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ULS1 | ATP-dependent helicase ULS1; Swi2/Snf2-related translocase, SUMO-Targeted Ubiquitin Ligase (STUbL); required for maintenance of NHEJ inhibition at telomeres; functions at telomeres to translocate and ubiquitinylate poly-sumoylated Rap1p for proteosomal degradation; plays role in antagonizing silencing during mating-type switching; only known STUbL with a translocase activity; contains RING finger domain; relocalizes from nucleus to cytoplasm upon DNA replication stress. (1619 aa) | ||||
MCM16 | Inner kinetochore subunit MCM16; Component of the Ctf19 complex and the COMA subcomplex; involved in kinetochore-microtubule mediated chromosome segregation; binds to centromere DNA; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-H and fission yeast fta3. (181 aa) | ||||
KAR9 | Karyogamy protein KAR9; Spindle positioning factor; orients astral microtubules, connecting them to actin cables at the cortex with Bim1p and Myo2, resulting in proper spindle positioning; targeted for StuBL-dependent degradation at kinetochores by Slx5p-Slx8p, ensuring chromosome transmission fidelity and correct spindle positioning; role in karyogamy; localizes to the shmoo tip, the growing bud-tip, the nucleus, the kinetochore, the spindle and microtubules; homolog of adenomatous polyposis coli. (644 aa) | ||||
PLC1 | 1-phosphatidylinositol 4,5-bisphosphate phosphodiesterase 1; Phospholipase C; hydrolyzes phosphatidylinositol 4,5-biphosphate (PIP2) to generate the signaling molecules inositol 1,4,5-triphosphate (IP3) and 1,2-diacylglycerol (DAG); involved in regulating many cellular processes; Plc1p and inositol polyphosphates are required for acetyl-CoA homeostasis which regulates global histone acetylation. (869 aa) | ||||
NSL1 | Kinetochore-associated protein NSL1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; required for accurate chromosome segregation; complex consists of Mtw1p Including Nnf1p-Nsl1p-Dsn1p (MIND). (216 aa) | ||||
IPL1 | Spindle assembly checkpoint kinase; Aurora kinase of chromosomal passenger complex; mediates release of mono-oriented kinetochores from microtubules in meiosis I, and kinetochore release from SPB clusters at meiotic exit; helps maintain condensed chromosomes during anaphase; required for SPB cohesion and prevention of multipolar spindle formation; promotes telomerase release at G2/M; Iocalizes to nuclear foci that diffuse upon DNA replication stress; required for inhibition of karyopherin Pse1p upon SAC arrest; Belongs to the protein kinase superfamily. Ser/Thr protein kinase family. A [...] (367 aa) | ||||
HRR25 | Conserved casein kinase; regulates diverse events including: vesicular traffic, DNA repair, the CVT pathway, monopolar attachment of sister kinetochores at meiosis I, and ribosomal subunit biogenesis; monopolin subunit; binds the RNAPII CTD; phosphorylates COPII coat subunits; interacts with Sit4p phosphatase; antagonizes calcineurin signaling, reducing nuclear accumulation of Crz1p; phosphorylates Dsn1p, the kinetochore receptor for monopolin; homolog of mammalian CK1delta. (494 aa) | ||||
CTF19 | Inner kinetochore subunit CTF19; Outer kinetochore protein, needed for accurate chromosome segregation; component of kinetochore sub-complex COMA (Ctf19p, Okp1p, Mcm21p, Ame1p) that functions as platform for kinetochore assembly; required for spindle assembly checkpoint; minimizes potentially deleterious centromere-proximal crossovers by preventing meiotic DNA break formation proximal to centromere; homolog of human centromere constitutive-associated network (CCAN) subunit CENP-P and fission yeast fta2. (369 aa) | ||||
SLK19 | Kinetochore-associated protein; required for chromosome segregation and kinetochore clustering; required for normal segregation of chromosomes in meiosis and mitosis; component of the FEAR regulatory network, which promotes Cdc14p release from the nucleolus during anaphase; potential Cdc28p substrate. (821 aa) | ||||
