S Phase (Homo sapiens)

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1, 37, 4929, 31, 40, 41, 525132, 3343615622, 27, 47, 585, 20, 54, 585, 2730, 5819, 44318, 42, 53483655271238, 39145117, 29, 40, 52, 57...12, 6, 8, 39, 45...8, 10, 25519, 245051527, 30, 5821, 315734562831611346Processive complexOkazaki fragment complex DNA Polymerase delta tetramer DNA polymerase alphaprimase RNA primer-DNA primerorigin duplexPCNA RNA primer-DNA primerorigin duplex Processive complex PCNA homotrimer Processive complexOkazaki fragmentFlap CohesinPDS5WAPAL Processive complex RFC Heteropentamer DNA Polymerase delta tetramer PCNA homotrimer PCNA homotrimer RPA heterotrimer DNA polymerase alphaprimase PCNA homotrimer Cyclin ACdk2 complex RNA primerorigin duplexDNA polymerase alphaprimase complex RNA primer-DNA primerorigin duplex PDS5 RNA primerorigin duplexDNA polymerase alphaprimase complex PDS5 Ac-CohesinPDS5WAPALCentromere RFC HeteropentamerRNA primer-DNA primerorigin duplex DNA polymerase epsilon DNA Polymerase delta tetramer Cyclin E/ACdk2phospho-p27/p21 CDK Mcm2-7 complex Cyclin Aphospho-Cdk2DNA Polymerase delta tetramer ubiquitinated Cdc6 ubiquitinated Orc1 phospho-p27/p21 RFC HeteropentamerRNA primer-DNA primerorigin duplexPCNA homotrimer Sister Chromosomal ArmsAc-CohesinPDS5CDCA5WAPAL RFC Heteropentamer Cyclin E/ACdk2phospho-p27/p21 Cyclin ACdk2 complex DNA polymerase alphaprimase RNA primerorigin duplexDNA polymerase alphaprimase complex ORCorigin SCFRNA primer-DNA primerorigin duplex RPA heterotrimer PCNA homotrimer PDS5 Ac-CohesinPDS5CDCA5WAPAL RNA primerorigin duplexDNA polymerase alphaprimase complex phosphoCyclin ACdk2 complex Cyclin E/A Cyclin E/ACdk2 nucleoplasmCDT1CDC6ORCorigin complex CentromereAc-CohesinPDS5CDCA5WAPAL RNA primer-DNA primerorigin duplexPCNA phospho-p27/p21 GINS complex CDC6ORCorigin complex Orc2 associated with MCM8 Mcm2-7 complex DNA polymerase alphaprimase PDS5 DNA polymerase alphaprimase Cyclin E/ACdk2 SCFCohesinPDS5WAPALChromosomal Arm Processive complexOkazaki fragmentFlap Processive complexOkazaki fragmentsRemaining Flap Cyclin E/A Processive complexnicked DNA from adjacent Okazaki fragments SCFubiquitinated Orc1 DNA Polymerase delta tetramer DNA polymerase alphaprimaseDNA polymerase alphaorigin complex p21/p27 CAK Ac-CohesinPDS5CDCA5WAPAL SCFPCNA homotrimer RNA primer-DNA primerorigin duplexPCNA DNA Polymerase delta tetramer RNA primerorigin duplexDNA polymerase alphaprimase complex Ub phosphorylated Cyclin ACdk2 substrate proteins Cyclin E/ACdk2 RNA primer-DNA primerorigin duplex RNA primerorigin duplexDNA polymerase alphaprimase complex p21/p27 Processive complex RNA primer-DNA primerorigin duplex DNA polymerase epsilon PDS5 RNA primer-DNA primerorigin duplexPCNA GINS complex phospho-p27/p21 Sister CentromeresAc-CohesinPDS5CDCA5WAPAL RNA primerorigin duplexDNA polymerase alphaprimase complex Cyclin ACdk1 substrate proteins RNA primer-DNA primerorigin duplexPCNA PDS5 pre-replicative complex RNA primer-DNA primerorigin duplex RNA primer-DNA primerorigin duplexPCNA 26S proteasome multi-ubiquitinated phospho-Processive complex RPA heterotrimer Processive complexOkazaki fragmentFlapRPA heterotrimer DNA polymerase alphaprimase SCFCyclin E/ACdk2p27/p21 complex RNA primerorigin duplexDNA polymerase alphaprimase complex Ac-Cohesin Complex ubiquitin PCNA homotrimer DNA Polymerase delta tetramer RNA primer-DNA primerorigin duplex Cohesin Complex 26S proteasome Cyclin E/ACdk2 Cyclin ACdk2phosphorylated substrate complex Cyclin E/A Cyclin E/ACdk2multiubiquitinated phospho-p27/p21SCFProcessive complexOkazaki fragmentFlap Cyclin ACdk2substrate complex DNA polymerase alphaprimase Cyclin E/ACdk2phospho-p27/p21 Cyclin E/A PCNA homotrimer RNA primer-DNA primerorigin duplexPCNA CohesinPDS5WAPALCentromere Ub Cyclin D1Cdk4 Processive complexOkazaki fragment complex Mcm4,6,7 complex Processive complexOkazaki fragmentFlapRPA heterotrimerdna2 DNA Polymerase delta tetramer Cyclin Aphospho-Cdk2Cyclin Aphospho-Cdk2SCFDNA polymerase alphaprimase Cyclin E/ACdk2 PCNA homotrimer Ac-Cohesin Complex Processive complex Processive complexOkazaki fragment complex DNA polymerase alphaprimase DNA polymerase alphaprimase DNA polymerase alphaprimase Cyclin E/ACdk2phospho-p27/p21SCFCyclin E/A RNA primerorigin duplexDNA polymerase alphaprimase complex RNA primer-DNA primerorigin duplex Cyclin ACdk2p21/p27 complex Processive complexOkazaki fragmentFlapRPA heterotrimer Ac-CohesinPDS5WAPALChromosomal Arm RFC HeteropentamerRNA primer-DNA primerorigin duplex Myc/Max heterodimer Processive complex DNA polymerase alphaprimase Ac-Cohesin Complex CohesinPDS5WAPAL RNA primer-DNA primerorigin duplex RNA primer-DNA primerorigin duplex Cyclin E/ACdk2phospho-p27/p21SCFUb DNA polymerase alphaprimase DNA polymerase alphaprimase RNA primerorigin duplexDNA polymerase alphaprimase complex ubiquitin RNA primerorigin duplexDNA polymerase alphaprimase complex SCFRNA primerorigin duplexDNA polymerase alphaprimase complex Mcm2-7 complex phospho-p27/p21 Unwinding complex at replication fork RNA primer-DNA primerorigin duplex