(and subcomplexes) / Protein names used in this article / Alternative names / Organisms / Phosphorylation sites / Kinase / Function of phosphorylation / Ref.
Inner centromere / Nucleosome / Histone H3 / H. sapiens,
X. laevis,
S. pombe / T3 / Haspin / Survivin binding / (Dai, et al. 2005)(Wang, et al. 2010)(Yamagishi, et al. 2010)(Kelly, et al. 2010)
Histone H2A / H. sapiens,
M. musculus / T120 / Bub1 / Recruitment of Sgo1 (mitosis) and Sgo2 (mouse meiosis) / (Kawashima, et al. 2010)
S. pombe / S121 / Bub1 / Sgo2 recruitment / (Kawashima, et al. 2010)
S. cerevisiae / S121 / Bub1 / Sgo1 recruitment / (Kawashima, et al. 2010)
-- / Haspin / H. sapiens / S93, S108, S143, S185, S216, T231, S269, T295, S366, S387, S430 / Aurora B / Support effective H3T3 phosphorylation / (Wang, et al. 2011)
Chromosomal passenger complex (CPC) / Aurora B / Aurora B / H. sapiens / T232 / Aurora B / Kinase activation / (Yasui, et al. 2004)
S331 / Chk1 (and unknown) / INCENP phosphorylation / (Petsalaki, et al. 2011)
xAurora B / X. laevis / T248 / Aurora B / Kinase activation / (Eyers, et al. 2005)(Sessa, et al. 2005)(Tseng, et al. 2010)
Ipl1 / S. cerevisiae / T260 / Aurora B (Ipl1) / Kinase activation / (Cheeseman, et al. 2002)(Zimniak, et al. 2012)
S50, S76 / Cdk1(Cdc28) / Inhibiting binding to Bim1, which supports recruitment of the CPC to spindle / (Zimniak, et al. 2012)
INCENP / INCENP / H. sapiens / T893, S894, S895 / Aurora B / Aurora B kinase activation / (Honda, et al. 2003)
T59 / Cdk1 / Inhibiting binding to Mklp2 and premature association with spindle / (Goto, et al. 2006)(Hummer and Mayer 2009)
T388 / Cdk1 / Recruitment of Plk1 to kinetochores / (Goto, et al. 2006)
xINCENP / X. laevis / T848, S849, S850 / Aurora B kinase activation / (Knowlton, et al. 2006)(Kelly, et al. 2007)(Sessa, et al. 2005)
ICP1 / C. elegans / S598, S599 / Aurora B(AIR-2) / Aurora B kinase activation / (Bishop and Schumacher 2002)
Sli15 / S. cerevisiae / S335, S373, S427, S437, S462, T474 / Cdk1 (Cdc28) / Inhibiting targeting the CPC to spindle during pre-anaphase, and inhibiting SAC reactivation during anaphase / (Pereira and Schiebel 2003)(Mirchenko and Uhlmann 2010)
S137, S177, S276, T277, S298, S305, S326, S382, S383, S391, T395, S413, S432, S460, T564, S578, T589 / Aurora B (Ipl1) / Inhibiting targeting the CPC to spindle / (Nakajima, et al. 2011)
Survivin / Survivin, BIRC5 / H. sapiens /
T117
/ Aurora B* / Suppressing Survivin and Aurora B localization at the centromere / (Wheatley, et al. 2004)(Wheatley, et al. 2007)T34 / Cdk1 (Cdc2) / Supporting apoptosis** / (O'Connor, et al. 2000)(Barrett, et al. 2009)
Bir1, Cut17 / S. pombe / S244 and S202, S229, S278, S294, T393, T831, T908 / Cdk1 (Cdc2) / Sgo2 binding / (Tsukahara, et al. 2010)
Bir1 / S. cerevisiae / S383, S395, S552, S587, S667, T684, S688, T735, T747 / Cdk1 (Cdc28) / Ndc10 binding and anaphase spindle elongation / (Widlund, et al. 2006)
Borealin / Dasra B, CDCA8 / H. sapiens / T106, T172, T185, T189, T199, T204, S219 / Cdk1 / Sgo1 and Sgo2 binding, and centromere targeting of the CPC / (Tsukahara, et al. 2010)(Kaur, et al. 2010)
T88, T94, T169, T230 / Mps1 / Supporting Aurora B activity on kinetochore substrates for kinetochore microtubule regulation, but not the SAC*** / (Jelluma, et al. 2008)(Bourhis, et al. 2009)
Shugoshin-PP2A complex / Sgo2 / Tripin / H. sapiens / S26, T29, S53, T76, S146, S236, S247, S266, S278 / Aurora B / Supporting PP2A binding / (Tanno, et al. 2010)
T537 (T620, T720, S816, T914) / Aurora B / Supporting MCAK binding / (Tanno, et al. 2010)
-- / MCAK / MCAK (Kinesin-13) / H. sapiens / S92, S106, S108, S112, S186 / Aurora B / Suppressing microtubule depolymerization activity of MCAK, and supporting MCAK localization at the inner centromere / (Andrews, et al. 2004)
XKCM1 / X. laevis / S196, / Aurora B / Suppressing microtubule depolymerization activity of MCAK, inhibiting MCAK binding to chromosome arms / (Lan, et al. 2004)(Zhang, et al. 2007)(Ohi, et al. 2004)
T95 / Aurora B / Promoting chromosome arm binding of MCAK / (Zhang, et al. 2007)
S110 / Aurora B / Promoting centromere targeting of MCAK / (Zhang, et al. 2007)
S719 / Aurora A* / Promoting spindle pole localization / (Zhang, et al. 2008)
Inner kinetochore / CENP-A nucleosome / CENP-A / Cen-H3 / H. sapiens / Ser7 / Aurora B / Cytokinesis?
