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| | SPO13 | Meiosis-specific protein SPO13; Meiotic regulator; involved in maintaining sister chromatid cohesion during meiosis I as well as promoting proper attachment of kinetochores to the spindle during meiosis I and meiosis II; anaphase-promoting complex (APC) substrate that is degraded during anaphase I; expressed only in meiotic cells. (291 aa) |
| | RRM3 | DNA helicase involved in rDNA replication and Ty1 transposition; binds to and suppresses DNA damage at G4 motifs in vivo; relieves replication fork pauses at telomeric regions; structurally and functionally related to Pif1p; Belongs to the helicase family. (723 aa) |
| | DNA2 | Tripartite DNA replication factor; single-stranded DNA-dependent ATPase, ATP-dependent nuclease, helicase; tracking protein for flap cleavage during Okazaki fragment maturation; involved in DNA repair/processing of meiotic DNA double strand breaks; component of telomeric chromatin with cell-cycle dependent localization; required for telomerase-dependent telomere synthesis; forms nuclear foci upon DNA replication stress; human homolog DNA2 complements yeast dna2 mutant. (1522 aa) |
| | CDC23 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); APC/C is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition. (626 aa) |
| | EST3 | Telomere replication protein EST3; Component of the telomerase holoenzyme; involved in telomere replication; synthesis of the full-length protein results from a programmed +1 ribosomal frameshift. (181 aa) |
| | DOT5 | Peroxiredoxin DOT5; Nuclear thiol peroxidase; functions as an alkyl-hydroperoxide reductase during post-diauxic growth; Belongs to the peroxiredoxin family. BCP/PrxQ subfamily. (215 aa) |
| | IRR1 | Subunit of the cohesin complex; which is required for sister chromatid cohesion during mitosis and meiosis and interacts with centromeres and chromosome arms; relocalizes to the cytosol in response to hypoxia; essential for viability; Belongs to the SCC3 family. (1150 aa) |
| | MOB1 | DBF2 kinase activator protein MOB1; Component of the mitotic exit network; associates with and is required for the activation and Cdc15p-dependent phosphorylation of the Dbf2p kinase; required for cytokinesis and cell separation; component of the CCR4 transcriptional complex; relocalizes from cytoplasm to the nuclear periphery upon DNA replication stress; Belongs to the MOB1/phocein family. (314 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) |
| | MCM10 | Minichromosome maintenance protein 10; Essential chromatin-associated protein; involved in initiation of DNA replication; required for association of MCM2-7 complex with replication origins; required to stabilize catalytic subunit of DNA polymerase-alpha; self-associates through its N-terminal domain. (571 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) |
| | MPS3 | Spindle pole body assembly component MPS3; Nuclear envelope protein; required for SPB insertion, SPB duplication, Kar5p localization near the SPB and nuclear fusion; interacts with Mps2p to tether half-bridge to core SPB; N-terminal acetylation by Eco1p regulates its role in nuclear organization; localizes to the SPB half bridge and telomeres during meiosis; required with Ndj1p and Csm4p for meiotic bouquet formation and telomere-led rapid prophase movement; member of the SUN protein family (Sad1-UNC-84 homology). (682 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) |
| | DLS1 | Protein DLS1; Subunit of ISW2/yCHRAC chromatin accessibility complex; ISW2/yCHRAC also includes Itc1p, Isw2p, and Dpb4p; involved in inheritance of telomeric silencing; DLS1 has a paralog, DPB3, that arose from the whole genome duplication. (167 aa) |
| | SCP160 | Protein SCP160; Essential RNA-binding G protein effector of mating response pathway; ligand-activated RNA-binding protein that delivers RNAs involved in polarization and perpetualizing mating signal to shmoo tip during pheromone signaling; Scp160p-mediated RNA trafficking essential for chemotropism and successful mating; mainly associated with nuclear envelope and ER, interacts in mRNA-dependent manner with translating ribosomes via multiple KH domains, similar to vertebrate vigilins. (1222 aa) |
| | ASF1 | Histone chaperone ASF1; Nucleosome assembly factor; involved in chromatin assembly, disassembly; required for recovery after DSB repair; role in H3K56 acetylation required for expression homeostasis, buffering mRNA synthesis rate against gene dosage changes in S phase; anti-silencing protein, derepresses silent loci when overexpressed; role in regulating Ty1 transposition; relocalizes to cytosol under hypoxia; growth defect of asf1 null is functionally complemented by either human ASF1A or ASF1B. (279 aa) |
| | SPT10 | Protein SPT10; Histone H3 acetylase with a role in transcriptional regulation; sequence-specific activator of histone genes, binds specifically and cooperatively to pairs of UAS elements in core histone promoters, functions at or near TATA box; involved in S phase-specific acetylation of H3K56 at histone promoters, which is required for recruitment of SWI/SNF nucleosome remodeling complex and subsequent transcription. (640 aa) |
| | RFA3 | Subunit of heterotrimeric Replication Protein A (RPA); RPA is a highly conserved single-stranded DNA binding protein complex involved in DNA replication, repair, recombination; RPA protects against inappropriate telomere recombination, and upon telomere uncapping, prevents cell proliferation by a checkpoint-independent pathway; with Sgs1p-Top2p-Rmi1p, stimulates DNA catenation/decatenation activity of Top3p; protein abundance increases in response to DNA replication stress. (122 aa) |
| | GON7 | Component of the EKC/KEOPS protein complex; EKC/KEOPS complex is required for t6A tRNA modification and telomeric TG1-3 recombination; may have role in transcription; implicated in osmotic stress response; other complex members are Kae1p, Cgi121p, Pcc1p, and Bud32p; Belongs to the GON7 family. (123 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) |
| | 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) |
| | 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) |
| | 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) |
| | SWD2 | COMPASS component SWD2; Subunit of the COMPASS (Set1C) histone H3K4 methyltransferase complex; required for Set1C stability and optimal activity; COMPASS methylates histone H3 on lys 4 and is involved in telomeric silencing; subunit of CPF (cleavage and polyadenylation factor), a complex involved in RNAP II transcription termination; Belongs to the WD repeat SWD2 family. (329 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) |
