Mitotic Prometaphase (Homo sapiens)

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Bibliography

1. Okada M, Cheeseman IM, Hori T, Okawa K, McLeod IX, Yates JR, Desai A, Fukagawa T.; ''The CENP-H-I complex is required for the efficient incorporation of newly synthesized CENP-A into centromeres.''; PubMed Europe PMC Scholia
2. Chen D, Ito S, Yuan H, Hyodo T, Kadomatsu K, Hamaguchi M, Senga T.; ''EML4 promotes the loading of NUDC to the spindle for mitotic progression.''; PubMed Europe PMC Scholia
3. Takemoto A, Kimura K, Yanagisawa J, Yokoyama S, Hanaoka F.; ''Negative regulation of condensin I by CK2-mediated phosphorylation.''; PubMed Europe PMC Scholia
4. Kotak S, Busso C, Gönczy P.; ''NuMA phosphorylation by CDK1 couples mitotic progression with cortical dynein function.''; PubMed Europe PMC Scholia
5. Hsu HL, Yeh NH.; ''Dynamic changes of NuMA during the cell cycle and possible appearance of a truncated form of NuMA during apoptosis.''; PubMed Europe PMC Scholia
6. Compton DA, Luo C.; ''Mutation of the predicted p34cdc2 phosphorylation sites in NuMA impair the assembly of the mitotic spindle and block mitosis.''; PubMed Europe PMC Scholia
7. Houtman SH, Rutteman M, De Zeeuw CI, French PJ.; ''Echinoderm microtubule-associated protein like protein 4, a member of the echinoderm microtubule-associated protein family, stabilizes microtubules.''; PubMed Europe PMC Scholia
8. Roig J, Mikhailov A, Belham C, Avruch J.; ''Nercc1, a mammalian NIMA-family kinase, binds the Ran GTPase and regulates mitotic progression.''; PubMed Europe PMC Scholia
9. St-Pierre J, Douziech M, Bazile F, Pascariu M, Bonneil E, Sauvé V, Ratsima H, D'Amours D.; ''Polo kinase regulates mitotic chromosome condensation by hyperactivation of condensin DNA supercoiling activity.''; PubMed Europe PMC Scholia
10. Hauf S, Waizenegger IC, Peters JM.; ''Cohesin cleavage by separase required for anaphase and cytokinesis in human cells.''; PubMed Europe PMC Scholia
11. Kimura K, Cuvier O, Hirano T.; ''Chromosome condensation by a human condensin complex in Xenopus egg extracts.''; PubMed Europe PMC Scholia
12. Salic A, Waters JC, Mitchison TJ.; ''Vertebrate shugoshin links sister centromere cohesion and kinetochore microtubule stability in mitosis.''; PubMed Europe PMC Scholia
13. Shintomi K, Hirano T.; ''Releasing cohesin from chromosome arms in early mitosis: opposing actions of Wapl-Pds5 and Sgo1.''; PubMed Europe PMC Scholia
14. Richards MW, O'Regan L, Roth D, Montgomery JM, Straube A, Fry AM, Bayliss R.; ''Microtubule association of EML proteins and the EML4-ALK variant 3 oncoprotein require an N-terminal trimerization domain.''; PubMed Europe PMC Scholia
15. Zhou T, Aumais JP, Liu X, Yu-Lee LY, Erikson RL.; ''A role for Plk1 phosphorylation of NudC in cytokinesis.''; PubMed Europe PMC Scholia