Chromosome alignment? / (Zeitlin, et al. 2001)(Kunitoku, et al. 2003)
Aurora A / Aurora B recruitment at the centromere? / (Kunitoku, et al. 2003)
Constitutive centromere-associated network (CCAN) / CENP-O/P/Q/R/U group, CAD, CENP-H/I interacting complex / CENP-U / PBIP1, MLF1IP, KLIP1 / H. sapiens / S77, T78 / Plk1 / Plk1 recruitment to kinetochore / (Kang, et al. 2006)
S349, S350*
/Aurora B*
/ Compromising Hec1 interaction and kinetochore-microtubule attachment / (Hua, et al. 2011)CENP-50 /
G. gallus
/ S62, T63 / Plk1 / CENP-Q and Plk1 recruitment to kinetochore / (Hori, et al. 2008)CENP-T/W/S/X / CENP-T / CENP-T / H. sapiens / T11, T27, S47, T85, T195 / Cdk1 / Ndc80 complex recruitment / (Gascoigne, et al. 2011)
Outer kinetochore / KMN network / Mis12 complex (MIND) / Dsn1 / HsMis13/KNL-3 / H. sapiens / S100, S109 / Aurora B / Supporting chromosome alignment by compromising microtubule attachment / (Yang, et al. 2008)(Welburn, et al. 2010)
KNL-3 / C. elegans /
T61, R133
/ Aurora B / (Welburn, et al. 2010)-- / KNL1 / hKNL1, Blinkin / H. sapiens / S24, S33, S60 / Aurora B / Supporting chromosome alignment by compromising microtubule attachment. Weakening PP1-interaction. / (Welburn, et al. 2010)
T875 (S1037, T1087, T1089, S1022, T1075, T1129, T1154, T1167) / Mps1 / Recruiting Bub1 to support the SAC and bipolar kinetochore-microtubule attachment / (Yamagishi, et al. 2012)
xKNL1 / X. laevis / S23, S54, S58 / Aurora B / Weakening PP1-interaction. / (Rosenberg, et al. 2011)
KNL-1 / C. elegans / S11, S41,
Spc7 / S. pombe / T257 (T77, S221, T338, T366, T395, T413, T422, T453, T507, T529, T552) / Mps1 (Mph1) / Recruiting Bub1 to support the SAC and bipolar kinetochore-microtubule attachment / (Yamagishi, et al. 2012)(Shepperd, et al. 2012)
Spc105 / S. cerevisiae / T149, T172, T211, T235, T284, T313 / Mps1 / Recruiting Bub1 to support the SAC and normal chromosome segregation / (London, et al.)