| | 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) |
| | DEF1 | RNAPII degradation factor; forms a complex with Rad26p in chromatin, enables ubiquitination and proteolysis of RNAPII present in an elongation complex; mutant is deficient in Zip1p loading onto chromosomes during meiosis. (738 aa) |
| | NUP120 | Nucleoporin NUP120; Subunit of the Nup84p subcomplex of the nuclear pore complex (NPC); contributes to nucleocytoplasmic transport and NPC biogenesis and is involved in establishment of a normal nucleocytoplasmic concentration gradient of the GTPase Gsp1p; also plays roles in several processes that may require localization of genes or chromosomes at the nuclear periphery, including double-strand break repair, transcription and chromatin silencing; homologous to human NUP160. (1037 aa) |
| | MIF2 | Inner kinetochore subunit MIF2; Protein required for structural integrity of elongating spindles; localizes to the kinetochore; interacts with histones H2A, H2B, and H4; phosphorylated by Ipl1p; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-C and fission yeast cnp3; Belongs to the CENP-C/MIF2 family. (549 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) |
| | 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) |
| | NUP133 | Nucleoporin NUP133; Subunit of Nup84p subcomplex of nuclear pore complex (NPC); contributes to nucleocytoplasmic transport, NPC biogenesis; is involved in establishment of a normal nucleocytoplasmic concentration gradient of GTPase Gsp1p; also plays roles in several processes that may require localization of genes or chromosomes at nuclear periphery, including double-strand break repair, transcription and chromatin silencing; relocalizes to cytosol in response to hypoxia; homolog of human NUP133; Belongs to the nucleoporin Nup133 family. (1157 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) |
| | PCC1 | Component of the EKC/KEOPS protein complex; EKC/KEOPS complex is required for t6A tRNA modification and telomeric TG1-3 recombination; may have role in transcription; other complex members are Kae1p, Gon7p, Bud32p, and Cgi121p. (88 aa) |
| | SIR1 | Regulatory protein SIR1; Protein involved in silencing at mating-type loci HML and HMR; recruitment to silent chromatin requires interactions with Orc1p and with Sir4p, through a common Sir1p domain; binds to centromeric chromatin. (654 aa) |
| | HIF1 | HAT1-interacting factor 1; Non-essential component of the HAT-B histone acetyltransferase complex; localized to the nucleus; has a role in telomeric silencing; other members are Hat1p and Hat2p; Belongs to the NASP family. (385 aa) |
| | TEN1 | Telomere length regulation protein TEN1; Protein that regulates telomeric length; protects telomeric ends in a complex with Cdc13p and Stn1p; similar to human Ten1 which is critical for the telomeric function of the CST (Cdc13p-Stn1p-Ten1p) complex. (160 aa) |
| | BRE2 | COMPASS component BRE2; Subunit of COMPASS (Set1C) complex; COMPASS methylates Lys4 of histone H3 and functions in silencing at telomeres; has a C-terminal Sdc1 Dpy-30 Interaction (SDI) domain that mediates binding to Sdc1p; similar to trithorax-group protein ASH2L. (505 aa) |
| | RAD5 | DNA repair protein RAD5; DNA helicase/Ubiquitin ligase; involved in error-free DNA damage tolerance (DDT), replication fork regression during postreplication repair by template switching, error-prone translesion synthesis; promotes synthesis of free and PCNA-bound polyubiquitin chains by Ubc13p-Mms2p; forms nuclear foci upon DNA replication stress; associates with native telomeres, cooperates with homologous recombination in senescent cells; human homolog HLTF can complement yeast null mutant. (1169 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) |
| | SIR2 | Conserved NAD+ dependent histone deacetylase of the Sirtuin family; deacetylation targets are primarily nuclear proteins; required for telomere hypercluster formation in quiescent yeast cells; involved in regulation of lifespan; plays roles in silencing at HML, HMR, telomeres, and rDNA; negatively regulates initiation of DNA replication; functions as regulator of autophagy like mammalian homolog SIRT1, and also of mitophagy. (562 aa) |
| | BRE1 | E3 ubiquitin ligase; forms heterodimer with Rad6p to regulate K63 polyubiquitination in response to oxidative stress and to monoubiquinate histone H2B-K123, which is required for the subsequent methylation of histone H3-K4 and H3-K79; required for DSBR, transcription, silencing, and checkpoint control; interacts with RNA-binding protein Npl3p, linking histone ubiquitination to mRNA processing; Bre1p-dependent histone ubiquitination promotes pre-mRNA splicing. (700 aa) |
| | SUB2 | ATP-dependent RNA helicase SUB2; Component of the TREX complex required for nuclear mRNA export; member of the DEAD-box RNA helicase superfamily and is involved in early and late steps of spliceosome assembly; homolog of the human splicing factor hUAP56; relocalizes from nucleus to cytoplasm upon DNA replication stress. (446 aa) |
| | NUP84 | Subunit of the Nup84p subcomplex of the nuclear pore complex (NPC); contributes to nucleocytoplasmic transport and NPC biogenesis; also plays roles in several processes that may require localization of genes or chromosomes at the nuclear periphery, including double-strand break repair, transcription and chromatin silencing; homologous to human NUP107. (726 aa) |
| | CDC53 | Cell division control protein 53; Cullin; structural protein of SCF complexes (which also contain Skp1p, Cdc34p, Hrt1p and an F-box protein) involved in ubiquitination; SCF promotes the G1-S transition by targeting G1 cyclins and the Cln-CDK inhibitor Sic1p for degradation; human homolog CUL1 can complement yeast cdc53 null mutant. (815 aa) |
| | SCM3 | Protein SCM3; Nonhistone component of centromeric chromatin; binds to histone H3 variant, Cse4p, and recruits it to centromeres; involved in the assembly and maintenance of Cse4-H4 at centromeres; required for kinetochore assembly and G2/M progression; may protect Cse4p from ubiquitination; homolog of mammalian HJURP. (223 aa) |
| | PPH22 | Catalytic subunit of protein phosphatase 2A (PP2A); functionally redundant with Pph21p; methylated at C terminus; forms alternate complexes with several regulatory subunits; involved in signal transduction and regulation of mitosis; protein abundance increases in response to DNA replication stress; dephosphorylates Tel1p/Mec1p-phosphorylated Cdc13p to promote telomerase release from telomeres at G2/M; PPH22 has a paralog, PPH21, that arose from the whole genome duplication; Belongs to the PPP phosphatase family. PP-2A subfamily. (377 aa) |