Cyclin Aphospho-Cdk2Cohesin Complex Cyclin ACdk2 complex DNA polymerase epsilonorigin complex pre-replicative complex Ac-Cohesin Complex Ac-CohesinPDS5WAPAL phosphoRFC Heteropentamer Processive complex PCNA homotrimer Processive complexOkazaki fragment complex RNA primer-DNA primerorigin duplexPCNA Orc2 associated with MCM8 Mcm2-7 complex Ac-CohesinPDS5WAPAL Cyclin ACdk2phospho-p27/p21 complex ubiquitinated Cdc62xAcK-SMC3 ATPPSMD2 POLD3 ATPPOLA2 2xAcK-SMC3 POLE2 PSMB7 PRIM1 PSMB2 POLD2 CDKN1A POLD3 POLE p-S,T-ORC1RPA2 PRIM1 Processive complexnicked DNA from adjacent Okazaki fragmentsPSMB8 CCNH PSMA1 POLD1 Mitotic Telophase/CytokinesisPCNA UBCPRIM1 PSMB6 PSMA4 PRIM2 RPA1 PRIM2 PSMB3 PSMD10 UBA52UBCPOLD4 p-T160-CDK2 PSF3pUBCCohesinPDS5WAPALCentromereSMC3 UBCPSMB11 GINS2 RPS27AADPUbPCNA POLA2 STAG2 UBCPOLA1 CDK2 p-T286-CCND1RAD21 CDK4CDKN1B pre-replicative complexPRIM2 POLD3 PSMC4 PSME1 RAD21 p-T160-CDK2 MCM2 RFC1 PSMD7DNA Polymerase delta tetramerAMPp-T187-CDKN1B PSMA6 CDK2 PSMA5 PSMD1 CDKN1BRNA primer-DNA primerorigin duplexPCNAGINS1POLD2 POLA2 POLA1 POLA2 p-FZR1 POLD4 PiRFC3 Sister CentromeresAc-CohesinPDS5CDCA5WAPALUBBWAPAL Okazaki fragmentADPPOLE2 POLD3 CDK2 SKP1 p-S130-CDKN1A MCM8POLA1 MCM7 STAG1 PRIM1 PSMD14 UBCPDS5B PRIM2 UBCCDKN1A Cyclin ACdk2phosphorylated substrate complexPOLD1 RFC5 GINS1 ORC1 Sister Chromosomal ArmsAc-CohesinPDS5CDCA5WAPALPCNA POLA1 PDS5B PSMA2 PSME1 STAG1 CDCA5CDK2 PSMD11 PSMD6UBCADPATPPOLA1 Cyclin AReplication ForkCDC6UBBPSMC5 MNAT1 SMC1A CDT1 POLA2 POLD4 STAG2 MYC SKP2 CDK7 ORC2 CDK2 PCNA UBCMCM5 Cyclin ACdk2 complexPSMA4 PSMD3 UBBPSME2 Ac-CohesinPDS5WAPALCentromerePSMD2 Processive complexOkazaki fragmentFlapORC3 CDK4UMPPSMD14 ATPRAD21 ADPWEE1PDS5A RB1 RFC2 POLA2 PRIM2 UBCPSME3 CDKphosphoUBCPSMD7CDCA5 LIG1UBCCDC45UBCUBBRPA1 Cyclin E/ACdk2p27/p21 complexSMC1A PRIM1 PRIM1 ligated okazaki fragmentPOLD1 SMC1A PSMB2 UBCp-S130-CDKN1A MCM2 SCFPOLA1 SKP2 UBCPOLD2 Processive complexOkazaki fragmentFlapRPA heterotrimerMcm2-7 complexCDK2 ubiquitinated Orc1RPA1 UBBADPPRIM1 UBCMCM6 POLD2 PSMB9 POLD3 STAG1 MCM6 ORC1 p-T286-CCND1 POLD3 POLA1 POLD1 POLD2 PSMA1 Unwinding complex at replication forkCDC6Unwound forkCDKN1B2xAcK-SMC3 MCM3 Sister Chromosomal ArmCDC6 dGTPPOLA1 26S proteasomeRPA2 MCM5RFC4 PRIM2 POLD2 CDK2 Myc/Max heterodimerMCM4Cyclin E/ACdk2phospho-p27/p21POLA2 PSMD8 MCM7 PSMD11 PRIM1 Cyclin D1Cdk4MCM6 PDS5A Processive complexOkazaki fragmentsRemaining FlapSMC1A POLD4 Cyclin ACdk2substrate complexSTAG1 CDCA5 UBCUBBSKP1 ORC4 POLA1 POLD3 PSMB4 RPA3 CDKN1B NTPSMC3 PSMB5 POLD2 DNA22xAcK-SMC3 CDK4CDKN1B CDK2 SMC1A RFC2 UBCDNA polymerase alphaprimaseMCM6 PRIM2 CAKPOLA2 WAPAL Ac-CohesinPDS5WAPALChromosomal ArmPSMD12 MAXPOLD1 PCNA SKP2 RPA heterotrimerUBA52RPA3 Processive complexOkazaki fragment complexanaphase-promoting complex RFC5 GINS complexProcessive complexOkazaki fragmentFlapRPA heterotrimerdna2ATPGINS4PSMA2 CUL1 UBBUBCORC2 POLA1 RFC HeteropentamerRNA primer-DNA primerorigin duplexCyclin E/ACdk2multiubiquitinated phospho-p27/p21SCFPSMC1PDS5B PSMD4RAD21 PSMB6 Cyclin E/ACdk2phospho-p27/p21SCFUBCMCM2 WAPAL PSMB5 CDKN1A PSMB10 SKP2 CDC25APDS5A UBBCKS1B UBA52STAG2 UBCPOLA1 MCM8 PRIM1 POLA1 UBCdCTPUBCCUL1 PSMD4RFC2 DNA2 UBCMCM7 UBBPSMB10 SKP1 PDS5B PRIM2 RFC1 ORC5 CKS1B PSMC5 UBCPOLA2 PRIM2 PSMD5 UBCPSMC6UBCUBCCUL1 GINS1 PCNA PSME2 UBCMCM2p21/p27PRIM2 MCM4DNA polymerase alphaprimaseDNA polymerase alphaorigin complexORC4 MCM4PSMD5 ADPPOLD2 RAD21 PDS5B SMC1A MCM4CDK2 MCM3 PDS5A RFC4 CMPPSMD12 p-S130-CDKN1A PSMB7 dTTPUBCMCM3 STAG1 SCFPOLD4 Mitotic PrometaphaseORC3 PSMD13 RFC4 Cyclin Aphospho-Cdk2WAPAL RFC5 PSMD3 PDS5B Cyclin ACdk2p21/p27 complexPSMC1p-T160-CDK2 26S proteasomeSTAG2 UBCADPUBCPRIM1 GINS2UBCPSMA5 p-Y15-CDK2 MCM5 PSMD10 CDC6 p-T187-CDKN1B MCM7 RPS27ACKS1Bp-T187-CDKN1B MCM2 UBA52RFC HeteropentamerORC5 p-T286-CCND1 UBBUbiquitin ligasePSMA7PCNA homotrimerPRIM2 UBCPSMD9 PRIM1 POLA2 UBCATPMCM6 PSMD13 PSME3 RNA primerorigin duplexDNA polymerase alphaprimase complexCCND1 Cyclin Aphospho-Cdk2ATPUBA52CDC45 phosphop-T187-CDKN1B p-T187-CDKN1B PSMC3 POLD1 CDT1 ATPPOLD3 ADPATPMCM8 CDK2 Cdc25 A/BESCODNA polymerase epsilonPSMC6RFC HeteropentamerRNA primer-DNA primerorigin duplexPCNA homotrimerSKP1 p-RB1 MCM4Processive complexPSMC2 UBCPSMD6PSMD8 RPS27AdATPPSME4 PDS5A RPA2 RFC3 MCM7 GINS4ubiquitinSTAG2 PSMB1 PSMB4 PSMA3 GMPPRIM1 POLA2 PSMB9 PSMF1CDKN1AWAPAL MCM3PSMC4 PSMB3 POLD4 Mcm4,6,7 complexRFC3 PSMA3 UBCMCM3 CDC6 POLD4 UBBUBCPOLE POLD1 multi-ubiquitinated phospho-RAD21 RPS27ARPA3 PSMA6 UBCUBBPOLA1 CDKN1Aubiquitinated Orc1H2OPSMB8 ORC6 CohesinPDS5WAPALChromosomal ArmPSMC2 PSMB1 GINS2 CDK2GINS4UBCPSMC3 PSMF1PSMD1 CKS1B PRIM1 POLA2 FEN1UBCMCM5 Cyclin ACdk2phospho-p27/p21 complexp-T286-CCND1 POLA1 RNA primer-DNA primerorigin duplexAc-CoAPCNA PSMD9 MCM5 POLD1 WAPAL p-S130-CDKN1A STAG2 UBCUBBCoA-SHORC1 Sister CentromerePCNA glycogen synthase kinase-3 betaPCNA STAG1 POLD4 UBCPOLA2 PCNA POLA2 CUL1 PRIM1 UBBRFC1 p-S130-CDKN1A ORC6 UBBPSMA8 UBCPRIM2 FZR1 PSMA7Cyclin ACdk2 complexp-CDC6RPS27APDS5A pre-replicative complex Cyclin E/ACdk2p-T286-CCND1UBCPRIM2 PRIM2 46362643511811, 2335431551