Ndc80 complex / Ndc80 / Hec1 / H. sapiens / S8, S44, S55 (S4, S5, S15, T49, S62, S69) / Aurora B / Weakening kinetochore-microtubule attachment / (DeLuca, et al. 2006)(Deluca, et al. 2011)(Ciferri, et al. 2008)
S165 / Nek2 / Supporting Mad1 and Mad2 recruitment to unattached kinetochores and normal chromosome segregation. / (Wei, et al. 2011)
NDC-80 / C. elegans / T8, S18, S44, S51 / Aurora B / Weakening microtubule binding / (Cheeseman, et al. 2006)
Ndc80 / S. cerevisiae / S37, T54, T74, S100 (T21, T71, S95) / Aurora B (Ipl1) / Nonessential, auxiliary roles on SAC activation and chromosome segregation / (Akiyoshi, et al. 2009)(Cheeseman, et al. 2002)
S4, T5, S6, T21, S22, S37, T38, T43, T74, T79, T82, S205, T248, T252 / Mps1 / Maintenance of the SAC / (Kemmler, et al. 2009)
Dam1 complex / Dam1 / Dam1 / S. cerevisiae / S20 / Aurora B (Ipl1) / Inhibiting binding to the Ndc80 complex, and supporting proper chromosome segregation / (Cheeseman, et al. 2002)(Gestaut, et al. 2008)(Tien, et al. 2010)(Keating, et al. 2009)
S257, S265, S292 / Aurora B (Ipl1) / Compromising oligomerization of the Dam1 complex, and supporting proper chromosome segregation. / (Cheeseman, et al. 2002)(Keating, et al. 2009)(Wang, et al. 2007)
S218, S221 / Mps1* / Phosphorylation site mutants are synthetic lethal with spc110-226. Mildly supporting the stable kinetochore-microtubule attachment. / (Shimogawa, et al. 2006)
S. pombe / S143 / Plk1 / Mildly supporting bioriented kinetochore-microtubule attachment. / (Buttrick, et al. 2012)
Ask1 / S. cerevisiae / S216, S250 / Cdk1 (Cdc28) / Auxiliary role to support Ask1 function. / (Li and Elledge 2003)
Ska complex / Ska1 / H. sapiens / T157, S185, T205, S242 / Aurora B / Inhibiting kinetochore recruitment of the Ska complex and destablize kinetochore-microtubule attachment / (Chan, et al. 2012)
Ska3 / H. sapiens / S110, S159 / Aurora B / Inhibiting kinetochore recruitment of the Ska complex and destablize kinetochore-microtubule attachment / (Chan, et al. 2012)
-- / Zwint-1 / H. sapiens / S250, T251, S262 / Aurora B* / Kinetochore recruitment of RZZ complex, Dynein, Dynactin, and Mad2, supporting chromosome alignment and SAC maintenance. / (Kasuboski, et al. 2011)
Dynein/dynactin / Dynein intermediate chain / H. sapiens / T89 / Plk1 / Decreasing binding affinity to p150Glued, increasing binding affinity to zw10, and contributing to dynein recruitment at kinetochores / (Whyte, et al. 2008)(Bader, et al. 2011)
-- / Mps1 / Mps1 / H. sapiens / T686 / Mps1 / Stimulating catalytic activity / (Kang, et al. 2007)(Mattison, et al. 2007)(Chu, et al. 2008)
T12, S15 / Mps1 / Kinetochore recruitment of Mps1 and Mad2, and SAC activation.** / (Xu, et al. 2009)
S821 / unknown / Kinetochore recruitment of Mps1 / (Xu, et al. 2009)
Mps1 / X. laevis / S844 / MAPK / Kinetochore recruitment of Mps1 and SAC activation / (Zhao and Chen 2006)
S283 / Cdk1 / Mps1 kinase activity / (Morin, et al. 2012)
-- / Bub1 / Bub1 / H. sapiens / T609 / Cdk1 / Generating Plk1-binding site, recruiting Plk1 to kinetochores / (Qi, et al. 2006)
xBub1 / X. laevis / T493, S500, S634, T650 / MAPK / Supporting the SAC / (Chen 2004)
Bub1 / S. pombe / T340, T423, T455, S466 / Cdk1 (Cdc2) / Supporting the SAC / (Yamaguchi, et al. 2003)
Bub1 / S.cerevisiae / T566 / Cdk1 (Cdc28) / Promoting Bub1 degradation to adapt the prolonged SAC activation / (Goto, et al. 2011)
Mitotic checkpoint protein complex (MCC) / Mad2 / Mad2 / H. sapiens / S170, S178, S195 / unknown / Disrupting Mad1-Mad2 interaction and/or Mad2-Cdc20 interaction, and promoting SAC silencing / (Wassmann, et al. 2003)(Kim, et al. 2010)
Mad2 / S. pombe / S92, /
Mps1 (Mph1)*
/ Supporting MCC-APC/C interaction / (Zich, et al. 2012)S187 / Mps1 (Mph1)* / Disrupting Mad1-Mad2 interaction and, promoting SAC silencing / (Zich, et al. 2012)
BubR1 / BubR1 / H. sapiens / T620 / Cdk1 / Generating Plk1-binding site, for Plk1-dependent phosphorylation of BubR1. Supporting stable kinetochore-microtubule attachment. / (Elowe, et al. 2007)