| | ASF2 | Anti-silencing protein; causes derepression of silent loci when overexpressed. (525 aa) |
| | CDC13 | Cell division control protein 13; Single stranded DNA-binding protein found at TG1-3 telomere G-tails; key roles in regulation of telomerase, telomere end protection, conventional telomere replication; regulates telomere replication through recruitment of specific sub-complexes, essential function is telomere capping; forms homodimer via N-terminus; disruption of dimerization leads to short telomeres; autophagy and proteasome are involved in Cdc13p degradation; differentially phosphorylated through cell cycle. (924 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) |
| | DBF4 | Regulatory subunit of Cdc7p-Dbf4p kinase complex; required for Cdc7p kinase activity and initiation of DNA replication; phosphorylates the Mcm2-7 family of proteins; cell cycle regulated; relative distribution to the nucleus increases upon DNA replication stress; co-expression of human CDC7 and DBF4 complements single cdc7 or dbf4 null mutations or the cdc7 dbf4 double null mutation. (704 aa) |
| | CDC34 | Ubiquitin-conjugating enzyme (E2); catalytic subunit of SCF ubiquitin-protein ligase complex (together with Skp1p, Rbx1p, Cdc53p, and an F-box protein) that regulates cell cycle progression by targeting key substrates for degradation; protein abundance increases in response to DNA replication stress; human CDC34 functionally complements the thermosensitivity of the cdc34-2 mutant. (295 aa) |
| | STN1 | Protein STN1; Telomere end-binding and capping protein; plays a key role with Pol12p in linking telomerase action with completion of lagging strand synthesis, and in a regulatory step required for telomere capping; similar to human Stn1. (494 aa) |
| | RRP8 | 25S rRNA (adenine(645)-N(1))-methyltransferase; Nucleolar S-adenosylmethionine-dependent rRNA methyltransferase; methylates adenine (m1A) of the large subunit (LSU) rRNA at position 645; involved in pre-rRNA cleavage at site A2; mutation is synthetically lethal with a gar1 mutation; deletion disrupts telomere maintenance by influencing the expression of neighboring gene STN1; Belongs to the methyltransferase superfamily. RRP8 family. (392 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) |
| | HPR1 | Subunit of THO/TREX complexes; this complex couple transcription elongation with mitotic recombination and with mRNA metabolism and export, subunit of an RNA Pol II complex; regulates lifespan; involved in telomere maintenance; similar to Top1p. (752 aa) |
| | SCC2 | Sister chromatid cohesion protein 2; Subunit of cohesin loading factor (Scc2p-Scc4p); a complex required for loading of cohesin complexes onto chromosomes; involved in establishing sister chromatid cohesion during DSB repair via histone H2AX; promotes gene expression program that supports translational fidelity; evolutionarily-conserved adherin; relocalizes to cytosol in response to hypoxia; human disorder Cornelia de Lange syndrome is caused by mutations in NIPBL, the human ortholog of SCC2. (1493 aa) |
| | SAS4 | Something about silencing protein 4; Subunit of the SAS complex (Sas2p, Sas4p, Sas5p); acetylates free histones and nucleosomes and regulates transcriptional silencing; required for the HAT activity of Sas2p. (481 aa) |
| | HST4 | NAD-dependent histone deacetylase HST4; NAD(+)-dependent protein deacetylase; deacetylation targets are primarily mitochondrial proteins; involved along with Hst3p in silencing at telomeres, cell cycle progression, radiation resistance, genomic stability and short-chain fatty acid metabolism; accumulates in mitochondria in response to biotin starvation and may link biotin metabolism with energy homeostasis; member of the Sir2 family and may be the functional equivalent of human SIRT3. (370 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) |
| | EBS1 | Protein involved in translation inhibition and nonsense-mediated decay; interacts with cap binding protein Cdc33p and with Nam7p; localizes to P-bodies upon glucose starvation; mRNA abundance regulated by mRNA decay factors; EBS1 has a paralog, EST1, that arose from the whole genome duplication. (884 aa) |
| | SIR4 | Regulatory protein SIR4; SIR protein involved in assembly of silent chromatin domains; silent information regulator (SIR) along with SIR2 and SIR3; involved in assembly of silent chromatin domains at telomeres and the silent mating-type loci; some alleles of SIR4 prolong lifespan; required for telomere hypercluster formation in quiescent yeast cells. (1358 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) |
| | ZIP1 | Transverse filament protein of the synaptonemal complex; required for normal levels of meiotic recombination and pairing between homologous chromosome during meiosis; required for meiotic recombination between non-allelc sites; potential Cdc28p substrate. (875 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | DOT1 | Histone-lysine N-methyltransferase, H3 lysine-79 specific; Nucleosomal histone H3-Lys79 methylase; methylation is required for telomeric silencing, meiotic checkpoint control, and DNA damage response. (582 aa) |
| | ADA2 | Transcriptional adapter 2; Transcription coactivator; component of the ADA and SAGA transcriptional adaptor/HAT (histone acetyltransferase) complexes. (434 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) |
| | FUN30 | ATP-dependent helicase FUN30; Snf2p family member with ATP-dependent chromatin remodeling activity; has a role in silencing at the mating type locus, telomeres and centromeres; enriched at centromeres and is required for correct chromatin structure around centromeres, as well as at the boundary element of the silent HMR; recruited to DNA double-strand breaks (DSBs) where it promotes 5' strand resection of DSBs; potential Cdc28p substrate. (1131 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) |
| | SWD1 | COMPASS component SWD1; Subunit of the COMPASS (Set1C) complex; COMPASS methylates histone H3 on lysine 4 and is required in transcriptional silencing near telomeres; WD40 beta propeller superfamily member with similarity to mammalian Rbbp7. (426 aa) |
| | RFA1 | Subunit of heterotrimeric Replication Protein A (RPA); RPA is a highly conserved single-stranded DNA binding protein involved in DNA replication, repair, and recombination; RPA protects against inappropriate telomere recombination, and upon telomere uncapping, prevents cell proliferation by a checkpoint-independent pathway; role in DNA catenation/decatenation pathway of chromosome disentangling; relocalizes to the cytosol in response to hypoxia. (621 aa) |