Description

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. Original Pathway at Reactome: http://www.reactome.org/PathwayBrowser/#DB=gk_current&FOCUS_SPECIES_ID=48887&FOCUS_PATHWAY_ID=69242

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Bibliography

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History

View all...
CompareRevisionActionTimeUserComment
118522view10:10, 28 May 2021EweitzOntology Term : 'S phase pathway' added !
115059view17:00, 25 January 2021ReactomeTeamReactome version 75
113503view11:58, 2 November 2020ReactomeTeamReactome version 74
112703view16:10, 9 October 2020ReactomeTeamReactome version 73
101618view11:48, 1 November 2018ReactomeTeamreactome version 66
101154view21:34, 31 October 2018ReactomeTeamreactome version 65
100681view20:07, 31 October 2018ReactomeTeamreactome version 64
100231view16:52, 31 October 2018ReactomeTeamreactome version 63
99783view15:18, 31 October 2018ReactomeTeamreactome version 62 (2nd attempt)
99337view12:47, 31 October 2018ReactomeTeamreactome version 62
93537view11:26, 9 August 2017ReactomeTeamreactome version 61
86636view09:22, 11 July 2016ReactomeTeamreactome version 56
83395view11:07, 18 November 2015ReactomeTeamVersion54
81589view13:07, 21 August 2015ReactomeTeamVersion53
77049view08:35, 17 July 2014ReactomeTeamFixed remaining interactions
76754view12:11, 16 July 2014ReactomeTeamFixed remaining interactions
76079view10:14, 11 June 2014ReactomeTeamRe-fixing comment source
75789view11:32, 10 June 2014ReactomeTeamReactome 48 Update
75139view14:08, 8 May 2014AnweshaFixing comment source for displaying WikiPathways description
74786view08:52, 30 April 2014ReactomeTeamNew pathway

External references

DataNodes

View all...
NameTypeDatabase referenceComment
26S proteasomeComplexREACT_2353 (Reactome)
26S proteasomeComplexREACT_7467 (Reactome)
2xAcK-SMC3 ProteinQ9UQE7 (Uniprot-TrEMBL)
ADPMetaboliteCHEBI:16761 (ChEBI)
AMPMetaboliteCHEBI:16027 (ChEBI)
ATPMetaboliteCHEBI:15422 (ChEBI)
Ac-CoAMetaboliteCHEBI:15351 (ChEBI)
Ac-Cohesin

PDS5 WAPAL

Centromere
ComplexREACT_152186 (Reactome)
Ac-Cohesin

PDS5 WAPAL

Chromosomal Arm
ComplexREACT_151929 (Reactome)
CAKComplexREACT_5717 (Reactome)
CCND1 ProteinP24385 (Uniprot-TrEMBL)
CCNH ProteinP51946 (Uniprot-TrEMBL)
CDC25AProteinP30304 (Uniprot-TrEMBL)
CDC45 ProteinO75419 (Uniprot-TrEMBL)
CDC45ProteinO75419 (Uniprot-TrEMBL)
CDC6 ProteinQ99741 (Uniprot-TrEMBL)
CDC6ProteinQ99741 (Uniprot-TrEMBL)
CDCA5 ProteinQ96FF9 (Uniprot-TrEMBL)
CDCA5ProteinQ96FF9 (Uniprot-TrEMBL)
CDK2 ProteinP24941 (Uniprot-TrEMBL)
CDK2ProteinP24941 (Uniprot-TrEMBL)
CDK4ProteinP11802 (Uniprot-TrEMBL)
CDK7 ProteinP50613 (Uniprot-TrEMBL)
CDKN1A ProteinP38936 (Uniprot-TrEMBL)
CDKN1AProteinP38936 (Uniprot-TrEMBL)
CDKN1B ProteinP46527 (Uniprot-TrEMBL)
CDKN1BProteinP46527 (Uniprot-TrEMBL)
CDKComplexREACT_5476 (Reactome)
CDT1 ProteinQ9H211 (Uniprot-TrEMBL)
CKS1B ProteinP61024 (Uniprot-TrEMBL)
CKS1BProteinP61024 (Uniprot-TrEMBL)
CMPMetaboliteCHEBI:17361 (ChEBI)
CUL1 ProteinQ13616 (Uniprot-TrEMBL)
Cdc25 A/BREACT_9277 (Reactome)
CoA-SHMetaboliteCHEBI:15346 (ChEBI)
Cohesin

PDS5 WAPAL

Centromere
ComplexREACT_151885 (Reactome)
Cohesin

PDS5 WAPAL

Chromosomal Arm
ComplexREACT_150643 (Reactome)
Cyclin A

Cdk2

p21/p27 complex
ComplexREACT_9335 (Reactome)
Cyclin A

Cdk2

phospho-p27/p21 complex
ComplexREACT_9145 (Reactome)
Cyclin A

Cdk2

phosphorylated substrate complex
ComplexREACT_9299 (Reactome)
Cyclin A

Cdk2

substrate complex
ComplexREACT_9192 (Reactome)
Cyclin A Cdk2 complexComplexREACT_4932 (Reactome)
Cyclin A Cdk2 complexComplexREACT_9350 (Reactome)
Cyclin A phospho-Cdk2ComplexREACT_9202 (Reactome)
Cyclin A phospho-Cdk2ComplexREACT_9292 (Reactome)
Cyclin AProteinREACT_6541 (Reactome)
Cyclin D1 Cdk4ComplexREACT_2478 (Reactome)
Cyclin E/A

Cdk2 multiubiquitinated phospho-p27/p21

SCF
ComplexREACT_9266 (Reactome)
Cyclin E/A

Cdk2

p27/p21 complex
ComplexREACT_9314 (Reactome)
Cyclin E/A

Cdk2 phospho-p27/p21

SCF
ComplexREACT_9193 (Reactome)
Cyclin E/A

Cdk2

phospho-p27/p21
ComplexREACT_9321 (Reactome)
Cyclin E/A Cdk2ComplexREACT_9091 (Reactome)
DNA Polymerase delta tetramerComplexREACT_5801 (Reactome)
DNA polymerase alpha

