Mitotic Prometaphase (Homo sapiens)

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312, 13, 14, 202, 62, 5, 7, 8, 13...3, 4, 12, 21, 23...4, 9, 329, 19221, 4, 9, 10, 16...11, 15, 25, 30, 319, 1923, 4, 32Ac-Cohesin Complex Kinetochore Mis12 complex Chromosome passenger complex Ac-CohesinPDS5p-CDCA5WAPAL NDC80 complex Mitotic checkpoint complex NDC80 complex CENP-CENP-PP2A p-STAG2,RAD21-Ac-Cohesin GO0003831Nup107-160 complex Sister CentromereKinetochore CCAN network PP2A CENP-O complex Mis12 complex Nup107-160 complex CENP-CCAN network Mis12 complex Condensin I Ac-Cohesin Complex KMN network CentromereAc-CohesinPDS5CDCA5WAPAL PP2A CENP-O complex PDS5 Ac-CohesinPDS5p-CDCA5WAPALSister CentromeresKinetochoresMicrotubules PDS5 Nup107-160 complex Ac-Cohesin Complex Nup107-160 complex Chromosome passenger complex PDS5 Ac-Cohesin Complex RZZ complex Sister Chromosomal ArmsAc-CohesinPDS5CDCA5WAPAL CCAN network CCAN network CENP-O complex Chromosome passenger complex ChEBI28061PP2A Mitotic checkpoint complex p-Ac-CohesinPDS5WAPAL Ac-CohesinPDS5CDCA5WAPALSister CentromeresKinetochoresMicrotubules PDS5 Casein kinase II p-CohesinPDS5WAPAL CK2 Phosphorylated Condensin I NDC80 complex p-STAG2,RAD21-Ac-Cohesin p-STAG2,RAD21-Ac-CohesinPDS5p-CDCA5WAPAL Ac-Cohesin Complex p-Cohesin Nup107-160 complex NDC80 complex PDS5 Microtubule-bound kinetochore Kinetochore Mis12 complex Kinetochore p-RAD21-Ac-Cohesin Mitotic checkpoint complex p-RAD21-Ac-CohesinPDS5p-CDCA5WAPALSister CentromeresKinetochoresMicrotubules p-STAG2,RAD21-Ac-CohesinPDS5p-CDCA5WAPALSister CentromeresKinetochoresMicrotubules KSPGSister Chromosomal ArmsAc-CohesinPDS5p-CDCA5WAPAL RZZ complex KMN network PDS5 Sister CentromeresAc-CohesinPDS5CDCA5WAPAL CCAN network Kinetochore RZZ complex PP2A Ac-CohesinPDS5CDCA5WAPAL PDS5 RZZ complex Ac-CohesinPDS5CDCA5WAPAL Kinetochore PP2A cytosolCENP-CENP-O complex Mitotic checkpoint complex Mitotic checkpoint complex CENP-Nup107-160 complex Chromosome passenger complex CDK1 Phosphorylated Condensin I CENP-Microtubule-bound kinetochore Chromosome passenger complex Ac-CohesinPDS5CDCA5WAPAL Gal Microtubule-bound kinetochore Kinetochore KMN network NDC80 complex PDS5 Microtubule-bound kinetochore Sister Chromosomal Armsp-STAG2,RAD21-Ac-CohesinPDS5p-CDCA5WAPAL Mis12 complex CENP-O complex Mitotic checkpoint complex CENP-O complex RZZ complex PDS5 Chromosome passenger complex