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| | KCC4 | Probable serine/threonine-protein kinase KCC4; Protein kinase of the bud neck involved in the septin checkpoint; associates with septin proteins, negatively regulates Swe1p by phosphorylation, shows structural homology to bud neck kinases Gin4p and Hsl1p; KCC4 has a paralog, GIN4, that arose from the whole genome duplication. (1037 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) |
| | 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) |
| | 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) |
| | 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) |
| | DUN1 | DNA damage response protein kinase DUN1; Cell-cycle checkpoint S/T protein kinase; required for transient G2/M arrest after DNA damage, damage-induced transcription, and nuclear-to-cytoplasmic redistribution of Rnr2p-Rnr4p after genotoxic stress and iron deprivation; phosphorylates repair protein Rad55p, transcriptional repressor Sml1p, superoxide dismutase, and ribonucleotide reductase inhibitors Crt1p and Dif1p; functions in the Mec1p pathway to regulate dNTP pools and telomere length; postreplicative repair role; Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase [...] (513 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) |
| | PPH3 | Catalytic subunit of protein phosphatase PP4 complex; Pph3p and Psy2p form active complex, Psy4p may provide substrate specificity; regulates recovery from the DNA damage checkpoint, the gene conversion- and single-strand annealing-mediated pathways of meiotic double-strand break repair and efficient Non-Homologous End-Joining (NHEJ) pathway; involved in activation of Gln3p to alleviate nitrogen catabolite repression; Pph3p and Psy2p localize to foci on meiotic chromosomes. (308 aa) |
| | BMH2 | 14-3-3 protein, minor isoform; controls proteome at post-transcriptional level, binds proteins and DNA, involved in regulation of many processes including exocytosis, vesicle transport, Ras/MAPK signaling, and rapamycin-sensitive signaling; protein increases in abundance and relative distribution to the nucleus increases upon DNA replication stress; abundance relative to Bmh1p increases during sporulation. (273 aa) |
| | RAD9 | DNA damage-dependent checkpoint protein; required for cell-cycle arrest in G1/S, intra-S, and G2/M, plays a role in postreplication repair (PRR) pathway; transmits checkpoint signal by activating Rad53p and Chk1p; hyperphosphorylated by Mec1p and Tel1p; multiple cyclin dependent kinase consensus sites and the C-terminal BRCT domain contribute to DNA damage checkpoint activation; Rad9p Chk1 Activating Domain (CAD) is phosphorylated at multiple sites by Cdc28p/Clb2p. (1309 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) |
| | GCN2 | eIF-2-alpha kinase GCN2; Protein kinase; phosphorylates the alpha-subunit of translation initiation factor eIF2 (Sui2p) in response to starvation; activated by uncharged tRNAs and the Gcn1p-Gcn20p complex; contributes to DNA damage checkpoint control. (1659 aa) |
| | ESC2 | Sumo-like domain protein; prevents accumulation of toxic intermediates during replication-associated recombinational repair; roles in silencing, lifespan, chromatid cohesion and the intra-S-phase DNA damage checkpoint; RENi family member. (456 aa) |
| | MUS81 | Subunit of structure-specific Mms4p-Mus81p endonuclease; cleaves branched DNA; involved in DNA repair, replication fork stability, and joint molecule formation/resolution during meiotic recombination; promotes template switching during break-induced replication (BIR), causing non-reciprocal translocations (NRTs); helix-hairpin-helix protein; phosphorylation of non-catalytic subunit Mms4p by Cdc28p and Cdcp during mitotic cell cycle activates function of Mms4p-Mus81p; Belongs to the XPF family. (632 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) |
| | LCD1 | Essential protein required for the DNA integrity checkpoint pathways; interacts physically with Mec1p; putative homolog of S. pombe Rad26 and human ATRIP; forms nuclear foci upon DNA replication stress. (747 aa) |
| | GIN4 | Serine/threonine-protein kinase GIN4; Protein kinase involved in bud growth and assembly of the septin ring; proposed to have kinase-dependent and kinase-independent activities; undergoes autophosphorylation; similar to Hsl1p; GIN4 has a paralog, KCC4, that arose from the whole genome duplication. (1142 aa) |
| | BIM1 | Protein BIM1; Microtubule plus end-tracking protein; together with Kar9p makes up the cortical microtubule capture site and delays the exit from mitosis when the spindle is oriented abnormally; homolog of human end binding protein 1 (EB1); Belongs to the MAPRE family. (344 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) |
| | BEM2 | GTPase-activating protein BEM2/IPL2; Rho GTPase activating protein (RhoGAP); involved in the control of cytoskeleton organization and cellular morphogenesis; required for bud emergence; potential GAP for Rho4p. (2167 aa) |
| | RAD24 | Checkpoint protein; involved in the activation of the DNA damage and meiotic pachytene checkpoints; subunit of a clamp loader that loads Rad17p-Mec3p-Ddc1p onto DNA; homolog of human and S. pombe Rad17 protein. (659 aa) |