primase DNA polymerase alpha

origin complex
ComplexREACT_3167 (Reactome)
DNA polymerase alpha primaseComplexREACT_3725 (Reactome)
DNA polymerase epsilonComplexREACT_4621 (Reactome)
DNA2 ProteinP51530 (Uniprot-TrEMBL)
DNA2ProteinP51530 (Uniprot-TrEMBL)
ESCOProteinREACT_151079 (Reactome)
FEN1ProteinP39748 (Uniprot-TrEMBL)
FZR1 ProteinQ9UM11 (Uniprot-TrEMBL)
GINS complexComplexREACT_7704 (Reactome)
GINS1 ProteinQ14691 (Uniprot-TrEMBL)
GINS1ProteinQ14691 (Uniprot-TrEMBL)
GINS2 ProteinQ9Y248 (Uniprot-TrEMBL)
GINS2ProteinQ9Y248 (Uniprot-TrEMBL)
GINS4ProteinQ9BRT9 (Uniprot-TrEMBL)
GMPMetaboliteCHEBI:17345 (ChEBI)
H2OMetaboliteCHEBI:15377 (ChEBI)
LIG1ProteinP18858 (Uniprot-TrEMBL)
MAXProteinP61244 (Uniprot-TrEMBL)
MCM2 ProteinP49736 (Uniprot-TrEMBL)
MCM2ProteinP49736 (Uniprot-TrEMBL)
MCM3 ProteinP25205 (Uniprot-TrEMBL)
MCM3ProteinP25205 (Uniprot-TrEMBL)
MCM4ProteinP33991 (Uniprot-TrEMBL)
MCM5 ProteinP33992 (Uniprot-TrEMBL)
MCM5ProteinP33992 (Uniprot-TrEMBL)
MCM6 ProteinQ14566 (Uniprot-TrEMBL)
MCM7 ProteinP33993 (Uniprot-TrEMBL)
MCM8 ProteinQ9UJA3 (Uniprot-TrEMBL)
MCM8ProteinQ9UJA3 (Uniprot-TrEMBL)
MNAT1 ProteinP51948 (Uniprot-TrEMBL)
MYC ProteinP01106 (Uniprot-TrEMBL)
Mcm2-7 complexComplexREACT_4763 (Reactome)
Mcm4,6,7 complexComplexREACT_3276 (Reactome)
Mitotic PrometaphasePathwayREACT_682 (Reactome) 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.
Mitotic Telophase/CytokinesisPathwayREACT_1932 (Reactome) In this final phase of mitosis, new membranes are formed around two sets of chromatids and two daughter cells are formed. The chromosomes and the spindle fibers disperse, and the fiber ring around the center of the cell, composed of actin, contracts, pinching the cell into two daughter cells.
Myc/Max heterodimerComplexREACT_9233 (Reactome)
NTPMetaboliteREACT_4491 (Reactome)
ORC1 ProteinQ13415 (Uniprot-TrEMBL)
ORC2 ProteinQ13416 (Uniprot-TrEMBL)
ORC3 ProteinQ9UBD5 (Uniprot-TrEMBL)
ORC4 ProteinO43929 (Uniprot-TrEMBL)
ORC5 ProteinO43913 (Uniprot-TrEMBL)
ORC6 ProteinQ9Y5N6 (Uniprot-TrEMBL)
Okazaki fragmentREACT_4386 (Reactome)
PCNA ProteinP12004 (Uniprot-TrEMBL)
PCNA homotrimerComplexREACT_2542 (Reactome)
PDS5A ProteinQ29RF7 (Uniprot-TrEMBL)
PDS5B ProteinQ9NTI5 (Uniprot-TrEMBL)
POLA1 ProteinP09884 (Uniprot-TrEMBL)
POLA2 ProteinQ14181 (Uniprot-TrEMBL)
POLD1 ProteinP28340 (Uniprot-TrEMBL)
POLD2 ProteinP49005 (Uniprot-TrEMBL)
POLD3 ProteinQ15054 (Uniprot-TrEMBL)
POLD4 ProteinQ9HCU8 (Uniprot-TrEMBL)
POLE ProteinQ07864 (Uniprot-TrEMBL)
POLE2 ProteinP56282 (Uniprot-TrEMBL)
PRIM1 ProteinP49642 (Uniprot-TrEMBL)
PRIM2 ProteinP49643 (Uniprot-TrEMBL)
PSF3pREACT_7682 (Reactome)
PSMA1 ProteinP25786 (Uniprot-TrEMBL)
PSMA2 ProteinP25787 (Uniprot-TrEMBL)
PSMA3 ProteinP25788 (Uniprot-TrEMBL)
PSMA4 ProteinP25789 (Uniprot-TrEMBL)
PSMA5 ProteinP28066 (Uniprot-TrEMBL)
PSMA6 ProteinP60900 (Uniprot-TrEMBL)
PSMA7ProteinO14818 (Uniprot-TrEMBL)
PSMA8 ProteinQ8TAA3 (Uniprot-TrEMBL)
PSMB1 ProteinP20618 (Uniprot-TrEMBL)
PSMB10 ProteinP40306 (Uniprot-TrEMBL)
PSMB11 ProteinA5LHX3 (Uniprot-TrEMBL)
PSMB2 ProteinP49721 (Uniprot-TrEMBL)
PSMB3 ProteinP49720 (Uniprot-TrEMBL)
PSMB4 ProteinP28070 (Uniprot-TrEMBL)
PSMB5 ProteinP28074 (Uniprot-TrEMBL)
PSMB6 ProteinP28072 (Uniprot-TrEMBL)
PSMB7 ProteinQ99436 (Uniprot-TrEMBL)
PSMB8 ProteinP28062 (Uniprot-TrEMBL)
PSMB9 ProteinP28065 (Uniprot-TrEMBL)
PSMC1ProteinP62191 (Uniprot-TrEMBL)
PSMC2 ProteinP35998 (Uniprot-TrEMBL)
PSMC3 ProteinP17980 (Uniprot-TrEMBL)
PSMC4 ProteinP43686 (Uniprot-TrEMBL)
PSMC5 ProteinP62195 (Uniprot-TrEMBL)
PSMC6ProteinP62333 (Uniprot-TrEMBL)
PSMD1 ProteinQ99460 (Uniprot-TrEMBL)
PSMD10 ProteinO75832 (Uniprot-TrEMBL)
PSMD11 ProteinO00231 (Uniprot-TrEMBL)
PSMD12 ProteinO00232 (Uniprot-TrEMBL)
PSMD13 ProteinQ9UNM6 (Uniprot-TrEMBL)
PSMD14 ProteinO00487 (Uniprot-TrEMBL)
PSMD2 ProteinQ13200 (Uniprot-TrEMBL)
PSMD3 ProteinO43242 (Uniprot-TrEMBL)
PSMD4ProteinP55036 (Uniprot-TrEMBL)
PSMD5 ProteinQ16401 (Uniprot-TrEMBL)
PSMD6ProteinQ15008 (Uniprot-TrEMBL)
PSMD7ProteinP51665 (Uniprot-TrEMBL)
PSMD8 ProteinP48556 (Uniprot-TrEMBL)
PSMD9 ProteinO00233 (Uniprot-TrEMBL)
PSME1 ProteinQ06323 (Uniprot-TrEMBL)
PSME2 ProteinQ9UL46 (Uniprot-TrEMBL)
PSME3 ProteinP61289 (Uniprot-TrEMBL)
PSME4 ProteinQ14997 (Uniprot-TrEMBL)
PSMF1ProteinQ92530 (Uniprot-TrEMBL)
PiMetaboliteCHEBI:18367 (ChEBI)
Processive complex

Okazaki fragment Flap RPA heterotrimer

dna2
ComplexREACT_3597 (Reactome)
Processive complex

Okazaki fragment Flap

RPA heterotrimer
ComplexREACT_4699 (Reactome)
Processive complex

Okazaki fragment

Flap
ComplexREACT_4984 (Reactome)
Processive complex Okazaki fragment complexComplexREACT_5537 (Reactome)
Processive complex

Okazaki fragments

Remaining Flap
ComplexREACT_4338 (Reactome)
Processive complex nicked DNA from adjacent Okazaki fragmentsComplexREACT_4371 (Reactome)
Processive complexComplexREACT_3120 (Reactome)
RAD21 ProteinO60216 (Uniprot-TrEMBL)
RB1 ProteinP06400 (Uniprot-TrEMBL)
RFC Heteropentamer

RNA primer-DNA primer origin duplex

PCNA homotrimer
ComplexREACT_5474 (Reactome)
RFC Heteropentamer

RNA primer-DNA primer

origin duplex
ComplexREACT_5241 (Reactome)
RFC HeteropentamerComplexREACT_4881 (Reactome)
RFC1 ProteinP35251 (Uniprot-TrEMBL)
RFC2 ProteinP35250 (Uniprot-TrEMBL)
RFC3 ProteinP40938 (Uniprot-TrEMBL)
RFC4 ProteinP35249 (Uniprot-TrEMBL)
RFC5 ProteinP40937 (Uniprot-TrEMBL)
RNA primer