RIO1 | Serine/threonine-protein kinase RIO1; Serine kinase involved in cell cycle regulation and rDNA integrity; associated with late pre-40S particles via its conserved C-terminal domain and participates in late 40S biogenesis; association with pre-40S particles regulated by its catalytic ATPase site and likely occurs after the release of Rio2p from these particles; involved in cell cycle progression and processing of the 20S pre-rRNA into mature 18S rRNA; phosphorylates Rpa43p in anaphase to remove Pol I from rDNA. (484 aa) | ||||
SGO1 | Shugoshin; Component of the spindle checkpoint; involved in sensing lack of tension on mitotic chromosomes; protects centromeric Rec8p at meiosis I; required for accurate chromosomal segregation at meiosis II and for mitotic chromosome stability; recruits condensin to the pericentric region of chromosomes during meiosis; dissociates from pericentromeres when sister kinetochores are under tension; Belongs to the shugoshin family. (590 aa) | ||||
BUB3 | Cell cycle arrest protein BUB3; Kinetochore checkpoint WD40 repeat protein; localizes to kinetochores during prophase and metaphase, delays anaphase in the presence of unattached kinetochores; forms complexes with Mad1p-Bub1p and with Cdc20p, binds Mad2p and Mad3p; functions at kinetochore to activate APC/C-Cdc20p for normal mitotic progression. (341 aa) | ||||
RTS1 | Serine/threonine-protein phosphatase 2A 56 kDa regulatory subunit delta isoform; B-type regulatory subunit of protein phosphatase 2A (PP2A); Rts1p and Cdc55p are alternative regulatory subunits for PP2A catalytic subunits, Pph21p and Pph22p; PP2A-Rts1p protects cohesin when recruited by Sgo1p to the pericentromere; highly enriched at centromeres in the absence of Cdc55p; required for maintenance of septin ring organization during cytokinesis, for ring disassembly in G1 and for dephosphorylation of septin, Shs1p; homolog of the mammalian B' subunit of PP2A. (757 aa) | ||||
NUF2 | Kinetochore protein NUF2; Component of the kinetochore-associated Ndc80 complex; involved in chromosome segregation, spindle checkpoint activity, and kinetochore clustering; evolutionarily conserved; other members include Ndc80p, Nuf2p, Spc24p, and Spc25p. (451 aa) | ||||
DSL1 | Peripheral membrane protein needed for Golgi-to-ER retrograde traffic; mediates Sey1p-independent homotypic ER fusion; forms Dsl1 tethering complex with Sec39p and Tip20p that forms a stable complex with ER SNAREs Sec20p, Ufe1p and Use1p and is functionally conserved from yeast to mammalian cells; component of the ER target site that interacts with coatomer; interacts with different subunits of COPI vesicle coat; interacts with Cin5p; homolog of fly and human ZW10 gene. (754 aa) | ||||
CEP3 | Essential kinetochore protein; component of the CBF3 complex that binds the CDEIII region of the centromere; contains an N-terminal Zn2Cys6 type zinc finger domain, a C-terminal acidic domain, and a putative coiled coil dimerization domain. (608 aa) | ||||
SPC24 | Kinetochore protein SPC24; Component of the kinetochore-associated Ndc80 complex; involved in chromosome segregation, spindle checkpoint activity, and kinetochore clustering; evolutionarily conserved; other members include Ndc80p, Nuf2p, Spc24p, and Spc25p. (213 aa) | ||||
CTF3 | Inner kinetochore subunit CTF3; Outer kinetochore protein that forms a complex with Mcm16p and Mcm22p; may bind the kinetochore to spindle microtubules; required for the spindle assembly checkpoint; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-I and fission yeast mis6; Belongs to the CENP-I/CTF3 family. (733 aa) | ||||
NKP2 | Central kinetochore protein and subunit of the Ctf19 complex; mutants have elevated rates of chromosome loss; orthologous to fission yeast kinetochore protein cnl2. (153 aa) | ||||