| | HIR1 | Protein HIR1; Subunit of the HIR complex; HIR is a nucleosome assembly complex involved in regulation of histone gene transcription; contributes to nucleosome formation, heterochromatic gene silencing, and formation of functional kinetochores. (840 aa) |
| | HEK2 | Heterogeneous nuclear rnp K-like protein 2; RNA binding protein involved in asymmetric localization of ASH1 mRNA; represses translation of ASH1 mRNA, an effect reversed by Yck1p-dependent phosphoryation; regulates telomere position effect and length; similarity to hnRNP-K. (381 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) |
| | SAS3 | Histone acetyltransferase catalytic subunit of NuA3 complex; acetylates histone H3, involved in transcriptional silencing; homolog of the mammalian MOZ proto-oncogene; mutant has aneuploidy tolerance; sas3gcn5 double mutation is lethal; Belongs to the MYST (SAS/MOZ) family. (831 aa) |
| | TEL1 | Serine/threonine-protein kinase TEL1; Protein kinase primarily involved in telomere length regulation; contributes to cell cycle checkpoint control in response to DNA damage; acts with Red1p and Mec1p to promote interhomolog recombination by phosphorylation of Hop1; functionally redundant with Mec1p; regulates P-body formation induced by replication stress; homolog of human ataxia-telangiectasia mutated (ATM) gene. (2787 aa) |
| | HHT1 | Histone H3; core histone protein required for chromatin assembly, part of heterochromatin-mediated telomeric and HM silencing; one of two identical histone H3 proteins (see HHT2); regulated by acetylation, methylation, and phosphorylation; H3K14 acetylation plays an important role in the unfolding of strongly positioned nucleosomes during repair of UV damage. (136 aa) |
| | ORC2 | Subunit of the origin recognition complex (ORC); ORC directs DNA replication by binding to replication origins and is also involved in transcriptional silencing; interacts with Spp1p and with trimethylated histone H3; phosphorylated by Cdc28p. (620 aa) |
| | POL30 | Proliferating cell nuclear antigen (PCNA); functions as the sliding replication clamp for DNA polymerase delta; may function as a docking site for other proteins required for mitotic and meiotic chromosomal DNA replication and for DNA repair; PCNA ubiquitination at K164 plays a crucial role during Okazaki fragment processing. (258 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) |
| | SLI15 | Inner centromere protein-related protein SLI15; Subunit of the conserved chromosomal passenger complex (CPC); complex regulates kinetochore-microtubule attachments, activation of the spindle tension checkpoint, and mitotic spindle disassembly; other complex members are Ipl1p, Bir1p, and Nbl1p. (698 aa) |
| | SWD3 | COMPASS component SWD3; Essential subunit of the COMPASS (Set1C) complex; COMPASS methylates histone H3 on lysine 4 and is required in transcriptional silencing near telomeres; WD40 beta propeller superfamily member and ortholog of mammalian WDR5. (315 aa) |
| | MCM7 | DNA replication licensing factor MCM7; Component of the Mcm2-7 hexameric helicase complex; MCM2-7 primes origins of DNA replication in G1 and becomes an active ATP-dependent helicase that promotes DNA melting and elongation in S-phase; forms an Mcm4p-6p-7p subcomplex. (845 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) |
| | 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) |
| | SHG1 | COMPASS component SHG1; Subunit of the COMPASS (Set1C) complex; COMPASS methylates histone H3 on lysine 4 and is required in transcriptional silencing near telomeres. (142 aa) |
| | RIF1 | Telomere length regulator protein RIF1; Protein that binds to the Rap1p C-terminus; acts synergistically with Rif2p to help control telomere length and establish telomeric silencing; involved in control of DNA replication; contributes to resection of DNA double strand breaks (DSBs); deletion results in telomere elongation; Belongs to the RIF1 family. (1916 aa) |
| | GBP2 | Single-strand telomeric DNA-binding protein GBP2; Poly(A+) RNA-binding protein; key surveillance factor for the selective export of spliced mRNAs from the nucleus to the cytoplasm; preference for intron-containing genes; similar to Npl3p; also binds single-stranded telomeric repeat sequence in vitro; relocalizes to the cytosol in response to hypoxia; GBP2 has a paralog, HRB1, that arose from the whole genome duplication. (427 aa) |
| | DCC1 | Sister chromatid cohesion protein DCC1; Subunit of a complex with Ctf8p and Ctf18p; shares some components with Replication Factor C; required for sister chromatid cohesion and telomere length maintenance; Belongs to the DCC1 family. (380 aa) |
| | CDC7 | Cell division control protein 7; DDK (Dbf4-dependent kinase) catalytic subunit; required for origin firing and replication fork progression in mitotic S phase through phosphorylation of Mcm2-7p complexes and Cdc45p; kinase activity correlates with cyclical DBF4 expression; required for pre-meiotic DNA replication, meiotic DSB formation, recruitment of monopolin complex to kinetochores during meiosis I, regulation of meiosis-specific Ndt80p; mutation complemented by human CDC7 and DBF4 co-expression. (507 aa) |
| | SLX5 | Subunit of the Slx5-Slx8 SUMO-targeted Ub ligase (STUbL) complex; role in Ub-mediated degradation of histone variant Cse4p preventing mislocalization to euchromatin; role in proteolysis of spindle positioning protein Kar9p, and DNA repair proteins Rad52p and Rad57p; forms SUMO-dependent nuclear foci, including DNA repair centers; contains a RING domain and two SIM motifs; associates with the centromere; required for maintenance of genome integrity like human ortholog RNF4. (619 aa) |
| | MCD1 | Sister chromatid cohesion protein 1; Essential alpha-kleisin subunit of the cohesin complex; required for sister chromatid cohesion in mitosis and meiosis; apoptosis induces cleavage and translocation of a C-terminal fragment to mitochondria; expression peaks in S phase. (566 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) |
| | RLF2 | Largest subunit (p90) of the Chromatin Assembly Complex (CAF-1); chromatin assembly by CAF-1 is important for multiple processes including silencing at telomeres, mating type loci, and rDNA; maintenance of kinetochore structure; deactivation of the DNA damage checkpoint after DNA repair; chromatin dynamics during transcription; and repression of divergent noncoding transcription. (606 aa) |