origin duplex DNA polymerase alpha

primase complex
ComplexREACT_5709 (Reactome)
RNA primer-DNA primer

origin duplex

PCNA
ComplexREACT_4810 (Reactome)
RNA primer-DNA primer origin duplexComplexREACT_5497 (Reactome)
RPA heterotrimerComplexREACT_3427 (Reactome)
RPA1 ProteinP27694 (Uniprot-TrEMBL)
RPA2 ProteinP15927 (Uniprot-TrEMBL)
RPA3 ProteinP35244 (Uniprot-TrEMBL)
RPS27AProteinP62979 (Uniprot-TrEMBL)
Replication ForkREACT_7812 (Reactome)
SCFComplexREACT_9182 (Reactome)
SCFComplexREACT_9339 (Reactome)
SKP1 ProteinP63208 (Uniprot-TrEMBL)
SKP2 ProteinQ13309 (Uniprot-TrEMBL)
SMC1A ProteinQ14683 (Uniprot-TrEMBL)
SMC3 ProteinQ9UQE7 (Uniprot-TrEMBL)
STAG1 ProteinQ8WVM7 (Uniprot-TrEMBL)
STAG2 ProteinQ8N3U4 (Uniprot-TrEMBL)
Sister CentromereREACT_151483 (Reactome)
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)
UBA52ProteinP62987 (Uniprot-TrEMBL)
UBBProteinP0CG47 (Uniprot-TrEMBL)
UBCProteinP0CG48 (Uniprot-TrEMBL)
UMPMetaboliteCHEBI:16695 (ChEBI)
UbProteinREACT_3316 (Reactome)
Ubiquitin ligaseREACT_4282 (Reactome)
Unwinding complex at replication forkComplexREACT_7007 (Reactome)
Unwound forkREACT_7610 (Reactome)
WAPAL ProteinQ7Z5K2 (Uniprot-TrEMBL)
WEE1ProteinP30291 (Uniprot-TrEMBL)
anaphase-promoting complex REACT_3388 (Reactome)
dATPMetaboliteCHEBI:16284 (ChEBI)
dCTPMetaboliteCHEBI:16311 (ChEBI)
dGTPMetaboliteCHEBI:16497 (ChEBI)
dTTPMetaboliteCHEBI:18077 (ChEBI)
glycogen synthase kinase-3 betaREACT_3846 (Reactome)
ligated okazaki fragmentREACT_4394 (Reactome)
multi-ubiquitinated phospho-ComplexREACT_7367 (Reactome)
p-CDC6ProteinQ99741 (Uniprot-TrEMBL)
p-FZR1 ProteinQ9UM11 (Uniprot-TrEMBL)
p-RB1 ProteinP06400 (Uniprot-TrEMBL) The pRB C-terminus contains a cluster of seven candidate in vivo cdk phosphorylation sites (residues 795, 807, 811, 821, and 826) and is phosphorylated in vitro by cyclin A, cyclin E, and cyclin D-associated kinases.
p-S,T-ORC1ProteinQ13415 (Uniprot-TrEMBL)
p-S130-CDKN1A ProteinP38936 (Uniprot-TrEMBL)
p-T160-CDK2 ProteinP24941 (Uniprot-TrEMBL)
p-T187-CDKN1B ProteinP46527 (Uniprot-TrEMBL)
p-T286-CCND1 ProteinP24385 (Uniprot-TrEMBL)
p-T286-CCND1ProteinP24385 (Uniprot-TrEMBL)
p-Y15-CDK2 ProteinP24941 (Uniprot-TrEMBL)
p21/p27ProteinREACT_8306 (Reactome)
phosphoComplexREACT_4903 (Reactome)
phosphoComplexREACT_5446 (Reactome)
pre-replicative complex ComplexREACT_3695 (Reactome)
pre-replicative complexComplexREACT_4388 (Reactome)
ubiquitinProteinREACT_3995 (Reactome)
ubiquitinated Cdc6ComplexREACT_5308 (Reactome)
ubiquitinated Orc1ComplexREACT_5173 (Reactome)
ubiquitinated Orc1ComplexREACT_5402 (Reactome)

Annotated Interactions

View all...
SourceTargetTypeDatabase referenceComment
26S proteasomemim-catalysisREACT_1210 (Reactome)
26S proteasomemim-catalysisREACT_2142 (Reactome)
26S proteasomemim-catalysisREACT_480 (Reactome)
26S proteasomemim-catalysisREACT_9034 (Reactome)
ADPArrowREACT_1279 (Reactome)
ADPArrowREACT_132 (Reactome)
ADPArrowREACT_1994 (Reactome)
ADPArrowREACT_2111 (Reactome)
ADPArrowREACT_8998 (Reactome)
ADPArrowREACT_9007 (Reactome)
ADPArrowREACT_9057 (Reactome)
ADPArrowREACT_9070 (Reactome)
AMPArrowREACT_315 (Reactome)
ATPREACT_1279 (Reactome)
ATPREACT_132 (Reactome)
ATPREACT_1673 (Reactome)
ATPREACT_1994 (Reactome)
ATPREACT_2111 (Reactome)
ATPREACT_8998 (Reactome)
ATPREACT_9007 (Reactome)
ATPREACT_9057 (Reactome)
ATPREACT_9070 (Reactome)
Ac-CoAREACT_150144 (Reactome)
Ac-CoAREACT_150456 (Reactome)
Ac-Cohesin

PDS5 WAPAL

Centromere
ArrowREACT_150144 (Reactome)
Ac-Cohesin

PDS5 WAPAL

Centromere
REACT_150445 (Reactome)
Ac-Cohesin

PDS5 WAPAL

Chromosomal Arm
ArrowREACT_150456 (Reactome)
Ac-Cohesin

PDS5 WAPAL

Chromosomal Arm
REACT_150264 (Reactome)
CAKmim-catalysisREACT_9070 (Reactome)
CDC45REACT_6963 (Reactome)
CDC6ArrowREACT_1279 (Reactome)
CDC6REACT_1279 (Reactome)
CDCA5REACT_150264 (Reactome)
CDCA5REACT_150445 (Reactome)
CDK2REACT_8996 (Reactome)
CDKmim-catalysisREACT_1279 (Reactome)
CKS1BREACT_9017 (Reactome)
CMPArrowREACT_315 (Reactome)
Cdc25 A/Bmim-catalysisREACT_9062 (Reactome)
CoA-SHArrowREACT_150144 (Reactome)
CoA-SHArrowREACT_150456 (Reactome)
Cohesin

PDS5 WAPAL

Centromere
REACT_150144 (Reactome)
Cohesin

PDS5 WAPAL

Chromosomal Arm
REACT_150456 (Reactome)
Cyclin A

Cdk2

p21/p27 complex
mim-catalysisREACT_8995 (Reactome)
Cyclin A

Cdk2

phosphorylated substrate complex
ArrowREACT_9007 (Reactome)
Cyclin A

Cdk2

substrate complex
REACT_9007 (Reactome)
Cyclin A

Cdk2

substrate complex
mim-catalysisREACT_2111 (Reactome)
Cyclin A

Cdk2

substrate complex
mim-catalysisREACT_9007 (Reactome)
Cyclin A Cdk2 complexArrowREACT_9062 (Reactome)
Cyclin A Cdk2 complexREACT_9005 (Reactome)
Cyclin A Cdk2 complexREACT_9057 (Reactome)
Cyclin A Cdk2 complexREACT_9070 (Reactome)
Cyclin A phospho-Cdk2ArrowREACT_9057 (Reactome)
Cyclin A phospho-Cdk2ArrowREACT_9070 (Reactome)
Cyclin A phospho-Cdk2REACT_9062 (Reactome)
Cyclin AREACT_8996 (Reactome)
Cyclin D1 Cdk4REACT_132 (Reactome)
Cyclin E/A

Cdk2 multiubiquitinated phospho-p27/p21

SCF
mim-catalysisREACT_9026 (Reactome)
Cyclin E/A

Cdk2

p27/p21 complex
REACT_8998 (Reactome)
Cyclin E/A

Cdk2

p27/p21 complex
mim-catalysisREACT_8998 (Reactome)
Cyclin E/A

Cdk2 phospho-p27/p21

SCF
REACT_9026 (Reactome)
Cyclin E/A

Cdk2

phospho-p27/p21
ArrowREACT_8998 (Reactome)
Cyclin E/A

Cdk2

phospho-p27/p21
REACT_9060 (Reactome)
Cyclin E/A Cdk2ArrowREACT_9034 (Reactome)
DNA Polymerase delta tetramerREACT_1009 (Reactome)
DNA Polymerase delta tetramerREACT_149 (Reactome)
DNA Polymerase delta tetramermim-catalysisREACT_1024 (Reactome)
DNA polymerase alpha