S676, T680 / Plk1 / Supporting stable kinetochore-microtubule attachment through recruiting PP2A-B56 / (Elowe, et al. 2007)(Suijkerbuijk, et al. 2012)
T792, T1008 / Plk1 / Supporting stable kinetochore-microtubule attachment / (Matsumura, et al. 2007)
S670 / Mps1 (and other kinases) / Supporting correct kinetochore-microtubule attachment through recruiting PP2A-B56 / (Huang, et al. 2008)(Elowe, et al. 2007; Suijkerbuijk, et al. 2012)
T608 / BubR1-CENP-E / Supporting Aurora B-mediated Ndc80/Hec1 phosphorylation, Mad2 recruitment to unattached kinetochores, chromosome alignment and SAC activation. / (Guo, et al. 2012)
BubR1 / X. laevis / T605 / Cdk1 / Generating Plk1-binding site, recruiting Plk1 and Mad2 to kinetochores, supporting the SAC / (Wong and Fang 2007)
Mad3 / S. cerevisiae / S303, S337 / Aurora B (Ipl1) / Specifically required for SAC induced by lack of kinetochore tension but not by unattached kinetochores. / (King, et al. 2007)
Cdc20 / Cdc20 / H. sapiens / S41, T55, T59, T70, T157, S408, S452 /
Cdk1*
/ Inhibiting Cdc20-APC/C interaction / (Yudkovsky, et al. 2000)S41, S72, S92, S153, T157, S161 /
Bub1*
/ Catalytically inhibit APC/C activity / (Tang, et al. 2004)Cdc20/Fizzy / X. laevis / T64, T68 / MAPK / MCC formation and SAC activation / (Chung and Chen 2003)
S50, T64, T68, T79, S114 / Cdk1 / Supporting Mad2-Cdc20 interaction and inhibiting Cdc20-APC/C interaction / {Chung, 2003 #3793}(D'Angiolella, et al. 2003)(Labit, et al. 2012)
-- / CENP-E / Kinesin-7 / H. sapiens / T422 / Aurora A and Aurora B / Reducing affinity for microtubules and PP1, reducing motor processivity, and controlling chromosome congression / (Kim, et al. 2010)
-- / Kif2b / Kif2b (Kinesin-13) / H. sapiens / T125 / Plk1 / Promoting microtubule depolymerization activity of Kif2b, and correcting erroneous kinetochore-microtubule attachment / (Hood, et al. 2012)
S204 / Plk1 / Kif2b localization at the kinetochore, promoting microtubule depolymerization activity, and correcting erroneous kinetochore-microtubule attachment / (Hood, et al. 2012)
-- / CLIP-170 / H. sapiens / S195 / Plk1 / Promoting CK2 interaction and S1318 phosphorylation / (Li, et al. 2010)
S1318 / CK2 / Promoting dynein/dynactin interaction and kinetochore recruitment, and supporting stable kinetochore-microtubule attachment / (Li, et al. 2010)
-- / PP1 / H. sapiens / T320 / Cdk1 (Cdc2) / Suppression of the phosphatase activity / (Dohadwala, et al. 1994)
Dis2 / S. pombe / T316 / Cdk1 (Cdc2) / Suppression of the phosphatase activity / (Yamano, et al. 1994)
-- / Plk1 / Plk1 / H. sapiens / T210 / Aurora A / Kinase activation / (Seki, et al. 2008)(Macurek, et al. 2008)
Polo / D. melanogaster / T182 / Aurora B / Kinase activation / (Carmena and Earnshaw 2003)
-- / NudC / H. sapiens / S274, S326 / Plk1 / Recruiting Plk1 to kinetochores and supporting end-on kinetochore microtubule attachments and cytokinesis / (Nishino, et al. 2006)(Zhou, et al. 2003)
-- / MYPT1 / H. sapiens / S473 / Cdk1 / Recruiting Plk1 and facilitating Plk1 dephosphorylation and inactivation by PP1 / (Yamashiro, et al. 2008)
-- / mDia3 / H. sapiens / T66, S196 / Aurora B / Activation of mDia3? / (Cheng, et al. 2011)
S820, T882 / Aurora B / Reducing microtubule binding and stabilizing activity of mDia3, and supporting proper kinetochore-microtubule attachment / (Cheng, et al. 2011)
-- / Nsk1 / S. pombe / S71, S103, S113, S140, S171, S195, S206, S221, S236, T240, T255, S257, S289, S344, S379, S399, T411, T437 / Cdk1 (Cdc2)* / Suppressing kinetochore binding and correcting erroneous kinetochore-microtubule attachment / (Chen, et al. 2011)
Italic residues indicate predicted phosphorylation sites, whose individual phosphorylation in vivo has not been established.
* Only in vitro phosphorylation data are currently available.
** Contradictory to these reports, this phosphorylation site does not influence apoptosis in chicken DT cells (Yue, et al. 2008)
*** Contradictory to these reports, it was shown that Mps1 activity is not required for kinetochore localization of Mps1 (Maciejowski, et al. 2010)(Hewitt, et al. 2010).
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