| | REC8 | Meiotic recombination protein REC8; Meiosis-specific component of the sister chromatid cohesion complex; alpha-kleisin family member that maintains cohesion between sister chromatids during meiosis I; maintains cohesion between centromeres of sister chromatids until meiosis II; independent of its role in sister chromatid cohesion, Rec8p promotes allelic collisions and prevents nonspecific chromosome interactions; homolog of S. pombe Rec8p. (680 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) |
| | CSM4 | Protein required for accurate chromosome segregation during meiosis; involved in meiotic telomere clustering (bouquet formation) and telomere-led rapid prophase movements; functions with meiosis-specific telomere-binding protein Ndj1p; CSM4 has a paralog, MPS2, that arose from the whole genome duplication. (156 aa) |
| | SPP1 | COMPASS component SPP1; Subunit of COMPASS (Set1C); a complex which methylates histone H3 on lysine 4 and is required in telomeric transcriptional silencing; promotes meiotic DSB formation by interacting with H3K4me3 and Rec107p, a protein required for Spo11p-catalyzed DSB formation located on chromosome axes; interacts with Orc2p; PHD finger domain protein similar to human CGBP, an unmethylated CpG binding protein; relocalizes to cytosol in response to hypoxia. (353 aa) |
| | TBF1 | Protein TBF1; Telobox-containing general regulatory factor; binds TTAGGG repeats within subtelomeric anti-silencing regions (STARs), blocking silent chromatin propagation; binds majority of snoRNA gene promoters, required for full snoRNA expression; caps DSB flanked by long T2AG3 repeats and blocks checkpoint activation. (562 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) |
| | HAT1 | Catalytic subunit of the Hat1p-Hat2p histone acetyltransferase complex; uses the cofactor acetyl coenzyme A to acetylate free nuclear and cytoplasmic histone H4; involved in telomeric silencing and DNA double-strand break repair. (374 aa) |
| | ISW2 | ISWI chromatin-remodeling complex ATPase ISW2; ATP-dependent DNA translocase involved in chromatin remodeling; ATPase component that, with Itc1p, forms a complex required for repression of a-specific genes, INO1, and early meiotic genes during mitotic growth; the Isw2 complex exhibits basal levels of chromatin binding throughout the genome as well as target-specific chromatin interactions; targeted by Ume6p- and Sua7p-dependent DNA looping to many loci genome-wide. (1120 aa) |
| | PAC1 | Nuclear distribution protein PAC1; Involved in nuclear migration, part of the dynein/dynactin pathway; targets dynein to microtubule tips, which is necessary for sliding of microtubules along bud cortex; serves at interface between dynein's ATPase site and its microtubule binding stalk, causing individual dynein motors to remain attached to microtubules for long periods; synthetic lethal with bni1; homolog of human LIS1, mutations in which cause the severe brain disorder lissencephaly. (494 aa) |
| | SAS5 | Something about silencing protein 5; Subunit of the SAS complex (Sas2p, Sas4p, Sas5p); acetylates free histones and nucleosomes and regulates transcriptional silencing; stimulates Sas2p HAT activity. (248 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) |
| | NPT1 | Nicotinate phosphoribosyltransferase; acts in the salvage pathway of NAD+ biosynthesis; required for silencing at rDNA and telomeres and has a role in silencing at mating-type loci; localized to the nucleus. (429 aa) |
| | 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) |
| | ELG1 | Telomere length regulation protein ELG1; Subunit of an alternative replication factor C complex; important for DNA replication and genome integrity; suppresses spontaneous DNA damage; involved in homologous recombination-mediated repair and telomere homeostasis; required for PCNA (Pol30p) unloading during DNA replication; Belongs to the ELG1 family. (791 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) |
| | DIA2 | Protein DIA2; Origin-binding F-box protein; forms SCF ubiquitin ligase complex with Skp1p and Cdc53p; functions in ubiquitination of silent chromatin structural protein Sir4p; required to target Cdc6p for destruction during G1 phase; required for deactivation of Rad53 checkpoint kinase, completion of DNA replication during recovery from DNA damage, assembly of RSC complex, RSC-mediated transcription regulation, and nucleosome positioning; involved in invasive and pseudohyphal growth; Belongs to the DIA2 family. (732 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) |
| | SLD7 | Mitochondrial morphogenesis protein SLD7; Protein with a role in chromosomal DNA replication; interacts with Sld3p and reduces its affinity for Cdc45p; deletion mutant has aberrant mitochondria. (257 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) |
| | HST3 | NAD-dependent histone deacetylase HST3; Member of the Sir2 family of NAD(+)-dependent protein deacetylases; involved along with Hst4p in telomeric silencing, cell cycle progression, radiation resistance, genomic stability and short-chain fatty acid metabolism. (447 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) |
| | HRT1 | RING-box protein HRT1; RING-H2 domain core subunit of multiple ubiquitin ligase complexes; subunit of Skp1-Cullin-F-box (SCF) that tethers the Cdc34p (E2) and Cdc53p (cullin) SCF subunits, and is required for degradation of Gic2p, Far1p, Sic1p and Cln2p; subunit of the Rtt101p-Mms1p-Mms22p ubiquitin ligase that stabilizes replication forks after DNA lesions; subunit of the Cul3p-Elc1p-Ela1p ubiquitin ligase involved in Rpb1p degradation as part of transcription-coupled repair; Belongs to the RING-box family. (121 aa) |
| | NDJ1 | Non-disjunction protein 1; Protein that regulates meiotic SPB cohesion and telomere clustering; localizes to both spindle pole bodies (SPBs) and telomeres; required for bouquet formation, effective homolog pairing, ordered cross-over distribution, sister chromatid cohesion at meiotic telomeres, chromosomal segregation and telomere-led rapid prophase movement. (352 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) |
| | MRC1 | S-phase checkpoint protein required for DNA replication; couples DNA helicase and polymerase; interacts with and stabilizes Pol2p at stalled replication forks during stress, where it forms a pausing complex with Tof1p and is phosphorylated by Mec1p; defines a novel S-phase checkpoint with Hog1p that coordinates DNA replication and transcription upon osmostress; protects uncapped telomeres; Dia2p-dependent degradation mediates checkpoint recovery; mammalian claspin homolog. (1096 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) |
