-2545253 (Reactome)
KinetochoreR-HSA-375302 (Reactome)
Microtubule-bound kinetochoreArrowR-HSA-375302 (Reactome)
Microtubule-bound kinetochoreR-HSA-2484822 (Reactome)
PLK1mim-catalysisR-HSA-1638803 (Reactome)
PLK1mim-catalysisR-HSA-2466068 (Reactome)
PiArrowR-HSA-1638821 (Reactome)
PiArrowR-HSA-2529015 (Reactome)
R-HSA-1638803 (Reactome) Prior to anaphase onset, sister-chromatids are held together by cohesin complexes distributed along chromosomal arms and at centromeres. In prometaphase, PLK1, likely recruited to cohesin complexes by binding to CDK1-phosphorylated CDCA5 (Sororin) (Zhang et al. 2011), phosphorylates cohesin subunits STAG2 (SA2) and RAD21 (Hauf et al. 2005). PLK1-mediated phosphorylation of cohesin subunits at centromeres is counteracted by the phosphatase activity of PP2A complex (containing the regulatory subunit B56 i.e. PPP2R5), which is recruited to the kinetochore by shugoshin proteins, SGOL1 and SGOL2 (Kitajima et al. 2006). Therefore, while cohesin complexes dissociate from chromosomal arms in prometaphase (Hauf et al. 2001), they remain bound to centromeres until anaphase onset (Hauf et al. 2001, Hauf et al. 2005, Kitajima et al. 2006). When separase is activated after its inhibitor securin is degraded by APC/C at the onset of anaphase, RAD21 is cleaved by separase. Phosphorylation of RAD21 by PLK1 facilitates subsequent cleavage of RAD21 by separase (Hauf et al. 2005). There are several potential PLK1 phosphorylation sites in STAG2 and RAD21, but the exact positions of in vivo phosphorylation of STAG2 and RAD21 by PLK1 have not been explicitly established (Hauf et al. 2005).
R-HSA-1638821 (Reactome) PLK1-mediated phosphorylation of the STAG2 subunit of centromeric cohesin (Hauf et al. 2005) is counteracted by the kinetochore PP2A phosphatase, containing the 56 kDa regulatory B subunit (PP2A-B56 i.e. PP2A-PPP2R5). PP2A-B56 is recruited to the centromeric cohesin complex by shugoshin proteins (SGOL1 and SGOL2) (Kitajima et al. 2006), which are also kinetochore constituents (Cheeseman and Desai 2008). SGOL1 localization to centromeres is sustained by the interaction with histone H2A possessing the phosphorylation of T120 which is introduced by the protein kinase BUB1, and heterochromatin protein HP1 (Kitajima et al. 2005, Kawashima et al. 2010, Yamagishi et al. 2008). Shugoshin- and PP2A-B56-regulated dephosphorylation of centromeric STAG2 ensures that the cohesin complex remains bound to centromeres throughout prometaphase and metaphase, thereby preventing premature separation of sister chromatids (Salic et al. 2004, Kitajima et al. 2004, Kitajima et al. 2005, Kitajima et al. 2006).
R-HSA-2466068 (Reactome) Prior to anaphase onset, sister-chromatids are held together by cohesin complexes. PLK1-dependent phosphorylation of the cohesin subunit STAG2 (SA2) (Hauf et al. 2005) promotes dissociation of cohesins from chromosomal arms in prometaphase (Hauf et al. 2001). Besides phosphorylating STAG2, PLK1 also phosphorylates RAD21 cohesin subunit, but the phosphorylation of RAD21 is not required for the dissociation of cohesin from chromosomal arms in early mitosis (Hauf et al. 2005). There are several potential PLK1 phosphorylation sites in STAG2 and RAD21, but the exact positions of in vivo phosphorylation of STAG2 and RAD21 by PLK1 have not been explicitly established (Hauf et al. 2005). It is likely that the phosphorylation of cohesin-bound CDCA5 (Sororin) by CDK1 creates a docking site for PLK1 at threonine T159 of CDCA5, thus enabling PLK1 to phosphorylate cohesin subunits (Zhang et al. 2011).
R-HSA-2467794 (Reactome) Cohesin complexes dissociate from chromosomal arms in prometaphase, leading to sister chromatid resolution. Sister chromatid resolution involves separation of sister chromosomal arms while cohesion at sister centromeres persists (Losada et al. 1998, Hauf et al. 2001, Hauf et al. 2005). Cohesin and CDCA5 (Sororin) simultaneously dissociate from chromosomal arms in prometaphase (Nishiyama et al. 2010, Zhang et al. 2011). This process, triggered by CDK1-mediated phosphorylation of CDCA5 (Dreier et al. 2011, Zhang et al. 2011) and PLK1-mediated phosphorylation of the STAG2 cohesin subunit (Hauf et al. 2005), is controlled by WAPAL (Gandhi et al. 2006, Kueng et al. 2006, Shintomi and Hirano 2009). WAPAL controls cohesion of sister chromatids likely through competing with CDCA5 for binding to cohesin-associated PDS5 (PDS5A and PDS5B) (Nishiyama et al. 2010). While the interaction of WAPAL with PDS5 depends on CDCA5 (Nishiyama et al. 2010), WAPAL maintains its association with cohesin through interaction with cohesin subunits (Kueng et al. 2006, Shintomi and Hirano 2009).
R-HSA-2468287 (Reactome) Phosphorylation of CDCA5 (Sororin) coincides with dissociation of CDCA5 from chromosomal arms in prometaphase, but phosphorylated CDCA5 persists on centromeres throughout prophase and metaphase. Several serine and threonine residues in CDCA5 are phosphorylated by CDK1 in prometaphase, but only the three sites that perfectly match the CDK1 consensus phosphorylation sequence are shown here - serines S21 and S75 and threonine T159 (Drier et al. 2011, Zhang et al. 2011).