primase DNA polymerase alpha

origin complex
REACT_1611 (Reactome)
DNA polymerase alpha primasemim-catalysisREACT_1611 (Reactome)
DNA polymerase alpha primasemim-catalysisREACT_91 (Reactome)
DNA polymerase epsilonArrowREACT_1611 (Reactome)
DNA2ArrowREACT_315 (Reactome)
DNA2REACT_1278 (Reactome)
ESCOmim-catalysisREACT_150144 (Reactome)
ESCOmim-catalysisREACT_150456 (Reactome)
FEN1mim-catalysisREACT_2024 (Reactome)
GINS complexREACT_6963 (Reactome)
GINS1REACT_6747 (Reactome)
GINS2REACT_6747 (Reactome)
GINS4REACT_6747 (Reactome)
GMPArrowREACT_315 (Reactome)
H2OREACT_9062 (Reactome)
LIG1mim-catalysisREACT_1889 (Reactome)
MCM2ArrowREACT_1303 (Reactome)
MCM3ArrowREACT_1303 (Reactome)
MCM5ArrowREACT_1303 (Reactome)
MCM8mim-catalysisREACT_6768 (Reactome)
Mcm2-7 complexREACT_6963 (Reactome)
Mcm2-7 complexmim-catalysisREACT_6922 (Reactome)
Mcm4,6,7 complexArrowREACT_1303 (Reactome)
Myc/Max heterodimerArrowREACT_9013 (Reactome)
NTPREACT_1611 (Reactome)
PCNA homotrimerREACT_1994 (Reactome)
PSF3pREACT_6747 (Reactome)
PiArrowREACT_9062 (Reactome)
Processive complex

Okazaki fragment Flap

RPA heterotrimer
REACT_1278 (Reactome)
Processive complex

Okazaki fragment

Flap
REACT_2026 (Reactome)
Processive complex

Okazaki fragments

Remaining Flap
ArrowREACT_315 (Reactome)
REACT_1009 (Reactome) The loading of proliferating cell nuclear antigen (PCNA) leads to recruitment of pol delta. Human PCNA is a homotrimer of 36 kDa subunits that form a toroidal structure. The loading of PCNA by RFC is a key event in the transition from the priming mode to the extension mode of DNA synthesis. The processive complex is composed of the pol delta holoenzyme and PCNA.
REACT_1024 (Reactome) After RFC initiates the assembly of the primer recognition complex, the complex of pol delta and PCNA is responsible for incorporating the additional nucleotides prior to the position of the next downstream initiator RNA primer. On the lagging strand, short discontinuous segments of DNA, called Okazaki fragments, are synthesized on RNA primers. The average length of the Okazaki fragments is 100 nucleotides. Polymerase switching is a key event that allows the processive synthesis of DNA by the pol delta and PCNA complex.
REACT_1180 (Reactome) In this reaction, 1 molecule of 'phospho(T286)-Cyclin D1' is translocated from nucleoplasm to cytosol.

This reaction takes place in the 'nuclear envelope'.

REACT_1210 (Reactome) At the beginning of this reaction, 1 molecule of 'ubiquitinated Cdc6' is present. At the end of this reaction, 1 molecule of 'ubiquitin' is present.

This reaction takes place in the 'cytosol' and is mediated by the 'endopeptidase activity' of '26S proteasome'.

REACT_1278 (Reactome) After RPA binds the long flap, it recruits the Dna2 endonuclease. Dna2 endonuclease removes most of the flap, but the job of complete removal of the flap is then completed by FEN-1.
REACT_1279 (Reactome) At the beginning of this reaction, 1 molecule of 'CDC6', and 1 molecule of 'ATP' are present. At the end of this reaction, 1 molecule of 'ADP', and 1 molecule of 'phosphorylated Cdc6' are present.

This reaction takes place in the 'nucleus' and is mediated by the 'kinase activity' of 'CDK'.

REACT_1303 (Reactome) At the start of the elongation phase of DNA replication, the Mcm2-7 complex may re-arrange to function as the replicative helicase associated with the replication fork. In general, a replicative helicase is associated with the replication fork and unwinds DNA ahead of the polymerase. In yeast, the Mcm proteins associate with origin DNA in G1 phase and then exit the origin upon replication initiation, consistent with moving out of the origin with the replication fork. The Mcm2-7 complex is a ring-shaped hexamer. Complexes of Mcm4, Mcm6 and Mcm7 proteins from humans or S. pombe display a modest ATP-dependent helicase activity in vitro. Consistent with the hypothesis that eukaryotic Mcm complexes function as helicases, an archaeal Mcm homolog is a ring-shaped double hexamer that has a processive DNA unwinding activity. Mcm proteins may have additional functions during elongation, as uninterrupted function of all six is required for replication fork progression in budding yeast. Mcm4,6,7 helicase activity may be negatively regulated in two ways. Mcm2, Mcm4, Mcm6, and Mcm7 also form a stable complex which, however, has no helicase activity, suggesting that Mcm2 inhibits DNA unwinding by Mcm4,6,7. In addition, phosphorylation of human Mcm4,6,7 complex by CDK inhibits its helicase activity.
REACT_132 (Reactome) At the beginning of this reaction, 1 molecule of 'Cyclin D1:Cdk4', and 1 molecule of 'ATP' are present. At the end of this reaction, 1 molecule of 'phospho(T286)-Cyclin D1:Cdk4', and 1 molecule of 'ADP' are present.

This reaction takes place in the 'nucleus' and is mediated by the 'kinase activity' of 'glycogen synthase kinase-3 beta'.

REACT_149 (Reactome) Polymerase switching is a key event that allows the processive synthesis of DNA by the pol delta and PCNA complex. Polymerase delta possesses polymerization and proofreading activities, which increases the overall fidelity of DNA replication. The pol delta holoenzyme is a heterotetrameric complex that contains p125, p66, p50, and p12 subunits, in human cells.
REACT_150144 (Reactome) Acetyltransferases ESCO1 and ESCO2 are homologs of the S. cerevisiae acetyltransferase Eco1, essential for viability in yeast. ESCO1 and ESCO2 share sequence homology in the C-terminal region, consisting of a H2C2 zinc finger motif and an acetyltransferase domain (Hou and Zou 2005). Both ESCO1 and ESCO2 acetylate the cohesin subunit SMC3 on two lysine residues, K105 and K106 (Zhang et al. 2008), an important step in the establishment of sister-chromatid cohesion during the S-phase of the cell cycle. Divergent N-termini of ESCO1 and ESCO2, necessary for chromatin binding, suggest that ESCO1 and ESCO2 may perform distinct functions in sister chromatid cohesion (Hou and Zou 2005). Several studies suggest that ESCO2 may be predominantly involved in acetylation of the SMC3 subunit of centromeric cohesin. A conditional targeting of Esco2 locus in mice leads to pre-implantational loss of homozygous Esco2 -/- embryos at the eight-cell stage. Prometaphase chromosomes isolated from two-cell stage Esco2 knockout embryos show marked cohesion defect at centromeres (Whelan et al. 2012). ESCO2 protein appears in the S-phase (Hou and Zou 2005, Whelan et al. 2012) and in mouse embryonic fibroblasts Esco2 predominantly localizes to pericentric heterochromatin (Whelan et al. 2012). Mutations in the ESCO2 gene (Vega et al. 2005) that impair ESCO2 acetyltransferase activity (Gordillo et al. 2008) are the cause of the Roberts syndrome, an autosomal recessive disorder characterized by craniofacial and limb abnormalities, and intellectual disability. Metaphase chromosomes of Roberts syndrome patients exhibit loss of cohesion at heterochromatic regions of centromeres and the Y chromosome, with a characteristic 'railroad track appearance' (Van den Berg and Francke 1993, Vega et al. 2005).
REACT_150264 (Reactome) CDCA5 (Sororin) is essential for the establishment of sister chromatid cohesion in mammalian cells (Rankin et al. 2005) in the S-phase of the cell cycle (Nishiyama et al. 2010). Several factors contribute to the recruitment of CDCA5 to chromatin-associated cohesin: DNA replication (i.e. presence of two sister chromatids), association of cohesin complex with PDS5, and acetylation of the SMC3 cohesin subunit by ESCO1/ESCO2 acetyltransferases. Experiments in which a recombinant tagged mouse CDCA5 was expressed in human HeLa cell line showed that CDCA5 starts to accumulate on chromatin in S-phase and dissociates from chromosomal arms in prophase (Nishiyama et al. 2010).