| | 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) |
| | IES3 | Subunit of the INO80 chromatin remodeling complex. (250 aa) |
| | CBF5 | H/ACA ribonucleoprotein complex subunit CBF5; Pseudouridine synthase catalytic subunit of box H/ACA snoRNPs; acts on large and small rRNAs, on snRNA U2, and on some mRNAs; mutations in human ortholog dyskerin cause the disorder dyskeratosis congenita; small nucleolar ribonucleoprotein particles are also known as snoRNPs. (483 aa) |
| | IFH1 | Protein IFH1; Coactivator, regulates transcription of ribosomal protein (RP) genes; recruited to RP gene promoters during optimal growth conditions via Fhl1p; subunit of CURI, a complex that coordinates RP production and pre-rRNA processing; regulated by acetylation and phosphorylation at different growth states via TORC1 signaling; IFH1 has a paralog, CRF1, that arose from the whole genome duplication. (1085 aa) |
| | EST1 | Telomere elongation protein EST1; TLC1 RNA-associated factor involved in telomere length regulation; recruitment subunit of telomerase; has G-quadruplex promoting activity required for telomere elongation; role in activating telomere-bound Est2p-TLC1-RNA; EST1 has a paralog, EBS1, that arose from the whole genome duplication. (699 aa) |
| | CMG1 | Uncharacterized protein YLR271W; Putative protein of unknown function; green fluorescent protein (GFP)-fusion protein localizes to the cytoplasm and the nucleus and is induced in response to the DNA-damaging agent MMS. (274 aa) |
| | YCS4 | Subunit of the condensin complex; required for establishment and maintenance of chromosome condensation, chromosome segregation, chromatin binding of condensin, tRNA gene clustering at the nucleolus, and silencing at the mating type locus; required for replication slow zone (RSZ) breakage following Mec1p inactivation; Belongs to the CND1 (condensin subunit 1) family. (1176 aa) |
| | MCM5 | Minichromosome maintenance protein 5; Component of the Mcm2-7 hexameric helicase complex; MCM complex is important for priming origins of DNA replication in G1 and becomes an active ATP-dependent helicase that promotes DNA melting and elongation when activated by Cdc7p-Dbf4p in S-phase. (775 aa) |
| | MEC3 | DNA damage and meiotic pachytene checkpoint protein; subunit of a heterotrimeric complex (Rad17p-Mec3p-Ddc1p) that forms a sliding clamp, loaded onto partial duplex DNA by a clamp loader complex; homolog of human and S. pombe Hus1. (474 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) |
| | EST2 | Reverse transcriptase subunit of the telomerase holoenzyme; essential for telomerase core catalytic activity, involved in other aspects of telomerase assembly and function; mutations in human homolog are associated with aplastic anemia. (884 aa) |
| | NUP2 | Nucleoporin involved in nucleocytoplasmic transport; binds to either the nucleoplasmic or cytoplasmic faces of the nuclear pore complex depending on Ran-GTP levels; also has a role in chromatin organization. (720 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) |
| | CST9 | Chromosome stability protein 9; SUMO E3 ligase; required for synaptonemal complex formation; localizes to synapsis initiation sites on meiotic chromosomes; associates with centromeres early in meiosis, then with chromosome axes and finally with double-strand break sites that are engaged in repair by crossovers; potential Cdc28p substrate. (482 aa) |
| | SIR3 | Regulatory protein SIR3; Silencing protein; interacts with Sir2p, Sir4p, and histone H3/H4 tails to establish transcriptionally silent chromatin; required for spreading of silenced chromatin; recruited to chromatin through interaction with Rap1p; C-terminus assumes variant winged helix-turn-helix (wH) fold that mediates homodimerization, which is critical for holo-SIR complex loading; required for telomere hypercluster formation in quiescent yeast cells; has paralog ORC1 from whole genome duplication. (978 aa) |
| | RIF2 | Protein that binds to the Rap1p C-terminus; acts synergistically with Rif1p to help control telomere length and establish telomeric silencing; deletion results in telomere elongation; RIF2 has a paralog, ORC4, that arose from the whole genome duplication. (395 aa) |
| | CGI121 | Component of the EKC/KEOPS complex; EKC/KEOPS complex is required for t6A tRNA modification and telomeric TG1-3 recombination; may have role in transcription; Cgi121p is dispensable for tRNA modification; other complex members are Bud32p, Kae1p, Pcc1p, and Gon7p; Belongs to the CGI121/TPRKB family. (181 aa) |
| | MFT1 | Subunit of the THO complex; THO is a nuclear complex comprised of Hpr1p, Mft1p, Rlr1p, and Thp2p, that is involved in transcription elongation and mitotic recombination; involved in telomere maintenance. (392 aa) |
| | CAC2 | Subunit of chromatin assembly factor I (CAF-1), with Rlf2p and Msi1p; chromatin assembly by CAF-1 is important for multiple processes including silencing at telomeres, mating type loci, and rDNA; maintenance of kinetochore structure, deactivation of the DNA damage checkpoint after DNA repair, chromatin dynamics during transcription; and repression of divergent transcription; relocalizes to the cytosol in response to hypoxia. (468 aa) |
| | GTR1 | GTP-binding protein GTR1; Subunit of a TORC1-stimulating GTPase complex; subunit of the heterodimeric Gtr1-Gtr2 GTPase complex that stimulates TORC1 in response to amino acid stimulation; tethered to the vacuolar membrane as part of the EGOC, a complex required for sorting of Gap1p and microautophagy; involved in phosphate transport and telomeric chromatin silencing; activated by the the Iml1p (GAP) subunit of the SEACIT complex; similar to human RagA and RagB. (310 aa) |
| | CTF13 | Subunit of the CBF3 complex; CBF3 binds to the CDE III element of centromeres, bending the DNA upon binding, and may be involved in sister chromatid cohesion during mitosis. (478 aa) |
| | YKU80 | ATP-dependent DNA helicase II subunit 2; Subunit of telomeric Ku complex (Yku70p-Yku80p); involved in telomere length maintenance, structure and telomere position effect; required for localization of telomerase ribonucleoprotein via interaction with TLC1 guide RNA; relocates to sites of double-strand cleavage to promote nonhomologous end joining during DSB repair; colocalizes with quiescent cell telomere hyperclusters; Belongs to the ku80 family. (629 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) |
| | SAS2 | Histone acetyltransferase (HAT) catalytic subunit of the SAS complex; acetylates free histones and nucleosomes and regulates transcriptional silencing; member of the MYSTacetyltransferase family; other members are Sas4p and Sas5p. (338 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) |
