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. Author manuscript; available in PMC: 2023 Apr 7.
Published in final edited form as: Annu Rev Genet. 2022 Sep 2;56:279–314. doi: 10.1146/annurev-genet-072820-034559

Table 1. Parts list of budding yeast and human kinetochores and centromeres.

Protein/component Sub-complex Complex Sub- Notes
H. sapiens S. cerevisiae   assembly
CenpA Cse4 - Nucleosome I Non-specific DNA binding (wraps AT-
rich CDEII in Sc)
Mis18α MIS18C MIS18-HJURP I CenpA loading machinery
Mis18β MIS18C MIS18-HJURP I CenpA loading machinery
Mis18bp MIS18C MIS18-HJURP I CenpA loading machinery
HJURP Scm3 - MIS18-HJURP CenpA chaperone (binds CenpA:H4)
Shugoshin-1 Sgo1^ - - - Cohesion protection; PP2A receptor
Shugoshin-2 Sgo1^ - - - Cohesion protection; PP2A receptor
CenpB n/a - n/a I DNA binding (CenpB box)
n/a Cbf1 - n/a DNA binding (CDEI)
n/a Ndc10 - CBF3 I DNA binding (sequence independent)
n/a Ctf13 CBF3core CBF3 I DNA binding (CDEIII)
n/a Cep3 CBF3core CBF3 I DNA binding (CCG motif in CDEIII)
n/a Skp1 CBF3core CBF3 I F-box protein
CenpC Mif2 - CCAN/Ctf19C II DNA binding (AT hook); Dimer
CenpH McmI6 HIKM CCAN/Ctf19C II
CenpI Ctf3 HIKM CCAN/Ctf19C II
CenpK Mcm22 HIKM CCAN/Ctf19C II
CenpL Iml3 NL CCAN/Ctf19C II
CenpM McmI6 HIKM CCAN/Ctf19C II
CenpN Chl4 NL CCAN/Ctf19C II
CenpO Mcm21 OPQUR/COMA CCAN/Ctf19C II
CenpP Ctf19 OPQUR/COMA CCAN/Ctf19C II
CenpQ Okp1 OPQUR/COMA CCAN/Ctf19C II Ndc80-like MT binding
CenpR n/a OPQUR CCAN/Ctf19C II
CenpS Mhf1* SX TWSX II Histone-fold; also DNA repair role
CenpT Cnn1 TW TWSX II Histone-fold
CenpU Ame1 OPQUR/COMA CCAN/Ctf19C II Receptor for Plk1 (Hs)
CenpV n/a - - GFA domain, CEN chromatin structure;
meiosis, MT binding
CenpW Wip1 TW TWSX II Histone-fold
CenpX Mfh2* SX TWSX II Histone-fold; also DNA repair role
n/a Nkp1 Nkp1/2 CCAN/Ctf19C II
n/a Nkp2 Nkp1/2 CCAN/Ctf19C II
Spc24 Spc24 NDC80 KMN-S III RWD domains
Spc25 Spc25 NDC80 KMN-S III RWD domains
Ndc80 Ndc80 NDC80 KMN-S III CH domain/MT lattice binding
Nuf1 Nuf2 NDC80 KMN-S III CH domain/MT lattice binding
Knl1 Spc105 KNL1 KMN-S III MELT array as platform for SAC; PP1
receptor and MT binding (N-term)
Zwint Kre28 KNL1 KMN-S III
Mis12 Mtw1 MIS12 KMN-S III
Nnf1 (Pmf1) Nnf1 MIS12 KMN-S III
Nsl1 Nsl1 MIS12 KMN-S III
Dsn1 Dsn1 MIS12 KMN-S III Receptor for monopolin
Ska1 n/a SKA KMN-S III Load bearing device; MT Tip tracking -
Ska2 n/a SKA KMN-S III binding curved protofilaments
Ska3 n/a SKA KMN-S III
Cdt1 Tah11* III Also a replication factor; can bind
Ndc80 loop
Ch-TOG Stu2 - - III MT polymerase; docks Ndc80 4-way
junction
n/a Dam1 - Dam1C III MT Encircling-coupler
n/a Duo1 - Dam1C III
n/a Dad1 - Dam1C III
n/a Dad2 - Dam1C III
n/a Dad3 - Dam1C III
n/a Dad4 - Dam1C III
n/a Spc34 - Dam1C III
n/a Spc19 - Dam1C III
n/a Ask1 - Dam1C III