CCAN network KMN network PDS5 Mis12 complex Ac-CohesinPDS5WAPALCentromereKinetochore RZZ complex KMN network NDC80 complex KMN network p-Ac-Cohesin CENPP 2xAcK-SMC3 SGOL1 CoA-SHKIF2C CENPTp-S21,S75,T159-CDCA5 CENPA SPDL1 MIS12 STAG1 p-T1339,T1384,T1389-NCAPD2 PDS5B NUP160 SEC13 CENPM Condensin Ip-S-NCAPD2 SPC25 NUF2 ADPMAD2L1 WAPAL APITD1 NSL1 PPP1CC SPC25 CK2 Phosphorylated Condensin INUF2 ITGB3BP ATPXPO1 NSL1 KIF2C CDCA8 NUP37 CENPA PMF1AURKBPDS5B ZWILCH NDC80 NUP98-5 STAG2 NUP160 CLIP1 2xAcK-SMC3 NUP98-5 NUP85 NDEL1 CDCA5 SPDL1 MAPRE1 MAD2L1 MAD2L1 PLK1HDAC8STAG2 TAOK1 PPP1CC NUP107 NSL1 PAFAH1B1NUP98-5 RPS27 RCC2 Sister Chromosomal ArmsAc-CohesinPDS5p-CDCA5WAPALSEC13 ZW10 CLASP2 ZWINT CENPC1 SMC2 XPO1 PDS5B PDS5B APITD1 CENPH PAFAH1B1KIF2B ZW10 NUP85 CLIP1 SPC25 CKAP5 B9D2 p-S21,S75,T159-CDCA5 KIF18A SGOL2 BUB1 CENPE SMC1A SKA1SEH1L-1 CLASP1 CKAP5 KIF18A CSNK2BKIF2C NUP133 CENPF KIF18A CENPM PDS5A STAG2 ERCC6L NUP43 RANGAP1 NUP133 CENPN PDS5B KNTC1 CLASP2 ADPSGOL1 SEC13 PDS5A CKAP5 MAD1L1 MLF1IP CENPA ERCC6L CENPH CLIP1 KIF2C SEH1L-1 CLIP1 ADPPLK1 NUP160 AURKBKNTC1 Sister Chromosomal Armsp-STAG2,RAD21-Ac-CohesinPDS5p-CDCA5WAPALNDE1 RCC2 CENPP CDC20 CENPL NUDC NUP43 KIF18A Sister Chromosomal ArmCENPM Microtubule-bound kinetochoreAHCTF1 CENPN 2xAcK-SMC3 ADPB9D2 NUDC PDS5A SEC13 KIF2A SKA2 ZWILCH PMF1NUP85 WAPAL SGOL2 MAD1L1 CENPO MAD2L1 NUP160 ITGB3BP KNTC1 CLIP1 SMC1A NCAPG CENPO CENPC1 CENPM CENPL p-S21,S75,T159-CDCA5 SMC1A SKA2 KIF2B AURKBNDE1 KIF2A ERCC6L CENPN SMC3 NDE1 SPC24 CASC5 SKA2 ZWILCH CSNK2A2 NSL1 STAG1 NDC80 CENPK NCAPD2 DSN1 CDCA8 RCC2 KIF2A RANBP2 CSNK2A1 SMC2 STAG1 DSN1 NUP43 CENPQ RANBP2 ZWILCH B9D2 ZW10 CENPQ RAD21 NDEL1 PAFAH1B1p-STAG2 SGOL1 PAFAH1B1BIRC5 CENPO BUB1B KNTC1 NUP133 NUP98-5 NUP85 XPO1 CDC20 TAOK1 RAD21 CDCA8 MIS12 NDC80 MIS12 NUP107 CENPP CENPO 2xAcK-SMC3 Ac-CoARANGAP1 NUP98-5 CENPH RPS27 p-S-NCAPG CENPP STAG2 NUDC KNTC1 MIS12 CENPL CENPI RANGAP1 CLASP2 PDS5A BUB1B AURKBSGOL2 BIRC5 ERCC6L PLK1 SMC4 CENPC1 KIF2B NSL1 CDC20 CENPI CENPO XPO1 CENPP NDE1 KinetochoreCASC5 CENPN CENPL p-CohesinPDS5WAPALp-RAD21-Ac-CohesinPDS5p-CDCA5WAPALSister CentromeresKinetochoresMicrotubulesSMC2 p-T308,T332-NCAPG S PhaseSGOL1 KIF2C MAPRE1 SMC1A B9D2 Unknown PhosphataseCENPC1 ZW10 PMF1BUB3 SEH1L-1 INCENP Ac-CohesinPDS5CDCA5WAPALSister CentromeresKinetochoresMicrotubulesZWILCH MLF1IP SPC25 RANBP2 PDS5A BUB3 AURKBp-S454-RAD21 CENPC1 CDC20 