| | SPT21 | Protein with a role in transcriptional silencing; required for normal transcription at several loci including HTA2-HTB2 and HHF2-HHT2, but not required at the other histone loci; functionally related to Spt10p; localizes to nuclear foci that become diffuse upon DNA replication stress. (758 aa) |
| | ESC1 | Silent chromatin protein ESC1; Protein involved in telomeric silencing; required for quiescent cell telomere hypercluster localization at nuclear membrane vicinity; interacts with PAD4-domain of Sir4p. (1658 aa) |
| | RNA1 | GTPase activating protein (GAP) for Gsp1p; involved in nuclear transport; Belongs to the RNA1 family. (407 aa) |
| | RKR1 | RING domain E3 ubiquitin ligase; involved in ubiquitin-mediated degradation of non-stop proteins and translationally stalled ER membrane proteins; component of ribosome-bound RQC (ribosome quality control) complex; degrades products of mRNAs lacking a termination codon regardless of a poly(A) tail; functional connections to chromatin modification; homolog of mouse Listerin, mutations in which reported to cause neurodegeneration; Belongs to the LTN1 family. (1562 aa) |
| | YKU70 | ATP-dependent DNA helicase II subunit 1; Subunit of the telomeric Ku complex (Yku70p-Yku80p); involved in telomere length maintenance, structure and telomere position effect; required for localization of telomerase ribonucleoprotein to nucleus via interaction with the TLC1 guide RNA; relocates to sites of double-strand cleavage to promote nonhomologous end joining during DSB repair. (602 aa) |
| | HHT2 | Histone H3; core histone protein required for chromatin assembly, part of heterochromatin-mediated telomeric and HM silencing; one of two identical histone H3 proteins (see HHT1); regulated by acetylation, methylation, and phosphorylation; H3K14 acetylation plays an important role in the unfolding of strongly positioned nucleosomes during repair of UV damage. (136 aa) |
| | YAF9 | Protein AF-9 homolog; Subunit of NuA4 histone H4 acetyltransferase and SWR1 complexes; may function to antagonize silencing near telomeres; interacts directly with Swc4p; has homology to human leukemogenic protein AF9; contains a YEATS domain. (226 aa) |
| | FYV6 | Protein of unknown function; required for survival upon exposure to K1 killer toxin; proposed to regulate double-strand break repair via non-homologous end-joining. (173 aa) |
| | THO2 | Subunit of the THO complex; THO is required for efficient transcription elongation and involved in transcriptional elongation-associated recombination; required for LacZ RNA expression from certain plasmids; Belongs to the THOC2 family. (1597 aa) |
| | UBP10 | Ubiquitin carboxyl-terminal hydrolase 10; Ubiquitin-specific protease, deubiquitinates Ub-protein moieties; interacts with proteins that function in rRNA production and ribosome biogenesis via its intrinsically disordered regions; stabilizes Rpa190p by deubiquitination; controls PCNA deubiquitylation; may regulate silencing by acting on Sir4p; involved in posttranscriptionally regulating Gap1p, possibly other transporters; localized to the nucleolus; null mutant phenotypes are functionally complemented by human USP36; Belongs to the peptidase C19 family. (792 aa) |
| | RAP1 | DNA-binding protein RAP1; Essential DNA-binding transcription regulator that binds many loci; involved in transcription activation, repression, chromatin silencing, telomere length maintenance; relocalizes to cytosol under hypoxia; conserved protein with N-terminal BRCT domain, central region with homology to Myb DNA binding domain, and C-terminal Rap1-specific protein-interaction domain (RCT domain); recruits Sir complex to telomeric DNA; present in quiescent cell telomere hyperclusters. (827 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) |
| | RFA2 | Subunit of heterotrimeric Replication Protein A (RPA); RPA is a highly conserved single-stranded DNA binding protein involved in DNA replication, repair, and recombination; RPA protects against inappropriate telomere recombination, and upon telomere uncapping, prevents cell proliferation by a checkpoint-independent pathway; in concert with Sgs1p-Top2p-Rmi1p, stimulates DNA catenation/decatenation activity of Top3p; protein abundance increases in response to DNA replication s. (273 aa) |
| | PSH1 | RING finger protein PSH1; E3 ubiquitin ligase targeting centromere-binding protein Cse4p; mediates polyubiquitination and degradation of histone H3 variant Cse4p, preventing its mislocalization to euchromatin independent of Slx5p; ubiquitination of Cse4p may be antagonized by Scm3p. (406 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) |
| | SDC1 | COMPASS component SDC1; Subunit of the COMPASS (Set1C) complex; COMPASS methylates lysine 4 of histone H3 and is required in chromatin silencing at telomeres; contains a Dpy-30 domain that mediates interaction with Bre2p; similar to C. elegans and human DPY-30. (175 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) |
| | MCM3 | DNA replication licensing factor MCM3; Protein involved in DNA replication; component of the Mcm2-7 hexameric helicase complex that binds chromatin as a part of the pre-replicative complex. (971 aa) |
| | HAT2 | Subunit of the Hat1p-Hat2p histone acetyltransferase complex; required for high affinity binding of the complex to free histone H4, thereby enhancing Hat1p activity; similar to human RbAp46 and 48; has a role in telomeric silencing. (401 aa) |
| | CIN8 | Kinesin-like protein CIN8; Kinesin motor protein; involved in mitotic spindle assembly and chromosome segregation. (1000 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) |
| | DOT6 | Transcriptional regulatory protein DOT6; Protein involved in rRNA and ribosome biogenesis; activated in stochastic pulses of nuclear localization; binds polymerase A and C motif; subunit of the RPD3L histone deacetylase complex; has chromatin specific SANT domain; involved in telomeric gene silencing and filamentation; relative distribution to the nucleus increases upon DNA replication stress; Belongs to the DOT6 family. (670 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) |
| | 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) |