n/a Hsk3 - Dam1C III
Astrin n/a - Astrin/Skap III
Skap n/a - Astrin/Skap III
MYCBP n/a - Astrin/Skap III
LC8 n/a - Astrin/Skap III
Bub1 Bub1 BUB1-BUB3 SAC III Protein kinase
Bub3 Bub3 BUB1-BUB3 SAC III Phospho-MELT binding
BubR1 Mad3 - SAC III Mad3 lacking pseudo-kinase domain
found in BubR1
Mad1 Mad1 MAD1-MAD2 SAC III (& VI) Forms mitotic checkpoint complex
Mad2 Mad2 MAD1-MAD2 SAC III (& VI) (MCC) with BubR1 and Cdc20.
Mps1 Mps1 - SAC III Protein kinase (MELTs, xxx)
p31 comet n/a SAC SAC inhibitor
CenpE n/a - Corona IV Kinesin-7 MT plus-directed molecular
motor
CenpF Slk19 FNNL Corona IV Homology unclear; MT binding and DD
regulator in Hs.
Rod Sec39 and
Sec31*		 RZZ-S Corona IV Required for vesicle tethering in yeast
Zw10 Dsl1* and
Tip20		 RZZ-S Corona IV Required for vesicle tethering in yeast
Zwilch n/a RZZ-S Corona IV
Spindly n/a RZZ-S Corona IV Dynein adapter
- - Cytoplasmic
Dynein*		 DD; Corona IV Minus-directed molecular motor
(Dynein Heavy chain + 5
light/intermediate chains)
- - Dynactin* DD; Corona IV Dynein cofactor (11 subunits)
Lis1 Pac1* FNNL Corona IV Dynein cofactor
Nde1 Ndl1* FNNL Corona IV Dynein cofactor
Ndel1 Ndl1* FNNL Corona IV Dynein cofactor
Clasp Stu1 - Corona IV MT rescue factor
Nup107 Nup84* NUP107-160
(NPC-Y)		 Corona IV Also a nuclear pore component
Nup133 Nup133* NUP107-160
(NPC-Y)		 Corona IV Also a nuclear pore component
Nup96 Nup145C* NUP107-160
(NPC-Y)		 Corona IV Also a nuclear pore component
Sec13 Seh1* NUP107-160
(NPC-Y)		 Corona IV Also a nuclear pore component
Nup160 Nup120* NUP107-160
(NPC-Y)		 Corona IV Also a nuclear pore component
EB1 Bim1 - - - MT end-tracker
HSET† Kar3 - - - Kinesin-14 MT minus-end directed
molecular motor
Kif18a Kip3 - - Kinesin-8 MT plus-end directed
molecular motor and depolymerase;
located on KT proximal K-fibre
- Csm1 - Monopolin - Also in the nucleolus
- Lrs4 - Monopolin - Also in the nucleolus
- Mam1 - Monopolin - Meiosis-specific
CSKN1D† Hrr25 - Monopolin - Protein kinase CK1d
Meikin Spo13 - - - Meiosis-specific kinase regulaltors
(MOKIRs)
AuroraB Ipl1 - CPC - Protein kinase
Survivin Bir1 - CPC - CPC localisation
Borealin Nbl1 - CPC - CPC localisation
INCENP Sli15 - CPC - Kinase activation (IN-box)
MCAK n/a - - - Kinesin-13 MT catastrophe factor
PP1 Glc7 - - - Protein phosphatase 1
PP2A-B56 PP2A-Rts1 - - - Protein phosphatase 2A
Plk1 (Polo) Cdc5 - - - Protein kinase
Haspin Hsk1*/2* - - - Protein kinase (H3T3ph)
Cyclin B Cyclin B-Cdk1 Corona IV Binds Mad1
Bod1 - - - - Regulatory subunit for PP2A
SENP family Ulp2 Docks to Ctf3/CenpI; desumolase

Abbreviations: MT (Microtubule); n/a (no confirmed orthologue);

no evidence for kinetochore localisation in budding yeast* or humans†;

^

Budding yeast has a single shugoshin protein that has functions in common with mammalian Sgo1 and Sgo2.