| | BMH1 | 14-3-3 protein, major isoform; controls proteome at post-transcriptional level, binds proteins and DNA, involved in regulation of exocytosis, vesicle transport, Ras/MAPK and rapamycin-sensitive signaling, aggresome formation, spindle position checkpoint; protein increases in abundance and relative distribution to the nucleus increases upon DNA replication stress; antiapoptotic gene similar to human 14-3-3; BMH1 has a paralog, BMH2, that arose from whole genome duplication. (267 aa) |
| | YPI1 | Regulatory subunit of the type I protein phosphatase (PP1) Glc7p; Glc7p participates in the regulation of a variety of metabolic processes including mitosis and glycogen metabolism; in vitro evidence suggests Ypi1p is an inhibitor of Glc7p while in vivo evidence suggests it is an activator; overproduction causes decreased cellular content of glycogen; partial depletion causes lithium sensitivity, while overproduction confers lithium-tolerance. (155 aa) |
| | ALK1 | Serine/threonine-protein kinase Haspin homolog ALK1; Protein kinase; along with its paralog, ALK2, required for proper spindle positioning and nuclear segregation following mitotic arrest, proper organization of cell polarity factors in mitosis, proper localization of formins and polarity factors, and survival in cells that activate spindle assembly checkpoint; phosphorylated in response to DNA damage; ALK1 has a paralog, ALK2, that arose from the whole genome duplication; similar to mammalian haspins. (760 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) |
| | 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) |
| | 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) |
| | CDC20 | Activator of anaphase-promoting complex/cyclosome (APC/C); APC/C is required for metaphase/anaphase transition; directs ubiquitination of mitotic cyclins, Pds1p, and other anaphase inhibitors; cell-cycle regulated; potential Cdc28p substrate; relative distribution to the nucleus increases upon DNA replication stress; Belongs to the WD repeat CDC20/Fizzy family. (610 aa) |
| | RCK1 | Serine/threonine-protein kinase RCK1; Protein kinase involved in oxidative stress response; promotes pseudohyphal growth via activation of Ubp3p phosphorylation; identified as suppressor of S. pombe cell cycle checkpoint mutations; RCK1 has a paralog, RCK2, that arose from the whole genome duplication. (512 aa) |
| | CDC55 | Regulatory subunit B of protein phosphatase 2A (PP2A); Zds1p/2p-dependent localization to cytoplasm promotes mitotic entry; localization to nucleus prevents mitotic exit; required for correct nuclear division, chromosome segregation during achiasmate meiosis; maintains nucleolar sequestration of Cdc14p during early meiosis; limits formation of PP2A-Rts1p holocomplexes to ensure timely dissolution of sister chromosome cohesion; homolog of mammalian B55. (526 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) |
| | AMA1 | Meiosis-specific APC/C activator protein AMA1; Activator of meiotic anaphase promoting complex (APC/C); Cdc20p family member; required for initiation of spore wall assembly; required for Clb1p degradation during meiosis; prevents premature assembly of the meiosis I spindle, required for DSB induced prophase I arrest; Belongs to the WD repeat CDC20/Fizzy family. (593 aa) |
| | DMA1 | Ubiquitin-protein ligase (E3); controls septin dynamics, spindle position checkpoint (SPOC) with ligase Dma2p by regulating recruitment of Elm1p to bud neck; regulates levels of eIF2 subunit Gcd11p, as well as abundance, localization, and ubiquitination of Cdk inhibitory kinase Swe1p; ubiquitinates cyclin Pcl1p; ortholog of human RNF8, similar to human Chfr; contains FHA, RING fingers; DMA1 has a paralog, DMA2, that arose from the whole genome duplication. (416 aa) |
| | ARP1 | Centractin; Actin-related protein of the dynactin complex; required for spindle orientation and nuclear migration; forms actin-like short filament composed of 9 or 10 Arp1p monomers; putative ortholog of mammalian centractin. (384 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) |
| | ULP2 | Ubiquitin-like-specific protease 2; Peptidase that deconjugates Smt3/SUMO-1 peptides from proteins; plays a role in chromosome cohesion at centromeric regions and recovery from checkpoint arrest induced by DNA damage or DNA replication defects; potential Cdc28p substrate; human homolog PML implicated in promyelocytic leukemia can partially complement yeast null mutant; Belongs to the peptidase C48 family. (1034 aa) |
| | HOP1 | Meiosis-specific protein required for chromosome synapsis; displays Red1p dependent localization to the unsynapsed axial-lateral elements of the synaptonemal complex; required for chiasma formation; in vitro, displays the ability to promote intra- and intermolecular synapsis between double-stranded DNA molecules and to fold DNA into rigid protein-DNA filaments. (605 aa) |
| | MAD3 | Subunit of spindle-assembly checkpoint complex; involved in delaying anaphase onset in cells with defects in mitotic spindle assembly; pseudosubstrate inhibitor of APC(Cdc20), the anaphase promoting complex involved in securin (Pds1p) turnover; MAD3 has a paralog, BUB1, that arose from the whole genome duplication. (515 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) |