| | SCS2 | Integral ER membrane protein, regulates phospholipid metabolism; one of 6 proteins (Ist2p, Scs2p, Scs22p, Tcb1p, Tcb2p, Tcb3p) that connect ER to plasma membrane (PM) and regulate PI4P levels by controlling access of Sac1p phosphatase to substrate PI4P in the PM; interacts with FFAT motifs in Opi1p, Swh1p, Osh2p, and Osh3p; involved in telomeric silencing; VAP homolog; SCS2 has a paralog, SCS22, that arose from the whole genome duplication; Belongs to the VAMP-associated protein (VAP) (TC 9.B.17) family. (244 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) |
| | 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) |
| | PNC1 | Nicotinamidase that converts nicotinamide to nicotinic acid; part of the NAD(+) salvage pathway; required for life span extension by calorie restriction; lacks a peroxisomal targeting signal but is imported into peroxisomes via binding to Gpd1p; PNC1 expression responds to all known stimuli that extend replicative life span; protein increases in abundance and relative distribution to cytoplasmic foci decreases upon DNA replication stress. (216 aa) |
| | RAD6 | Ubiquitin-conjugating enzyme (E2); involved in postreplication repair as a heterodimer with Rad18p, regulation of K63 polyubiquitination in response to oxidative stress, DSBR and checkpoint control as a heterodimer with Bre1p, ubiquitin-mediated N-end rule protein degradation as a heterodimer with Ubr1p, ERAD with Ubr1p in the absence of canonical ER membrane ligases, and Rpn4p turnover as part of proteasome homeostasis, in complex with Ubr2p and Mub1p. (172 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) |
| | 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) |
| | MPS2 | Monopolar spindle protein 2; Essential membrane protein localized at nuclear envelope and SPBs; required for insertion of the newly duplicated spindle pole body into the nuclear envelope; potentially phosphorylated by Cdc28p; MPS2 has a paralog, CSM4, that arose from the whole genome duplication. (387 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) |
| | NUP145 | Nucleoporin NUP145C; Essential protein with distinct roles in two nuclear pore subcomplexes; catalyzes its own proteolytic cleavage in vivo to generate a C-terminal fragment that is a structural component of the Nup84p subcomplex (with roles in NPC biogenesis and localization of genes to the nuclear periphery), and an N-terminal fragment that is one of several FG-nucleoporins within the NPC central core directly responsible for nucleocytoplasmic transport; homologous to human NUP98. (1317 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) |
| | SLD3 | Protein involved in the initiation of DNA replication; required for proper assembly of replication proteins at the origins of replication; interacts with Cdc45p; localizes to nuclear foci that become diffuse upon DNA replication stress; homologous to the human Treslin/Ticrr protein. (668 aa) |
| | ITC1 | Imitation switch two complex protein 1; Subunit of ATP-dependent Isw2p-Itc1p chromatin remodeling complex; required for repression of a-specific genes, repression of early meiotic genes during mitotic growth, and repression of INO1; similar to mammalian Acf1p, the regulatory subunit of the mammalian ATP-utilizing chromatin assembly and modifying factor (ACF) complex; ITC1 has a paralog, YPL216W, that arose from the whole genome duplication. (1264 aa) |
| | INO80 | Chromatin-remodeling ATPase INO80; ATPase and nucleosome spacing factor; subunit of complex containing actin and actin-related proteins that has chromatin remodeling activity and 3' to 5' DNA helicase activity in vitro; promotes nucleosome shifts in the 3 prime direction; has a role in modulating stress gene transcription. (1489 aa) |
| | SUA5 | Threonylcarbamoyl-AMP synthase; Protein involved in threonylcarbamoyl adenosine biosynthesis; Sua5p and Qri7p are necessary and sufficient for RNA t6A modification in vitro; null mutant lacks N6-threonylcarbamoyl adenosine (t6A) modification in the anticodon loop of ANN-decoding tRNA; member of conserved YrdC/Sua5 family; binds single-stranded telomeric DNA and null mutant has abnormal telomere length. (426 aa) |
| | MTE1 | Uncharacterized protein YGR042W; Protein of unknown function; involved in maintenance of proper telomere length; green fluorescent protein (GFP)-fusion protein localizes to both the cytoplasm and the nucleus; forms nuclear foci upon DNA replication stress. (271 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) |
| | TEL2 | Telomere length regulation protein TEL2; Subunit of the ASTRA complex, involved in chromatin remodeling; subunit of the telomere cap complex DNA-binding protein specific to single-stranded yeast telomeric DNA repeats, required for telomere length regulation and telomere position effect; involved in the stability or biogenesis of PIKKs such as TORC1; Belongs to the TEL2 family. (688 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) |
| | CBF2 | Essential kinetochore protein; component of the CBF3 multisubunit complex that binds to the CDEIII region of the centromere; Cbf2p also binds to the CDEII region possibly forming a different multimeric complex, ubiquitinated in vivo; sumoylated in an Mms21p-dependent manner; relative distribution to the spindle pole body decreases upon DNA replication stress. (956 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) |
| | 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) |
| | CRM1 | Exportin-1; Major karyopherin; involved in export of proteins, RNAs, and ribosomal subunits from the nucleus; exportin. (1084 aa) |
| | GCN5 | Catalytic subunit of ADA and SAGA histone acetyltransferase complexes; modifies N-terminal lysines on histones H2B and H3; acetylates Rsc4p, a subunit of the RSC chromatin-remodeling complex, altering replication stress tolerance; relocalizes to the cytosol in response to hypoxia; mutant displays reduced transcription elongation in the G-less-based run-on (GLRO) assay; greater involvement in repression of RNAPII-dependent transcription than in activation; Belongs to the acetyltransferase family. GCN5 subfamily. (439 aa) |
| | BUD32 | Protein kinase; component of the EKC/KEOPS complex with Kae1p, Cgi121p, Pcc1p, and Gon7p; Pyrococcus Bud32 ortholog functions as a P-loop ATPase rather than a protein kinase in the context of the complex; EKC/KEOPS complex is required for t6A tRNA modification and telomeric TG1-3 recombination; may have role in transcription; mutation is functionally complemented by human TP53RK. (261 aa) |
| | YTA7 | Tat-binding homolog 7; Protein that localizes to chromatin; has a role in regulation of histone gene expression; has a bromodomain-like region that interacts with the N-terminal tail of histone H3, and an ATPase domain; relocalizes to the cytosol in response to hypoxia; potentially phosphorylated by Cdc28p. (1379 aa) |