STU2 | Protein STU2; Microtubule-associated protein (MAP) of the XMAP215/Dis1 family; regulates microtubule dynamics during spindle orientation and metaphase chromosome alignment; interacts with spindle pole body component Spc72p; Belongs to the TOG/XMAP215 family. (888 aa) | ||||
DAD2 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (133 aa) | ||||
SPC34 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; also localized to nuclear side of spindle pole body. (295 aa) | ||||
HSK3 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (69 aa) | ||||
ASK1 | Essential subunit of the Dam1 complex (aka DASH complex); couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; phosphorylated during the cell cycle by cyclin-dependent kinases; sumoylated in an Mms21p-dependent manner; protein abundance increases in response to DNA replication stress. (292 aa) | ||||
CSE4 | Histone H3-like centromere protein; associated with promoters, accessible chromatin, RNAPII-bound regions; phosphorylated Cse4p associates with centromeres; required for proper kinetochore function; levels regulated by E3 ubiquitin ligase Psh1p; phosphorylation may destabilize defective kinetochores to promote bi-orientation; ubiquitination of N-terminus regulates proteolysis for faithful chromosome segregation; yeast CSE4 can complement mutations in human homolog CENPA. (229 aa) | ||||
MCM22 | Inner kinetochore subunit MCM22; Outer kinetochore protein and component of the Ctf3 subcomplex; binds to centromeric DNA in a Ctf19p-dependent manner; involved in chromosome segregation and minichromosome maintenance; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-K and fission yeast sim4. (239 aa) | ||||
NNF1 | Kinetochore-associated protein NNF1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; required for accurate chromosome segregation; complex consists of Mtw1p Including Nnf1p-Nsl1p-Dsn1p (MIND). (201 aa) | ||||
BIR1 | Protein BIR1; Subunit of chromosomal passenger complex (CPC); CPC is comprised of Ipl1p-Sli15p-Bir1p-Nbl1p and regulates chromosome segregation; required for chromosome bi-orientation and for spindle assembly checkpoint activation upon reduced sister kinetochore tension; relative distribution to shortened microtubules increases upon DNA replication stress; sumoylated in an Mms21p-dependent manner; human survivin homolog. (954 aa) | ||||
CBF1 | Centromere-binding protein 1; Basic helix-loop-helix (bHLH) protein; forms homodimer to bind E-box consensus sequence CACGTG present at MET gene promoters and centromere DNA element I (CDEI); affects nucleosome positioning at this motif; associates with other transcription factors such as Met4p and Isw1p to mediate transcriptional activation or repression; associates with kinetochore proteins, required for chromosome segregation; protein abundance increases in response to DNA replication stress. (351 aa) | ||||
MAD2 | Component of the spindle-assembly checkpoint complex; delays onset of anaphase in cells with defects in mitotic spindle assembly; forms a complex with Mad1p; regulates APC/C activity during prometaphase and metaphase of meiosis I; gene dosage imbalance between MAD1 and MAD2 leads to chromosome instability. (196 aa) | ||||
DSN1 | Kinetochore-associated protein DSN1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; Dsn1p phosphorylation promotes interaction between outer and inner kinetochore proteins; kinetochore receptor for monopolin, via interaction with subunit Csm1p; essential for meiotic but not mitotic chromosome segregation; MIND complex consists of Mtw1p, Nnf1p, Nsl1p and Dsn1p; modified by sumoylation; phosphorylated by monopolin subunit Hrr25p. (576 aa) | ||||
NDC80 | Component of the kinetochore-associated Ndc80 complex; conserved coiled-coil protein involved in chromosome segregation, spindle checkpoint activity, and kinetochore assembly and clustering; evolutionarily conserved; complex members include Ndc80p, Nuf2p, Scp24p, and Spc25p; modified by sumoylation. (691 aa) | ||||