R-HSA-2468293 (Reactome) Phosphorylation of CDCA5 (Sororin) coincides with dissociation of CDCA5 from chromosomal arms in prometaphase. Several serine and threonine residues in CDCA5 are phosphorylated by CDK1 in prometaphase, but only the three sites that perfectly match the CDK1 consensus phosphorylation sequence are shown here - serines S21 and S75 and threonine T159 (Drier et al. 2011, Zhang et al. 2011).
R-HSA-2484822 (Reactome) The kinetochore assembly on centromeres of replicated chromosomes is completed by mitotic prometaphase. Some kinetochore components are associated with centromeres throughout the cell cycle while others associate with centromeres during mitosis. The sequential kinetochore assembly and kinetochore dynamics is not shown here. For a review of this process, please refer to Cheeseman and Desai 2008.
R-HSA-2514854 (Reactome) CDK1 (CDC2) in complex with CCNB (cyclin B) phosphorylates condensin I subunits NCAPD2, NCAPG and NCAPH in mitosis (Kimura et al. 2001, Takemoto et al. 2006), but other mitotic kinases may also be involved. CDK1 phosphorylation sites in NCAPH have not been established. NCAPD2 threonine residues T1339, T1384 and T1389 are inferred to be phosphorylated by CDK1 based on homologues sites in Xenopus laevis Ncapd2 (Kimura et al. 1998). NCAPG threonine residues T308 and T332 are phosphorylated by CDK1 in vitro and functionally important. The functional importance of threonine T931, also phosphorylated by CDK1 in vitro, has not been demonstrated (Murphy et al. 2008). Phosphorylation by CDK1 is required for mitotic activation of condensin I and promotes chromosomal binding, introduction of positive supercoils into DNA, and chromatin condensation (Kimura et al. 1998, Kimura et al. 2001, Takemoto et al. 2006).
R-HSA-2520883 (Reactome) While condensin II complex (consisting of subunits SMC2, SMC4, NCAPD3, NCAPG2 and NCAPH2), responsible for condensation of chromosomes in prophase (Hirota et al. 2004, Abe et al. 2011), is nuclear, condensin I is cytosolic and gains access to chromosomes only after the nuclear envelope breakdown at the start of prometaphase (Ono et al. 2004). Condensin I, activated by CDK1 phosphorylation (Kimura et al. 1998, Kimura et al. 2001, Takemoto et al. 2006, Murphy et al. 2008), promotes further condensation of chromosomes in prometaphase and metaphase, visible as longitudinal chromosome shortening (Hirota et al. 2004). Besides CDK1-mediated phosphorylation, association of condensin I with chromosomes may be regulated by AURKB (Lipp et al. 2007). In budding yeast, condensin phosphorylation by Cdc2 (CDK1 ortholog) is followed by Cdc5-mediated phosphorylation (Cdc5 is PLK1 ortholog), which is important for the sustained mitotic activity of condensin complex (St-Pierre et al. 2009). Phosphorylation by PLK1 is also important for the activation of human condensin II complex (Abe et al. 2011).
R-HSA-2529015 (Reactome) Inhibitory phosphate groups that were added to condensin I subunits by CK2 during interphase have to be removed for full mitotic activation of condensin I (Takemoto et al. 2006). The responsible phosphatase has not been identified.
R-HSA-2529020 (Reactome) Protein levels of condensin subunits are constant during the cell cycle. Four subunits, SMC4, NCAPD2, NCAPG and NCAPH, are phosphorylated in interphase cells (Takemoto et al. 2004) by CK2 i.e. casein kinase II (Takemoto et al. 2006). Except for the phosphorylation of NCAPH subunit on serine residue S570, CK2 phosphorylation sites in condensin I subunits have not been identified. Phosphorylation by CK2 inhibits condensin I-mediated introduction of positive supercoils into DNA and chromatin condensation. Mitotic activation of condensin I involves removal of phosphate groups added by CK2 (Takemoto et al. 2006), but the responsible phosphatase has not been identified.
R-HSA-2545253 (Reactome) HDAC8 deacetylates cohesin in prometaphase, after cohesin dissociates from chromosomal arms (Deardorff et al. 2012).
R-HSA-375302 (Reactome) The human kinetochore, is a complex proteinaceous structure that assembles on centromeric DNA and mediates the association of mitotic chromosomes with spindle microtubules in prometaphase. The molecular composition of the human kinetochore is reviewed in detail in Cheeseman et al., 2008. This complex structure is composed of numerous protein complexes and networks including: the constitutive centromere-associated network (CCAN) containing several sub-networks such as (CENP-H, I, K), (CENP-50/U, O, P, Q, R), the KMN network (containing KNL1, the Mis12 complex, and the Ndc80 complex), the chromosomal passenger complex, the mitotic checkpoint complex, the nucleoporin 107-160 complex and the RZZ complex.
At prometaphase, following breakdown of the nuclear envelope, the kinetochores of condensed chromosomes begin to interact with spindle microtubules. In humans, 15-20 microtubules are bound to each kinetochore (McEwen et al., 2001), and the attachment of 15 microtubules to the kinetochore is shown in this reaction. Recently, it was found that the core kinetochore-microtubule attachment site is within the KMN network and is likely to be formed by two closely apposed low-affinity microtubule-binding sites, one in the Ndc80 complex and a second in KNL1 (Cheeseman et al., 2006).
Sister Centromeres:Ac-Cohesin:PDS5:CDCA5:WAPALR-HSA-2484822 (Reactome)
Sister