16. Belham C, Roig J, Caldwell JA, Aoyama Y, Kemp BE, Comb M, Avruch J.; ''A mitotic cascade of NIMA family kinases. Nercc1/Nek9 activates the Nek6 and Nek7 kinases.''; PubMed Europe PMC Scholia
17. Losada A, Hirano M, Hirano T.; ''Identification of Xenopus SMC protein complexes required for sister chromatid cohesion.''; PubMed Europe PMC Scholia
18. Nishiyama T, Ladurner R, Schmitz J, Kreidl E, Kreidl E, Schleiffer A, Bhaskara V, Bando M, Shirahige K, Hyman AA, Mechtler K, Peters JM.; ''Sororin mediates sister chromatid cohesion by antagonizing Wapl.''; PubMed Europe PMC Scholia
19. Zhang N, Panigrahi AK, Mao Q, Pati D.; ''Interaction of Sororin protein with polo-like kinase 1 mediates resolution of chromosomal arm cohesion.''; PubMed Europe PMC Scholia
20. Cheeseman IM, Desai A.; ''Molecular architecture of the kinetochore-microtubule interface.''; PubMed Europe PMC Scholia
21. Watrin E, Schleiffer A, Tanaka K, Eisenhaber F, Nasmyth K, Peters JM.; ''Human Scc4 is required for cohesin binding to chromatin, sister-chromatid cohesion, and mitotic progression.''; PubMed Europe PMC Scholia
22. Lipp JJ, Hirota T, Poser I, Peters JM.; ''Aurora B controls the association of condensin I but not condensin II with mitotic chromosomes.''; PubMed Europe PMC Scholia
23. Kitajima TS, Sakuno T, Ishiguro K, Iemura S, Natsume T, Kawashima SA, Watanabe Y.; ''Shugoshin collaborates with protein phosphatase 2A to protect cohesin.''; PubMed Europe PMC Scholia
24. Abe S, Nagasaka K, Hirayama Y, Kozuka-Hata H, Oyama M, Aoyagi Y, Obuse C, Hirota T.; ''The initial phase of chromosome condensation requires Cdk1-mediated phosphorylation of the CAP-D3 subunit of condensin II.''; PubMed Europe PMC Scholia
25. Ono T, Fang Y, Spector DL, Hirano T.; ''Spatial and temporal regulation of Condensins I and II in mitotic chromosome assembly in human cells.''; PubMed Europe PMC Scholia
26. Kueng S, Hegemann B, Peters BH, Lipp JJ, Schleiffer A, Mechtler K, Peters JM.; ''Wapl controls the dynamic association of cohesin with chromatin.''; PubMed Europe PMC Scholia
27. Kimura K, Hirano M, Kobayashi R, Hirano T.; ''Phosphorylation and activation of 13S condensin by Cdc2 in vitro.''; PubMed Europe PMC Scholia
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

External references

DataNodes

Name	Type	Database reference	Comment
2xAcK-SMC3 [chromosome, centromeric region]	Protein	Q9UQE7 (Uniprot-TrEMBL)
2xAcK-SMC3 [chromosome]	Protein	Q9UQE7 (Uniprot-TrEMBL)
2xAcK-SMC3 [cytosol]	Protein	Q9UQE7 (Uniprot-TrEMBL)
ADP	Metabolite	CHEBI:16761 (ChEBI)
AHCTF1 [cytosol]	Protein	Q8WYP5 (Uniprot-TrEMBL)
APITD1 [cytosol]	Protein	Q8N2Z9 (Uniprot-TrEMBL)
ATP	Metabolite	CHEBI:15422 (ChEBI)
AURKB(2-344) [cytosol]	Protein	Q96GD4 (Uniprot-TrEMBL)
Ac-CoA	Metabolite	CHEBI:15351 (ChEBI)
Ac-Cohesin:PDS5:CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules	Complex	REACT_151019 (Reactome)
Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules	Complex	REACT_152183 (Reactome)
B9D2 [cytosol]	Protein	Q9BPU9 (Uniprot-TrEMBL)
BIRC5 [cytosol]	Protein	O15392 (Uniprot-TrEMBL)
BUB1 [cytosol]	Protein	O43683 (Uniprot-TrEMBL)
BUB1B [cytosol]	Protein	O60566 (Uniprot-TrEMBL)
BUB3 [cytosol]	Protein	O43684 (Uniprot-TrEMBL)
CASC5 [cytosol]	Protein	Q8NG31 (Uniprot-TrEMBL)
CCNB1 [cytosol]	Protein	P14635 (Uniprot-TrEMBL)
CCNB2 [cytosol]	Protein	O95067 (Uniprot-TrEMBL)
CCNB:p-T161-CDK1	Complex	REACT_148199 (Reactome)
CDC20 [cytosol]	Protein	Q12834 (Uniprot-TrEMBL)
CDCA5 [chromosome, centromeric region]	Protein	Q96FF9 (Uniprot-TrEMBL)
CDCA5 [chromosome]	Protein	Q96FF9 (Uniprot-TrEMBL)
CDCA8 [cytosol]	Protein	Q53HL2 (Uniprot-TrEMBL)
CDK1 Phosphorylated Condensin I	Complex	REACT_150472 (Reactome)
CENPA [cytosol]	Protein	P49450 (Uniprot-TrEMBL)
CENPC1 [cytosol]	Protein	Q03188 (Uniprot-TrEMBL)
CENPE [cytosol]	Protein	Q02224 (Uniprot-TrEMBL)
CENPF [cytosol]	Protein	P49454 (Uniprot-TrEMBL)
CENPH [cytosol]	Protein	Q9H3R5 (Uniprot-TrEMBL)
CENPI [cytosol]	Protein	Q92674 (Uniprot-TrEMBL)
CENPK [cytosol]	Protein	Q9BS16 (Uniprot-TrEMBL)
CENPL [cytosol]	Protein	Q8N0S6 (Uniprot-TrEMBL)
CENPM [cytosol]	Protein	Q9NSP4 (Uniprot-TrEMBL)
CENPN [cytosol]	Protein	Q96H22 (Uniprot-TrEMBL)
CENPO [cytosol]	Protein	Q9BU64 (Uniprot-TrEMBL)
CENPP [cytosol]	Protein	Q6IPU0 (Uniprot-TrEMBL)
CENPQ [cytosol]	Protein	Q7L2Z9 (Uniprot-TrEMBL)
CENPT(2-561) [cytosol]	Protein	Q96BT3 (Uniprot-TrEMBL)
CK2 Phosphorylated Condensin I	Complex	REACT_151678 (Reactome)
CKAP5 [cytosol]	Protein	Q14008 (Uniprot-TrEMBL)
CLASP1 [cytosol]	Protein	Q7Z460 (Uniprot-TrEMBL)
CLASP2 [cytosol]	Protein	O75122 (Uniprot-TrEMBL)
CLIP1 [cytosol]	Protein	P30622 (Uniprot-TrEMBL)
CSNK2A1 [cytosol]	Protein	P68400 (Uniprot-TrEMBL)
CSNK2A2 [cytosol]	Protein	P19784 (Uniprot-TrEMBL)
CSNK2B(1-215) [cytosol]	Protein	P67870 (Uniprot-TrEMBL)
Casein kinase II	Complex	REACT_23098 (Reactome)
CoA-SH	Metabolite	CHEBI:15346 (ChEBI)
Condensed prometaphase chromosomes		REACT_150762 (Reactome)
Condensed prophase chromosomes		REACT_151302 (Reactome)
Condensin I	Complex	REACT_151162 (Reactome)
DSN1 [cytosol]	Protein	Q9H410 (Uniprot-TrEMBL)
ERCC6L [cytosol]	Protein	Q2NKX8 (Uniprot-TrEMBL)
H2O	Metabolite	CHEBI:15377 (ChEBI)
HDAC8	Protein	REACT_151192 (Reactome)
INCENP [cytosol]	Protein	Q9NQS7 (Uniprot-TrEMBL)
ITGB3BP [cytosol]	Protein	Q13352 (Uniprot-TrEMBL)
KIF18A [cytosol]	Protein	Q8NI77 (Uniprot-TrEMBL)
KIF2A [cytosol]	Protein	O00139 (Uniprot-TrEMBL)
KIF2B [cytosol]	Protein	Q8N4N8 (Uniprot-TrEMBL)
KIF2C [cytosol]	Protein	Q99661 (Uniprot-TrEMBL)
KNTC1 [cytosol]	Protein	P50748 (Uniprot-TrEMBL)
Kinetochore	Complex	REACT_14970 (Reactome)
MAD1L1 [cytosol]	Protein	Q9Y6D9 (Uniprot-TrEMBL)
MAD2L1 [cytosol]	Protein	Q13257 (Uniprot-TrEMBL)