CENPI KIF2A NUP37 PPP1CC AHCTF1 CENPN CASC5 PiPiKIF2B NDEL1 NSL1 p-T-NCAPH H2OCENPF MAD2L1 CLASP2 CKAP5 DSN1 NUP107 SMC1A MAD1L1 WAPAL BUB1 PLK1 CENPQ DSN1 CENPTCENPE BUB3 SKA1SPC25 CASC5 CENPM Condensed prometaphase chromosomesCENPH INCENP NUP107 p-S454-RAD21 NUP133 BUB3 CDCA8 INCENP CDCA5 ATPCENPK SPC24 NDC80 BUB1B SMC1A TAOK1 ITGB3BP WAPAL INCENP MAD1L1 INCENP PAFAH1B1SGOL2 ZWINT NUP107 KIF18A SKA1STAG2 SGOL2 PMF1SKA1ERCC6L SKA1CLASP1 NCAPH ATPBUB1B CENPH DSN1 MAD1L1 CENPK XPO1 RCC2 CENPI p-STAG2,RAD21-Ac-CohesinPDS5p-CDCA5WAPALSister CentromeresKinetochoresMicrotubulesNUP37 CENPA Sister CentromeresAc-CohesinPDS5CDCA5WAPALTAOK1 CLASP2 KIF18A SKA2 STAG1 RPS27 KIF2A CENPE SPDL1 SMC1A AURKBSMC1A NUF2 SMC1A ITGB3BP PPP1CC BUB1B BUB3 NUP37 ZWINT INCENP 2xAcK-SMC3 CLASP1 CDCA8 STAG1 CENPH APITD1 CDC20 BUB1 CENPL SGOL2 WAPAL KNTC1 NUP160 CENPE CENPTSMC4 BIRC5 p-S21,S75,T159-CDCA5 SPC24 SKA1CENPN RPS27 CENPM CENPL KIF2B CDCA8 STAG1 PDS5A ITGB3BP SGOL1 BUB1B NDE1 NUP43 KIF2C CLASP1 WAPAL ZWINT STAG1 RANGAP1 SPDL1 ADPRPS27 SPC24 PDS5A PDS5B CENPTSEH1L-1 CENPQ ZWILCH NUF2 RPS27 AHCTF1 Sister Chromosomal ArmsAc-CohesinPDS5CDCA5WAPALPDS5B CENPQ ITGB3BP p-S21,S75,T159-CDCA5 SPDL1 NUP133 SEC13 CENPK PAFAH1B1WAPAL ADPMAPRE1 BUB1 PLK1 XPO1 KIF2A CENPO MAD2L1 PMF1RANGAP1 KIF2B CLASP1 WAPAL p-S454-RAD21 MIS12 PLK1 NUP37 PLK1 PDS5A PDS5B ATPPDS5B NUP85 DSN1 NUDC BIRC5 p-S570-NCAPH H2O2xAcK-SMC3 CENPC1 RCC2 NDEL1 NDC80 STAG1 CENPA NUP43 RANGAP1 CENPA Ac-CohesinPDS5p-CDCA5WAPALSister CentromeresKinetochoresMicrotubulesNUP85 CENPTNUP160 BIRC5 RCC2 ZW10 MLF1IP SGOL1 CKAP5 2xAcK-SMC3 SPC24 AHCTF1 PDS5A RAD21 2xAcK-SMC3 MLF1IP p-STAG2 p-S21,S75,T159-CDCA5MLF1IP CDK1 Phosphorylated Condensin INDEL1 ATPCENPF SPDL1 PPP1CC SEH1L-1 AHCTF1 microtubulep-S454-RAD21 NUP37 BIRC5 ZWINT CENPF p-Ac-CohesinPDS5WAPALCDC20 CENPP RAD21 MIS12 CENPK PDS5B NUDC CASC5 CENPQ PMF1ERCC6L B9D2 APITD1 CCNBp-T161-CDK1CENPE CENPF APITD1 NUP43 CLASP1 CENPI NDE1 AHCTF1 MAPRE1 BUB1 p-STAG2 CENPE CENPK CLASP2 CASC5 TAOK1 NUP107 RANBP2 CDCA5 NDEL1 MAPRE1 TAOK1 SEC13 STAG1 PDS5A BUB1 Casein kinase IINUP98-5 p-STAG2 SPC24 SKA2 Condensed prophase chromosomesRANBP2 ATPNDC80 WAPAL CENPI ZW10 PPP1CC p-S454-RAD21 MAD1L1 SPC25 p-S-SMC4 SMC1A APITD1 NUF2 ZWINT STAG1 NUDC CENPF STAG2 WAPAL CKAP5 SEH1L-1 B9D2 BUB3 CLIP1 RAD21 2xAcK-SMC3 NUP133 CENPTMAPRE1 MLF1IP SKA2 Mitotic Metaphase and AnaphaseNUF2 RANBP2 1414