Chromosomal

Arms:Ac-Cohesin:PDS5:CDCA5:WAPAL		R-HSA-2468293 (Reactome)
Sister

Chromosomal

Arms:Ac-Cohesin:PDS5:p-CDCA5:WAPAL		ArrowR-HSA-2468293 (Reactome)
Sister

Chromosomal

Arms:Ac-Cohesin:PDS5:p-CDCA5:WAPAL		R-HSA-2466068 (Reactome)
Sister

Chromosomal

Arms:p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL		ArrowR-HSA-2466068 (Reactome)
Sister

Chromosomal

Arms:p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL		R-HSA-2467794 (Reactome)
Sister Chromosomal ArmArrowR-HSA-2467794 (Reactome)
Unknown Phosphatasemim-catalysisR-HSA-2529015 (Reactome)
microtubuleR-HSA-375302 (Reactome)
p-Ac-Cohesin:PDS5:WAPALArrowR-HSA-2467794 (Reactome)
p-Ac-Cohesin:PDS5:WAPALR-HSA-2545253 (Reactome)
p-Cohesin:PDS5:WAPALArrowR-HSA-2545253 (Reactome)
p-RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesArrowR-HSA-1638821 (Reactome)
p-S21,S75,T159-CDCA5ArrowR-HSA-2467794 (Reactome)
p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesArrowR-HSA-1638803 (Reactome)
p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:MicrotubulesR-HSA-1638821 (Reactome)
p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubulesmim-catalysisR-HSA-1638821 (Reactome)
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