CRM1 | Exportin-1; Major karyopherin; involved in export of proteins, RNAs, and ribosomal subunits from the nucleus; exportin. (1084 aa) | ||||
BUB1 | Protein kinase involved in the cell cycle checkpoint into anaphase; in complex with Mad1p and Bub3p, prevents progression into anaphase in presence of spindle damage; Cdc28p-mediated phosphorylation at Bub1p-T566 is important for degradation in anaphase and adaptation of checkpoint to prolonged mitotic arrest; associates with centromere DNA via Skp1p; involved in Sgo1p relocalization in response to sister kinetochore tension; paralog MAD3 arose from whole genome duplication. (1021 aa) | ||||
OKP1 | Inner kinetochore subunit OKP1; Outer kinetochore protein required for accurate chromosome segregation; component of COMA (Ctf19p, Okp1p, Mcm21p, Ame1p) a kinetochore sub-complex which functions as a platform for kinetochore assembly; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-Q and fission yeast fta7. (406 aa) | ||||
DAM1 | Essential subunit of the Dam1 complex (aka DASH complex); cooperates with Duo1p to connect the DASH complex with the microtubules (MT); couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; Ipl1p target for regulating kinetochore-MT attachments. (343 aa) | ||||
SPT4 | Spt4p/5p (DSIF) transcription elongation factor complex subunit; the Spt4/5 complex binds to ssRNA in a sequence-specific manner, and along with RNAP I and II has multiple roles regulating transcriptional elongation, RNA processing, quality control, and transcription-coupled repair; localizes to kinetochores and heterochromatin, influencing chromosomal dynamics and silencing; required for transcription through long trinucleotide repeats in ORFs and non-protein coding regions. (102 aa) | ||||
SPC105 | Spindle pole body component SPC105; Subunit of a kinetochore-microtubule binding complex; complex bridges centromeric heterochromatin and kinetochore MAPs and motors; required for sister chromatid bi-orientation and kinetochore binding of SAC components; complex also includes Kre28p; modified by sumoylation. (917 aa) | ||||
MAD1 | Coiled-coil protein involved in spindle-assembly checkpoint; required for inhibition of karyopherin/importin Pse1p (aka Kap121p) upon spindle assembly checkpoint arrest; phosphorylated by Mps1p upon checkpoint activation which leads to inhibition of anaphase promoting complex activity; forms a complex with Mad2p; gene dosage imbalance between MAD1 and MAD2 leads to chromosome instability; Belongs to the MAD1 family. (749 aa) | ||||
DUO1 | Essential subunit of the Dam1 complex (aka DASH complex); cooperates with Dam1p to connect the DASH complex with microtubules (MT); couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (247 aa) | ||||
YBP2 | YAP1-binding protein 2; Central kinetochore associated protein; mediates mitotic progression; interacts with several central kinetochore proteins and centromeric histone Cse4p; role in resistance to oxidative stress; similar to Slk19p; YBP2 has a paralog, YBP1, that arose from the whole genome duplication. (641 aa) | ||||
CNN1 | Inner kinetochore subunit CNN1; Kinetochore protein; associated with the essential kinetochore proteins Nnf1p and Spc24p; phosphorylated by Clb5-Cdk1, Mps1p, Ipl1p and to a lesser extent by Clb2-Cdk1; localizes to the lower region of the Ndc80 complex during anaphase and regulates KMN activity by inhibiting the Mtw1 and Spc105 complexes from binding to the Ndc80 complex; similar to metazoan CENP-T; Belongs to the CENP-T/CNN1 family. (361 aa) | ||||