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. Original Pathway at Reactome: http://www.reactome.org/PathwayBrowser/#DB=gk_current&FOCUS_SPECIES_ID=48887&FOCUS_PATHWAY_ID=68877

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Bibliography

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  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
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  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
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History

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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

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NameTypeDatabase referenceComment
2xAcK-SMC3 ProteinQ9UQE7 (Uniprot-TrEMBL)
ADPMetaboliteCHEBI:16761 (ChEBI)
AHCTF1 ProteinQ8WYP5 (Uniprot-TrEMBL)
APITD1 ProteinQ8N2Z9 (Uniprot-TrEMBL)
ATPMetaboliteCHEBI:15422 (ChEBI)
AURKBProteinQ96GD4 (Uniprot-TrEMBL)
Ac-CoAMetaboliteCHEBI:15351 (ChEBI)
Ac-Cohesin

PDS5 CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		ComplexREACT_151019 (Reactome)
Ac-Cohesin

PDS5 p-CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		ComplexREACT_152183 (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)
CCNB p-T161-CDK1ComplexREACT_148199 (Reactome)
CDC20 ProteinQ12834 (Uniprot-TrEMBL)
CDCA5 ProteinQ96FF9 (Uniprot-TrEMBL)
CDCA8 ProteinQ53HL2 (Uniprot-TrEMBL)
CDK1 Phosphorylated Condensin IComplexREACT_150472 (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)
CENPTProteinQ96BT3 (Uniprot-TrEMBL)
CK2 Phosphorylated Condensin IComplexREACT_151678 (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)
CSNK2BProteinP67870 (Uniprot-TrEMBL)
Casein kinase IIComplexREACT_23098 (Reactome)
CoA-SHMetaboliteCHEBI:15346 (ChEBI)
Condensed prometaphase chromosomesREACT_150762 (Reactome)
Condensed prophase chromosomesREACT_151302 (Reactome)
Condensin IComplexREACT_151162 (Reactome)
DSN1 ProteinQ9H410 (Uniprot-TrEMBL)
ERCC6L ProteinQ2NKX8 (Uniprot-TrEMBL)
H2OMetaboliteCHEBI:15377 (ChEBI)
HDAC8ProteinREACT_151192 (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)
KinetochoreComplexREACT_14970 (Reactome)
MAD1L1 ProteinQ9Y6D9 (Uniprot-TrEMBL)
MAD2L1 ProteinQ13257 (Uniprot-TrEMBL)
MAPRE1 ProteinQ15691 (Uniprot-TrEMBL)
MIS12 ProteinQ9H081 (Uniprot-TrEMBL)
MLF1IP ProteinQ71F23 (Uniprot-TrEMBL)
Microtubule-bound kinetochoreComplexREACT_15175 (Reactome)
Mitotic Metaphase and AnaphasePathwayWP2757 (WikiPathways) 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)
PAFAH1B1ProteinP43034 (Uniprot-TrEMBL)
PDS5A ProteinQ29RF7 (Uniprot-TrEMBL)
PDS5B ProteinQ9NTI5 (Uniprot-TrEMBL)
PLK1 ProteinP53350 (Uniprot-TrEMBL)
PLK1ProteinP53350 (Uniprot-TrEMBL)
PMF1ProteinQ6P1K2 (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 PhasePathwayWP2772 (WikiPathways) 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)
SKA1ProteinQ96BD8 (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

WAPAL		ComplexREACT_151874 (Reactome)
Sister Chromosomal ArmREACT_150847 (Reactome)
Sister Chromosomal Arms

Ac-Cohesin PDS5 CDCA5

WAPAL		ComplexREACT_150576 (Reactome)
Sister Chromosomal Arms

Ac-Cohesin PDS5 p-CDCA5

WAPAL		ComplexREACT_151061 (Reactome)
Sister Chromosomal Arms

p-STAG2,RAD21-Ac-Cohesin PDS5 p-CDCA5

WAPAL		ComplexREACT_151301 (Reactome)
TAOK1 ProteinQ7L7X3 (Uniprot-TrEMBL)
Unknown PhosphataseREACT_150933 (Reactome)
WAPAL ProteinQ7Z5K2 (Uniprot-TrEMBL)
XPO1 ProteinO14980 (Uniprot-TrEMBL)
ZW10 ProteinO43264 (Uniprot-TrEMBL)
ZWILCH ProteinQ9H900 (Uniprot-TrEMBL)
ZWINT ProteinO95229 (Uniprot-TrEMBL)
microtubuleREACT_10446 (Reactome)
p-Ac-Cohesin

PDS5

WAPAL		ComplexREACT_151851 (Reactome)
p-Cohesin

PDS5

WAPAL		ComplexREACT_151704 (Reactome)
p-RAD21-Ac-Cohesin

PDS5 p-CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		ComplexREACT_151499 (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

Microtubules		ComplexREACT_150708 (Reactome)
p-T-NCAPH ProteinQ15003 (Uniprot-TrEMBL)
p-T1339,T1384,T1389-NCAPD2 ProteinQ15021 (Uniprot-TrEMBL)
p-T308,T332-NCAPG ProteinQ9BPX3 (Uniprot-TrEMBL)