MAPRE1 [cytosol]	Protein	Q15691 (Uniprot-TrEMBL)
MIS12 [cytosol]	Protein	Q9H081 (Uniprot-TrEMBL)
MLF1IP [cytosol]	Protein	Q71F23 (Uniprot-TrEMBL)
Microtubule-bound kinetochore	Complex	REACT_15175 (Reactome)
Mitotic Metaphase and Anaphase	Pathway	WP2757 (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 [cytosol]	Protein	Q15021 (Uniprot-TrEMBL)
NCAPG [cytosol]	Protein	Q9BPX3 (Uniprot-TrEMBL)
NCAPH [cytosol]	Protein	Q15003 (Uniprot-TrEMBL)
NDC80 [cytosol]	Protein	O14777 (Uniprot-TrEMBL)
NDE1 [cytosol]	Protein	Q9NXR1 (Uniprot-TrEMBL)
NDEL1 [cytosol]	Protein	Q9GZM8 (Uniprot-TrEMBL)
NSL1 [cytosol]	Protein	Q96IY1 (Uniprot-TrEMBL)
NUDC [cytosol]	Protein	Q9Y266 (Uniprot-TrEMBL)
NUF2 [cytosol]	Protein	Q9BZD4 (Uniprot-TrEMBL)
NUP107 [cytosol]	Protein	P57740 (Uniprot-TrEMBL)
NUP133 [cytosol]	Protein	Q8WUM0 (Uniprot-TrEMBL)
NUP160 [cytosol]	Protein	Q12769 (Uniprot-TrEMBL)
NUP37 [cytosol]	Protein	Q8NFH4 (Uniprot-TrEMBL)
NUP43 [cytosol]	Protein	Q8NFH3 (Uniprot-TrEMBL)
NUP85 [cytosol]	Protein	Q9BW27 (Uniprot-TrEMBL)
NUP98-5 [cytosol]	Protein	P52948-5 (Uniprot-TrEMBL)
PAFAH1B1(2-410) [cytosol]	Protein	P43034 (Uniprot-TrEMBL)
PDS5A [chromosome, centromeric region]	Protein	Q29RF7 (Uniprot-TrEMBL)
PDS5A [chromosome]	Protein	Q29RF7 (Uniprot-TrEMBL)
PDS5A [cytosol]	Protein	Q29RF7 (Uniprot-TrEMBL)
PDS5B [chromosome, centromeric region]	Protein	Q9NTI5 (Uniprot-TrEMBL)
PDS5B [chromosome]	Protein	Q9NTI5 (Uniprot-TrEMBL)
PDS5B [cytosol]	Protein	Q9NTI5 (Uniprot-TrEMBL)
PLK1 [cytosol]	Protein	P53350 (Uniprot-TrEMBL)
PLK1	Protein	P53350 (Uniprot-TrEMBL)
PMF1(2-205) [cytosol]	Protein	Q6P1K2 (Uniprot-TrEMBL)
PPP1CC [cytosol]	Protein	P36873 (Uniprot-TrEMBL)
Pi	Metabolite	CHEBI:18367 (ChEBI)
RAD21 [chromosome, centromeric region]	Protein	O60216 (Uniprot-TrEMBL)
RAD21 [chromosome]	Protein	O60216 (Uniprot-TrEMBL)
RANBP2 [cytosol]	Protein	P49792 (Uniprot-TrEMBL)
RANGAP1 [cytosol]	Protein	P46060 (Uniprot-TrEMBL)
RCC2 [cytosol]	Protein	Q9P258 (Uniprot-TrEMBL)
RPS27 [cytosol]	Protein	P42677 (Uniprot-TrEMBL)
S Phase	Pathway	WP2772 (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 [cytosol]	Protein	P55735 (Uniprot-TrEMBL)
SEH1L-1 [cytosol]	Protein	Q96EE3-1 (Uniprot-TrEMBL)
SGOL1 [cytosol]	Protein	Q5FBB7 (Uniprot-TrEMBL)
SGOL2 [cytosol]	Protein	Q562F6 (Uniprot-TrEMBL)
SKA1(2-255) [cytosol]	Protein	Q96BD8 (Uniprot-TrEMBL)
SKA2 [cytosol]	Protein	Q8WVK7 (Uniprot-TrEMBL)
SMC1A [chromosome, centromeric region]	Protein	Q14683 (Uniprot-TrEMBL)
SMC1A [chromosome]	Protein	Q14683 (Uniprot-TrEMBL)
SMC1A [cytosol]	Protein	Q14683 (Uniprot-TrEMBL)
SMC2 [cytosol]	Protein	O95347 (Uniprot-TrEMBL)
SMC3 [cytosol]	Protein	Q9UQE7 (Uniprot-TrEMBL)
SMC4 [cytosol]	Protein	Q9NTJ3 (Uniprot-TrEMBL)
SPC24 [cytosol]	Protein	Q8NBT2 (Uniprot-TrEMBL)
SPC25 [cytosol]	Protein	Q9HBM1 (Uniprot-TrEMBL)
SPDL1 [cytosol]	Protein	Q96EA4 (Uniprot-TrEMBL)