GLC7 | Serine/threonine-protein phosphatase PP1-2; Type 1 S/T protein phosphatase (PP1) catalytic subunit; involved in glycogen metabolism, sporulation and mitotic progression; interacts with multiple regulatory subunits; regulates actomyosin ring formation; subunit of CPF; recruited to mating projections by Afr1p interaction; regulates nucleocytoplasmic shuttling of Hxk2p; import into the nucleus is inhibited during spindle assembly checkpoint arrest; involved in dephosphorylating Rps6a/b and Bnr1p. (312 aa) | ||||
SLX8 | Subunit of Slx5-Slx8 SUMO-targeted ubiquitin ligase (STUbL) complex; role in proteolysis of spindle positioning protein Kar9, DNA repair proteins Rad52p and Rad57p; stimulated by SUMO-modified substrates; contains a C-terminal RING domain; forms nuclear foci upon DNA replication stress; required for maintenance of genome integrity like human ortholog RNF. (274 aa) | ||||
MAM1 | Monopolin complex subunit MAM1; Monopolin; meiosis-specific kinetochore-associated protein involved in monopolar attachment of sister kinetochores to the meiotic spindle; subunit of monopolin, a complex that prevents biorientation of sister kinetochores to ensure homolog biorientation during meiosis I; regulates the conformation, enzyme kinetics and substrate specificity of the Dsn1p kinase, Hrr1p; expressed only during the first meiotic division. (302 aa) | ||||
SPC25 | Kinetochore protein SPC25; Component of the kinetochore-associated Ndc80 complex; involved in chromosome segregation, spindle checkpoint activity, and kinetochore clustering; evolutionarily conserved; other members include Ndc80p, Nuf2p, Spc24p, and Spc25p. (221 aa) | ||||
CIN8 | Kinesin-like protein CIN8; Kinesin motor protein; involved in mitotic spindle assembly and chromosome segregation. (1000 aa) | ||||
KRE28 | Spindle pole body component KRE28; Subunit of a kinetochore-microtubule binding complex; complex bridges centromeric heterochromatin and kinetochore MAPs and motors; required for sister chromatid bi-orientation and kinetochore binding of SAC components; complex also includes Spc105p; modified by sumoylation. (385 aa) | ||||
LRS4 | Monopolin complex subunit LRS4; Nucleolar protein that forms a complex with Csm1p; and then Mam1p at kinetochores during meiosis I to mediate accurate homolog segregation; required for condensin recruitment to the replication fork barrier site and rDNA repeat segregation. (347 aa) | ||||
NKP1 | Central kinetochore protein and subunit of the Ctf19 complex; mutants have elevated rates of chromosome loss; orthologous to fission yeast kinetochore protein fta4; Belongs to the NKP1 family. (238 aa) | ||||
WIP1 | Inner kinetochore subunit WIP1; Kinetochore localized protein of unknown function; interacts with Cnn1p (CENP-T); orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-W and fission yeast new1. (89 aa) | ||||
SKP1 | Evolutionarily conserved kinetochore protein; part of multiple protein complexes, including the SCF ubiquitin ligase complex, the CBF3 complex that binds centromeric DNA, and the RAVE complex that regulates assembly of the V-ATPase; protein abundance increases in response to DNA replication stress. (194 aa) | ||||
DAD4 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (72 aa) | ||||
MCM21 | Inner kinetochore subunit MCM21; Component of the kinetochore sub-complex COMA; COMA (Ctf19p, Okp1p, Mcm21p, Ame1p) bridges kinetochore subunits in contact with centromeric DNA with subunits bound to microtubules during kinetochore assembly; involved in minichromosome maintenance; modified by sumoylation; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-O and fission yeast mal2; Belongs to the CENP-O/MCM21 family. (368 aa) | ||||
CHL4 | Inner kinetochore subunit CHL4; Outer kinetochore protein required for chromosome stability; involved in new kinetochore assembly and sister chromatid cohesion; forms a stable complex with Iml3p; peripheral component of the Ctf19 kinetochore complex that interacts with Ctf19p, Ctf3p, and Mif2p; required for the spindle assembly checkpoint; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-N and fission yeast mis15; Belongs to the CENP-N/CHL4 family. (458 aa) | ||||