CDCA5 is essential for the establishment of chromosomal cohesion only in the presence of WAPAL, suggesting that the key role of CDCA5 (Sororin) is to antagonize WAPAL. Both CDCA5 and WAPAL contain an FGF (phenylalanine-glycine-phenylalanine) motif that is essential for PDS5 binding and is also essential for CDCA5 function in cohesion establishment. Indeed, CDCA5 is able to displace WAPAL from PDS5:WAPAL heterodimers in vitro. In vivo experiments in Xenopus egg extracts suggest that CDCA5 rearranges the topology of cohesin associated proteins so that WAPAL is no longer able to inhibit sister chromatid cohesion but remains associated with cohesin (Nishiyama et al. 2010).
REACT_150445 (Reactome) CDCA5 (Sororin) is essential for the establishment of sister chromatid cohesion at centromeres. Experiments in which a recombinant tagged mouse CDCA5 was expressed in human HeLa cell line showed that CDCA5 starts to accumulate on chromatin in S-phase and dissociates from centromeres in anaphase (Nishiyama et al. 2010).
REACT_150456 (Reactome) Acetyltransferases ESCO1 and ESCO2 are homologs of the S. cerevisiae acetyltransferase Eco1, essential for viability in yeast. ESCO1 and ESCO2 share sequence homology in the C-terminal region, consisting of a H2C2 zinc finger motif and an acetyltransferase domain (Hou and Zou 2005). Both ESCO1 and ESCO2 acetylate the cohesin subunit SMC3 on two lysine residues, K105 and K106 (Zhang et al. 2008), an important step in the establishment of sister-chromatid cohesion during the S-phase of the cell cycle. These dual acetylations on SMC3 are deacetylated by HDAC8 after the cohesin removal from chromatin for the dissociation and recycling of cohesin subunits (Deardorff et al. 2012). ESCO1 and ESCO2 differ in their N-termini, which are necessary for chromatin binding, and may perform distinct functions in sister chromatid cohesion (Hou and Zou 2005), as suggested by the study of Esco2 knockout mice (Whelan et al. 2012).
REACT_1572 (Reactome) When the polymerase delta:PCNA complex reaches a downstream Okazaki fragment, strand displacement synthesis occurs. The primer containing 5'-terminus of the downstream Okazaki fragment is folded into a single-stranded flap.
REACT_1611 (Reactome) At the beginning of this reaction, 1 molecule of 'DNA polymerase alpha:primase:DNA polymerase alpha:origin complex', and 1 molecule of 'NTP' are present. At the end of this reaction, 1 molecule of 'DNA polymerase epsilon', and 1 molecule of 'RNA primer:origin duplex:DNA polymerase alpha:primase complex' are present.

This reaction takes place in the 'nucleus' and is mediated by the 'DNA-directed RNA polymerase activity' of 'DNA polymerase alpha:primase'.

REACT_1626 (Reactome) At the beginning of this reaction, 1 molecule of 'ubiquitin', and 1 molecule of 'phosphorylated Orc1' are present. At the end of this reaction, 1 molecule of 'ubiquitinated Orc1' is present.

This reaction takes place in the 'nucleus'.

REACT_1673 (Reactome) At the beginning of this reaction, 1 molecule of 'phosphorylated Cdc6', 1 molecule of 'ubiquitin', and 1 molecule of 'ATP' are present. At the end of this reaction, 1 molecule of 'ubiquitinated Cdc6' is present.

This reaction takes place in the 'cytosol' and is mediated by the 'endopeptidase activity' of 'anaphase-promoting complex (APC)'.

REACT_1871 (Reactome) Once the RNA-DNA primer is synthesized, replication factor C (RFC) initiates a reaction called "polymerase switching"; pol delta, the processive enzyme replaces pol alpha, the priming enzyme. RFC binds to the 3'-end of the RNA-DNA primer on the Primosome, to displace the pol alpha primase complex. The binding of RFC triggers the binding of the primer recognition complex.
REACT_1889 (Reactome) Removal of the flap by FEN-1 leads to the generation of a nick between the 3'-end of the upstream Okazaki fragment and the 5'-end of the downstream Okazaki fragment. DNA ligase I then seals the nicks between adjacent processed Okazaki fragments to generate intact double-stranded DNA.
REACT_1994 (Reactome) The binding of the primer recognition complex involves the loading of proliferating cell nuclear antigen (PCNA). Replication Factor C transiently opens the PCNA toroid in an ATP-dependent reaction, and then allows PCNA to re-close around the double helix adjacent to the primer terminus. This leads to the formation of the "sliding clamp".
REACT_2024 (Reactome) The remaining flap, which is too short to support RPA binding, is then processed by FEN-1. There is evidence that binding of RPA to the displaced end of the RNA-containing Okazaki fragment prevents FEN-1 from accessing the substrate. FEN-1 is a structure-specific endonuclease that cleaves near the base of the flap at a position one nucleotide into the annealed region. Biochemical studies have shown that the preferred substrate for FEN-1 consists of a one-nucleotide 3'-tail on the upstream primer in addition to the 5'-flap of the downstream primer.
REACT_2026 (Reactome) The first step in the removal of the flap intermediate is the binding of Replication Protein A (RPA) to the long flap structure. RPA is a eukaryotic single-stranded DNA binding protein.
REACT_210 (Reactome) In this reaction, 1 molecule of 'ubiquitinated Orc1' is translocated from nucleoplasm to cytosol.

This movement of the molecule occurs through the 'nuclear pore'.

REACT_2111 (Reactome) At the beginning of this reaction, 1 molecule of 'ATP', and 1 molecule of 'pre-replicative complex' are present. At the end of this reaction, 1 molecule of 'phosphorylated Orc1', 1 molecule of 'pre-replicative complex (Orc1-minus)', and 1 molecule of 'ADP' are present.

This reaction takes place in the 'nucleus' and is mediated by the 'kinase activity' of 'Cyclin A:Cdk2 complex'.

REACT_2142 (Reactome) Phosphorylated Cyclin D1 is degraded during S phase by the 26S proteasome allowing for efficient DNA synthesis.
REACT_315 (Reactome) The Dna2 endonuclease removes the initiator RNA along with several downstream deoxyribonucleotides. The cleavage of the single-stranded RNA substrate results in the disassembly of RPA and Dna2. The current data for the role of the Dna2 endonuclease has been derived from studies with yeast and Xenopus Dna2.
REACT_351 (Reactome) Replication factor C is proposed to dissociate from PCNA following sliding clamp formation, and the DNA toroid alone tethers pol delta to the DNA.
REACT_448 (Reactome) In this reaction, 1 molecule of 'phosphorylated Cdc6' is translocated from nucleoplasm to cytosol.

This movement of the molecule occurs through the 'nuclear pore'.

REACT_480 (Reactome) At the beginning of this reaction, 1 molecule of 'ubiquitinated Orc1' is present. At the end of this reaction, 1 molecule of 'ubiquitin' is present.

This reaction takes place in the 'cytosol' and is mediated by the 'endopeptidase activity' of '26S proteasome'.

REACT_6747 (Reactome) At the beginning of this reaction, 1 molecule of 'PSF3p', 1 molecule of 'SLD5P', 1 molecule of 'PSF2p', and 1 molecule of 'PSF1p' are present. At the end of this reaction, 1 molecule of 'GINS complex' is present.

This reaction takes place in the 'nucleus'.