Annotated Interactions

View all...
SourceTargetTypeDatabase referenceComment
ADPArrowREACT_150152 (Reactome)
ADPArrowREACT_150232 (Reactome)
ADPArrowREACT_150239 (Reactome)
ADPArrowREACT_150246 (Reactome)
ADPArrowREACT_150414 (Reactome)
ADPArrowREACT_150454 (Reactome)
ATPREACT_150152 (Reactome)
ATPREACT_150232 (Reactome)
ATPREACT_150239 (Reactome)
ATPREACT_150246 (Reactome)
ATPREACT_150414 (Reactome)
ATPREACT_150454 (Reactome)
Ac-CoAArrowREACT_150215 (Reactome)
Ac-Cohesin

PDS5 CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		REACT_150414 (Reactome)
Ac-Cohesin

PDS5 p-CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		ArrowREACT_150414 (Reactome)
Ac-Cohesin

PDS5 p-CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		REACT_150152 (Reactome)
CCNB p-T161-CDK1REACT_150232 (Reactome)
CCNB p-T161-CDK1REACT_150414 (Reactome)
CCNB p-T161-CDK1REACT_150454 (Reactome)
CDK1 Phosphorylated Condensin IArrowREACT_150232 (Reactome)
CDK1 Phosphorylated Condensin IArrowREACT_150278 (Reactome)
CK2 Phosphorylated Condensin IArrowREACT_150239 (Reactome)
CK2 Phosphorylated Condensin IREACT_150403 (Reactome)
Casein kinase IIREACT_150239 (Reactome)
CoA-SHREACT_150215 (Reactome)
Condensin IArrowREACT_150403 (Reactome)
Condensin IREACT_150232 (Reactome)
Condensin IREACT_150239 (Reactome)
H2OREACT_150250 (Reactome)
H2OREACT_150403 (Reactome)
HDAC8REACT_150215 (Reactome)
KinetochoreREACT_14798 (Reactome)
Microtubule-bound kinetochoreREACT_150388 (Reactome)
PLK1REACT_150152 (Reactome)
PLK1REACT_150246 (Reactome)
PiArrowREACT_150250 (Reactome)
PiArrowREACT_150403 (Reactome)
REACT_14798 (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).
REACT_150152 (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).
REACT_150215 (Reactome) HDAC8 deacetylates cohesin in prometaphase, after cohesin dissociates from chromosomal arms (Deardorff et al. 2012).
REACT_150232 (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).
REACT_150239 (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.
REACT_150246 (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).
REACT_150250 (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).
REACT_150267 (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).
REACT_150278 (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).
REACT_150388 (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.
REACT_150403 (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.
REACT_150414 (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).
REACT_150454 (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).
Sister Centromeres

Ac-Cohesin PDS5 CDCA5

WAPAL		REACT_150388 (Reactome)
Sister Chromosomal ArmArrowREACT_150267 (Reactome)
Sister Chromosomal Arms

Ac-Cohesin PDS5 CDCA5

WAPAL		REACT_150454 (Reactome)
Sister Chromosomal Arms

Ac-Cohesin PDS5 p-CDCA5

WAPAL		ArrowREACT_150454 (Reactome)
Sister Chromosomal Arms

Ac-Cohesin PDS5 p-CDCA5

WAPAL		REACT_150246 (Reactome)
Sister Chromosomal Arms

p-STAG2,RAD21-Ac-Cohesin PDS5 p-CDCA5

WAPAL		ArrowREACT_150246 (Reactome)
Unknown PhosphataseREACT_150403 (Reactome)
microtubuleREACT_14798 (Reactome)
p-Ac-Cohesin

PDS5

WAPAL		ArrowREACT_150267 (Reactome)
p-Ac-Cohesin

PDS5

WAPAL		REACT_150215 (Reactome)
p-Cohesin

PDS5

WAPAL		ArrowREACT_150215 (Reactome)
p-RAD21-Ac-Cohesin

PDS5 p-CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		ArrowREACT_150250 (Reactome)
p-S21,S75,T159-CDCA5ArrowREACT_150267 (Reactome)
p-STAG2,RAD21-Ac-Cohesin

PDS5 p-CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		ArrowREACT_150152 (Reactome)
p-STAG2,RAD21-Ac-Cohesin

PDS5 p-CDCA5 WAPAL Sister Centromeres Kinetochores

Microtubules		REACT_150250 (Reactome)
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