SPC19 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; also localized to nuclear side of spindle pole body. (165 aa) | ||||
FIN1 | Filament protein FIN1; Spindle pole body-related intermediate filament protein; forms cell cycle-specific filaments between spindle pole bodies in dividing cells; localizes to poles and microtubules of spindle during anaphase and contributes to spindle stability; involved in Glc7p localization and regulation; relative distribution to the nucleus increases upon DNA replication stress. (291 aa) | ||||
DAD1 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (94 aa) | ||||
MPS1 | Serine/threonine-protein kinase MPS1; Dual-specificity kinase; autophosphorylation required for function; required for spindle pole body (SPB) duplication and spindle checkpoint function; contributes to bi-orientation by promoting formation of force-generating kinetochore-microtubule attachments in meiosis I; substrates include SPB proteins Spc42p, Spc110p, and Spc98p, mitotic exit network protein Mob1p, kinetochore protein Cnn1p, and checkpoint protein Mad1p; substrate of APCC(Cdh1); similar to human Mps1p. (764 aa) | ||||
CSM1 | Monopolin complex subunit CSM1; Nucleolar protein that mediates homolog segregation during meiosis I; forms a complex with Lrs4p and then Mam1p at kinetochores; required for condensin recruitment to the replication fork barrier site and rDNA repeat segregation. (190 aa) | ||||
PAT1 | DNA topoisomerase 2-associated protein PAT1; Deadenylation-dependent mRNA-decapping factor; also required for faithful chromosome transmission, maintenance of rDNA locus stability, and protection of mRNA 3'-UTRs from trimming; associated with topoisomerase II; binds to mRNAs under glucose starvation, most often in the 3' UTR; functionally linked to Pab1p; forms cytoplasmic foci upon DNA replication stress; phosphorylation by PKA inhibits P body foci formation; Belongs to the PAT1 family. (796 aa) | ||||
BIK1 | Nuclear fusion protein BIK1; Microtubule-associated protein; component of the interface between microtubules and kinetochore, involved in sister chromatid separation; essential in polyploid cells but not in haploid or diploid cells; ortholog of mammalian CLIP-170. (440 aa) | ||||
DAD3 | Essential subunit of the Dam1 complex (aka DASH complex); complex couples kinetochores to the force produced by MT depolymerization thereby aiding in chromosome segregation; is transferred to the kinetochore prior to mitosis. (94 aa) | ||||
AME1 | Inner kinetochore subunit AME1; Essential kinetochore protein associated with microtubules and SPBs; component of the kinetochore sub-complex COMA (Ctf19p, Okp1p, Mcm21p, Ame1p); involved in spindle checkpoint maintenance; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-U and fission yeast Mis17; relative distribution to the nucleus increases upon DNA replication stress; Belongs to the CENP-U/AME1 family. (324 aa) | ||||
IML3 | Inner kinetochore subunit IML3; Outer kinetochore protein and component of the Ctf19 complex; involved in the establishment of pericentromeric cohesion during mitosis; prevents non-disjunction of sister chromatids during meiosis II; forms a stable complex with Chl4p; required for localization of Sgo1p to pericentric sites during meiosis I; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-L and fission yeast fta1; Belongs to the CENP-L/IML3 family. (245 aa) | ||||
STU1 | Protein STU1; Component of the mitotic spindle; binds to interpolar microtubules via its association with beta-tubulin (Tub2p); required for interpolar microtubules to provide an outward force on the spindle poles; Belongs to the CLASP family. (1513 aa) | ||||
MTW1 | Kinetochore-associated protein MTW1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; critical to kinetochore assembly; complex consists of Mtw1p Including Nnf1p-Nsl1p-Dsn1p (MIND). (289 aa) |