REACT_6768 (Reactome) The MCM2-7 related protein, MCM8, is required to replicate chromosomal DNA in Xenopus egg extracts. MCM8 binds chromatin upon initiation of DNA synthesis. It may function as an helicase in the elongation step.
REACT_6922 (Reactome) In budding yeast, all MCM proteins have been proved to be essential for elongation. The active form of this protein complex may be a heterohexamer. A subcomplex of MCM proteins consisting fo MCM4,6, and -7 has a weak helicase activity that may contribute to DNA unwinding.
REACT_6963 (Reactome) By applying the chromatin immunoprecipitation technique to paused forks, certain proteins like DNA pol alpha, DNA pol delta, DNA pol epsilon, MCM2-7, CDC45, GINS and MCM10 were identified. By uncoupling a helicase at the site using a polymerase inhibitor, MCM2-7, GINS complex and CDC45 alone were found to be enriched at the paused fork suggesting these proteins may form a part of an "unwindosome" at the replicating fork.
REACT_715 (Reactome) Cyclin D is targeted for degradation by multi-ubiquitination.
REACT_752 (Reactome) In this reaction, 1 molecule of 'phospho(T286)-Cyclin D1:Cdk4' is translocated from nucleoplasm to cytosol.

This reaction takes place in the 'nuclear envelope'.

REACT_8995 (Reactome) Recognition of p27 by SCF(Skp2) and the subsequent ubiquitination of p27 is dependent upon Cyclin E/A:Cdk2-mediated phosphorylation of p27 at Thr 187 (Montagnoli et al., 1999). p21 is also phosphorylated at a specific site (Ser130) by Cyclin E/A:Cdk2, stimulating its ubiquitination. Unlike p27, however, p21 ubiquitination can take place in the absence of phosphorylation, although with less efficiency (Bornstein et al.,2003).
REACT_8996 (Reactome) During G1 phase of the cell cycle, cyclin A is synthesized and associates with Cdk2.
REACT_8998 (Reactome) The interaction between the Skp2 subunit of the SCF(Skp2) complex and p27 is dependent upon Cdk2:Cyclin A/E mediated phosphorylation of p27 at Thr 187 (Carrano et al, 1999; Tsvetkov et al, 1999). There is evidence that Cyclin A/B:Cdk1 can also bind and phosphorylate p27 on Thr 187 (Nakayama et al., 2004). This phosphorylation is also essential for the subsequent ubiquitination of p27.
REACT_9005 (Reactome) During G1, the activity of cyclin-dependent kinases (CDKs) is kept in check by the CDK inhibitors (CKIs) p27 and p21, thereby preventing premature entry into S phase (see Guardavaccaro and Pagano, 2006).
REACT_9007 (Reactome) Active Cyclin A:Cdk2 complexes phosphorylate and inactivate proteins required for maintaining the G1/S phase including: Cdh1, RB1, p21 and p27. All this creates auto-amplification loops that render Cdk2 increasingly more active. In G2, Cdk2, in association with cyclin A, phosphorylates E2F1 and E2F3 resulting in the inactivation and possibly degradation of these two transcription factors (Dynlacht et al., 1994; Krek et al., 1994).
REACT_9013 (Reactome) The Myc/Max heterodimer binds to elements in the cdc25A gene and activates transcription in mid to late G1.
REACT_9017 (Reactome) The accessory protein, Cks1 promotes efficient interaction between phosphorylated p27 and the SCF (Skp2) complex (Ganoth et al., 2001; Spruck et al., 2001). Cks1 binds to Skp2 in the leucine-rich repeat (LRR) domain and C-terminal tail (Hao et al., 2005). The phosphorylated Thr187 side chain of p27 associates with a phosphate binding site on Cks1, and the side chain containing Glu185 is positioned in the interface between Skp2 and Cks1 where it interacts with both (Hao et al., 2005).
REACT_9026 (Reactome) Once in tight contact with the SCF (Skp2):Cks1 complex, phosphorylated p27/p21 is ubiquitinated.
REACT_9034 (Reactome) Following ubiquitination by the SCF(Skp2):Cks1 complex, phospho-p27/p21 is degraded by the 26S proteasome.
REACT_9043 (Reactome) p27 translocates to the nucleoplasm where it associates with CyclinE:Cdk2 complexes. Localization of p27 to the nucleus is necessary to inhibit Cdk activation by Cdk-activating kinase.
REACT_9057 (Reactome) The CDK activity of the Cyclin A:Cdk2 complex is inhibited by phosphorylation at Tyr 15, presumably by the Wee1 kinase.
REACT_9060 (Reactome) The association of Cks1 with both Skp2 and phosphorylated p27 promotes a tight interaction between p27 and the SCF complex (Hao et al., 2005).
REACT_9062 (Reactome) Cdc25A, and probably Cdc25B, regulate the entry into S phase cell cycle by removing inhibitory phosphates from the Cdk2 subunit of Cyclin A:Cdk2.
REACT_9065 (Reactome) After forming in the cytoplasm, the Cyclin A:Cdk2 complexes are translocated to the nucleus.
REACT_9068 (Reactome) p21 associates with and inhibits Cyclin:Cdk complexes in the nucleus.
REACT_9070 (Reactome) Phosphorylation of cyclin-dependent kinases (CDKs) by the CDK-activating kinase (CAK) is required for the activation of the CDK kinase activity. The association of p21/p27 with the Cyclin A/E:Cdk2 complex prevents CAK mediated phosphorylation of Cdk2 (Aprelikova et al., 1995).
REACT_91 (Reactome) At the beginning of this reaction, 1 molecule of 'dTTP', 1 molecule of 'dGTP', 1 molecule of 'dATP', 1 molecule of 'RNA primer:origin duplex:DNA polymerase alpha:primase complex', and 1 molecule of 'dCTP' are present. At the end of this reaction, 1 molecule of 'RNA primer-DNA primer:origin duplex' is present.

This reaction takes place in the 'nucleus' and is mediated by the 'DNA-directed DNA polymerase activity' of 'DNA polymerase alpha:primase'.

RFC Heteropentamer

RNA primer-DNA primer origin duplex

PCNA homotrimer
ArrowREACT_1994 (Reactome)
RFC Heteropentamer

RNA primer-DNA primer

origin duplex
REACT_1994 (Reactome)
RFC HeteropentamerArrowREACT_351 (Reactome)
RFC HeteropentamerREACT_1871 (Reactome)
RNA primer

origin duplex DNA polymerase alpha

primase complex
ArrowREACT_1611 (Reactome)
RNA primer

origin duplex DNA polymerase alpha

primase complex
REACT_91 (Reactome)
RNA primer-DNA primer

origin duplex

PCNA
ArrowREACT_351 (Reactome)
RNA primer-DNA primer

origin duplex

PCNA
REACT_1009 (Reactome)
RNA primer-DNA primer

origin duplex

PCNA
REACT_149 (Reactome)
RNA primer-DNA primer origin duplexREACT_1871 (Reactome)
RPA heterotrimerArrowREACT_315 (Reactome)
RPA heterotrimerREACT_2026 (Reactome)
Replication ForkREACT_6963 (Reactome)
SCFArrowREACT_9034 (Reactome)
SCFREACT_9017 (Reactome)
SCFREACT_9060 (Reactome)
Sister CentromereREACT_150445 (Reactome)
Sister Chromosomal ArmREACT_150264 (Reactome)
UMPArrowREACT_315 (Reactome)
UbREACT_1673 (Reactome)
UbREACT_715 (Reactome)
Ubiquitin ligasemim-catalysisREACT_715 (Reactome)
WEE1mim-catalysisREACT_9057 (Reactome)
anaphase-promoting complex mim-catalysisREACT_1673 (Reactome)
dATPREACT_91 (Reactome)
dCTPREACT_91 (Reactome)
dGTPREACT_91 (Reactome)
dTTPREACT_91 (Reactome)
glycogen synthase kinase-3 betamim-catalysisREACT_132 (Reactome)
p-CDC6REACT_1673 (Reactome)
p-S,T-ORC1ArrowREACT_2111 (Reactome)
p-S,T-ORC1REACT_1626 (Reactome)
p-T286-CCND1REACT_715 (Reactome)
p21/p27REACT_9005 (Reactome)
p21/p27mim-catalysisREACT_9005 (Reactome)
phosphoArrowREACT_132 (Reactome)
pre-replicative complex ArrowREACT_2111 (Reactome)
pre-replicative complexREACT_2111 (Reactome)
ubiquitinArrowREACT_9034 (Reactome)
ubiquitinREACT_1626 (Reactome)
ubiquitinREACT_9026 (Reactome)
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