STAG1 [chromosome, centromeric region]	Protein	Q8WVM7 (Uniprot-TrEMBL)
STAG1 [chromosome]	Protein	Q8WVM7 (Uniprot-TrEMBL)
STAG1 [cytosol]	Protein	Q8WVM7 (Uniprot-TrEMBL)
STAG2 [chromosome, centromeric region]	Protein	Q8N3U4 (Uniprot-TrEMBL)
STAG2 [chromosome]	Protein	Q8N3U4 (Uniprot-TrEMBL)
Sister Centromeres:Ac-Cohesin:PDS5:CDCA5:WAPAL	Complex	REACT_151874 (Reactome)
Sister Chromosomal Arm		REACT_150847 (Reactome)
Sister Chromosomal Arms:Ac-Cohesin:PDS5:CDCA5:WAPAL	Complex	REACT_150576 (Reactome)
Sister Chromosomal Arms:Ac-Cohesin:PDS5:p-CDCA5:WAPAL	Complex	REACT_151061 (Reactome)
Sister Chromosomal Arms:p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL	Complex	REACT_151301 (Reactome)
TAOK1 [cytosol]	Protein	Q7L7X3 (Uniprot-TrEMBL)
Unknown Phosphatase		REACT_150933 (Reactome)
WAPAL [chromosome, centromeric region]	Protein	Q7Z5K2 (Uniprot-TrEMBL)
WAPAL [chromosome]	Protein	Q7Z5K2 (Uniprot-TrEMBL)
WAPAL [cytosol]	Protein	Q7Z5K2 (Uniprot-TrEMBL)
XPO1 [cytosol]	Protein	O14980 (Uniprot-TrEMBL)
ZW10 [cytosol]	Protein	O43264 (Uniprot-TrEMBL)
ZWILCH [cytosol]	Protein	Q9H900 (Uniprot-TrEMBL)
ZWINT [cytosol]	Protein	O95229 (Uniprot-TrEMBL)
microtubule		REACT_10446 (Reactome)
p-Ac-Cohesin:PDS5:WAPAL	Complex	REACT_151851 (Reactome)
p-Cohesin:PDS5:WAPAL	Complex	REACT_151704 (Reactome)
p-RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules	Complex	REACT_151499 (Reactome)
p-S-NCAPD2 [cytosol]	Protein	Q15021 (Uniprot-TrEMBL)
p-S-NCAPG [cytosol]	Protein	Q9BPX3 (Uniprot-TrEMBL)
p-S-SMC4 [cytosol]	Protein	Q9NTJ3 (Uniprot-TrEMBL)
p-S21,S75,T159-CDCA5 [chromosome, centromeric region]	Protein	Q96FF9 (Uniprot-TrEMBL)
p-S21,S75,T159-CDCA5 [chromosome]	Protein	Q96FF9 (Uniprot-TrEMBL)
p-S21,S75,T159-CDCA5	Protein	Q96FF9 (Uniprot-TrEMBL)
p-S454-RAD21 [chromosome, centromeric region]	Protein	O60216 (Uniprot-TrEMBL)
p-S454-RAD21 [chromosome]	Protein	O60216 (Uniprot-TrEMBL)
p-S454-RAD21 [cytosol]	Protein	O60216 (Uniprot-TrEMBL)
p-S570-NCAPH [cytosol]	Protein	Q15003 (Uniprot-TrEMBL)
p-STAG2 [chromosome, centromeric region]	Protein	Q8N3U4 (Uniprot-TrEMBL)
p-STAG2 [chromosome]	Protein	Q8N3U4 (Uniprot-TrEMBL)
p-STAG2 [cytosol]	Protein	Q8N3U4 (Uniprot-TrEMBL)
p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules	Complex	REACT_150708 (Reactome)
p-T-NCAPH [cytosol]	Protein	Q15003 (Uniprot-TrEMBL)
p-T1339,T1384,T1389-NCAPD2 [cytosol]	Protein	Q15021 (Uniprot-TrEMBL)
p-T161-CDK1 [cytosol]	Protein	P06493 (Uniprot-TrEMBL)
p-T308,T332-NCAPG [cytosol]	Protein	Q9BPX3 (Uniprot-TrEMBL)

Annotated Interactions

Source	Target	Type	Database reference	Comment
	ADP	Arrow	REACT_150152 (Reactome)
	ADP	Arrow	REACT_150232 (Reactome)
	ADP	Arrow	REACT_150239 (Reactome)
	ADP	Arrow	REACT_150246 (Reactome)
	ADP	Arrow	REACT_150414 (Reactome)
	ADP	Arrow	REACT_150454 (Reactome)
ATP			REACT_150152 (Reactome)
ATP			REACT_150232 (Reactome)
ATP			REACT_150239 (Reactome)
ATP			REACT_150246 (Reactome)
ATP			REACT_150414 (Reactome)
ATP			REACT_150454 (Reactome)
	Ac-CoA	Arrow	REACT_150215 (Reactome)
	Ac-Cohesin:PDS5:CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules	Arrow	REACT_150388 (Reactome)
Ac-Cohesin:PDS5:CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules			REACT_150414 (Reactome)
	Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules	Arrow	REACT_150414 (Reactome)
Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules			REACT_150152 (Reactome)
CCNB:p-T161-CDK1		mim-catalysis	REACT_150232 (Reactome)
CCNB:p-T161-CDK1		mim-catalysis	REACT_150414 (Reactome)
CCNB:p-T161-CDK1		mim-catalysis	REACT_150454 (Reactome)
	CDK1 Phosphorylated Condensin I	Arrow	REACT_150232 (Reactome)
CDK1 Phosphorylated Condensin I		Arrow	REACT_150278 (Reactome)
	CK2 Phosphorylated Condensin I	Arrow	REACT_150239 (Reactome)
CK2 Phosphorylated Condensin I			REACT_150403 (Reactome)
Casein kinase II		mim-catalysis	REACT_150239 (Reactome)
CoA-SH			REACT_150215 (Reactome)
	Condensed prometaphase chromosomes	Arrow	REACT_150278 (Reactome)
Condensed prophase chromosomes			REACT_150278 (Reactome)
	Condensin I	Arrow	REACT_150403 (Reactome)
Condensin I			REACT_150232 (Reactome)
Condensin I			REACT_150239 (Reactome)
H2O			REACT_150250 (Reactome)
H2O			REACT_150403 (Reactome)
HDAC8		mim-catalysis	REACT_150215 (Reactome)
Kinetochore			REACT_14798 (Reactome)
	Microtubule-bound kinetochore	Arrow	REACT_14798 (Reactome)
Microtubule-bound kinetochore			REACT_150388 (Reactome)
PLK1		mim-catalysis	REACT_150152 (Reactome)
PLK1		mim-catalysis	REACT_150246 (Reactome)
	Pi	Arrow	REACT_150250 (Reactome)
	Pi	Arrow	REACT_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 Arm	Arrow	REACT_150267 (Reactome)
Sister Chromosomal Arms:Ac-Cohesin:PDS5:CDCA5:WAPAL			REACT_150454 (Reactome)
	Sister Chromosomal Arms:Ac-Cohesin:PDS5:p-CDCA5:WAPAL	Arrow	REACT_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	Arrow	REACT_150246 (Reactome)
Sister Chromosomal Arms:p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL			REACT_150267 (Reactome)
Unknown Phosphatase		mim-catalysis	REACT_150403 (Reactome)
microtubule			REACT_14798 (Reactome)
	p-Ac-Cohesin:PDS5:WAPAL	Arrow	REACT_150267 (Reactome)
p-Ac-Cohesin:PDS5:WAPAL			REACT_150215 (Reactome)
	p-Cohesin:PDS5:WAPAL	Arrow	REACT_150215 (Reactome)
	p-RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules	Arrow	REACT_150250 (Reactome)
	p-S21,S75,T159-CDCA5	Arrow	REACT_150267 (Reactome)
	p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules	Arrow	REACT_150152 (Reactome)
p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules			REACT_150250 (Reactome)
p-STAG2,RAD21-Ac-Cohesin:PDS5:p-CDCA5:WAPAL:Sister Centromeres:Kinetochores:Microtubules		mim-catalysis	REACT_150250 (Reactome)