| | DPB11 | DNA replication initiation protein; loads DNA pol epsilon onto pre-replication complexes at origins; checkpoint sensor recruited to stalled replication forks by the checkpoint clamp complex where it activates Mec1p; along with Rfa1p, binds to ultrafine anaphase bridges in mitotic cells and prevents accumulation of chromatin bridges by stimulating the Mec1p kinase and suppressing homologous recombination; ortholog of human TopBP1; forms nuclear foci upon DNA replication stress. (764 aa) |
| | SWE1 | Mitosis inhibitor protein kinase SWE1; Protein kinase that regulates the G2/M transition; negative regulator of the Cdc28p kinase; morphogenesis checkpoint kinase; positive regulator of sphingolipid biosynthesis via Orm2p; phosphorylates a tyrosine residue in the N-terminus of Hsp90 in a cell-cycle associated manner, thus modulating the ability of Hsp90 to chaperone a selected clientele; localizes to the nucleus and to the daughter side of the mother-bud neck; homolog of S. pombe Wee1p; potential Cdc28p substrate. (819 aa) |
| | CDC6 | Cell division control protein 6; Essential ATP-binding protein required for DNA replication; component of the pre-replicative complex (pre-RC) which requires ORC to associate with chromatin and is in turn required for Mcm2-7p DNA association; homologous to S. pombe Cdc18p; relocalizes from nucleus to cytoplasm upon DNA replication stress; degraded in response to plasma membrane stress. (513 aa) |
| | BFA1 | Mitotic check point protein BFA1; Subunit of a two-component GTPase-activating protein, Bfa1p-Bub2p; contributes to GAP activity, inactivating Tem1 by stimulating GTP hydrolysis following damage or misalignment of the mitotic spindle; functions as a guanine-nucleotide exchange inhibitor (GDI) for Tem1p; involved in multiple cell cycle checkpoint pathways that control mitotic exit; required when telomeres are damaged, but not for all types of chromosomal DNA damage; phosphorylated by the Polo-like kinase Cdc5p; To S.pombe byr4. (574 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) |
| | RFC2 | Subunit of heteropentameric Replication factor C (RF-C); RF-C is a DNA binding protein and ATPase that acts as a clamp loader of the proliferating cell nuclear antigen (PCNA) processivity factor for DNA polymerases delta and epsilon. (353 aa) |
| | BUD2 | Inhibitory regulator protein BUD2/CLA2; GTPase activating factor for Rsr1p/Bud1p; plays a role in spindle position checkpoint distinct from its role in bud site selection; required for both axial and bipolar budding patterns; mutants exhibit random budding in all cell types; contains two PH-like domains. (1104 aa) |
| | HSL1 | Probable serine/threonine-protein kinase HSL1; Nim1p-related protein kinase; septin-binding kinase that localizes to the bud neck septin ring and regulates the morphogenesis checkpoint; phosphorylates Hsl7p and cooperates with Elm1p to recruit Hsl7p to the mother-bud neck, as a prerequisite for the subsequent recruitment, phosphorylation, and degradation of Swe1p; autophosphorylation enhances interactions with Hsl7p. (1518 aa) |
| | SLD2 | DNA replication regulator SLD2; Single-stranded DNA origin-binding and annealing protein; required for initiation of DNA replication; phosphorylated in S phase by cyclin-dependent kinases (Cdks), promoting origin binding, DNA replication and Dpb11p complex formation; component of the preloading complex; binds the Mcm2-7p complex to prevent inappropriate Mcm2-7p interaction with the GINS complex in G1; required for S phase checkpoint; relative distribution to the nucleus increases upon DNA replication stress; Belongs to the SLD2 family. (453 aa) |
| | RED1 | Protein component of the synaptonemal complex axial elements; involved in chromosome segregation during the first meiotic division; critical for coupling checkpoint signaling to SC formation; promotes interhomolog recombination by phosphorylating Hop1p; also interacts with Mec3p and Ddc1p. (827 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) |
| | 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) |
| | TSA1 | Peroxiredoxin TSA1; Thioredoxin peroxidase; acts as both ribosome-associated and free cytoplasmic antioxidant; self-associates to form high-molecular weight chaperone complex under oxidative stress; chaperone activity essential for growth in zinc deficiency; required for telomere length maintenance; binds and modulates Cdc19p activity; protein abundance increases, forms cytoplasmic foci during DNA replication stress; TSA1 has a paralog, TSA2, that arose from the whole genome duplication; Belongs to the peroxiredoxin family. AhpC/Prx1 subfamily. (196 aa) |
| | HUG1 | MEC1-mediated checkpoint protein HUG1; Ribonucleotide reductase inhibitor; intrinsically disordered protein that binds to and inhibits Rnr2p; involved in the Mec1p-mediated checkpoint pathway; transcription is induced by genotoxic stress and by activation of the Rad53p pathway; protein abundance increases in response to DNA replication stress. (68 aa) |
| | SPC72 | Spindle pole component SPC72; Gamma-tubulin small complex (gamma-TuSC) receptor; recruits the gamma-TuSC complex to the cytoplasmic side of the SPB, connecting nuclear microtubules to the SPB; involved in astral microtubule formation, stabilization, and with Stu2p, anchoring astral MTs at the cytoplasmic face of the SPB, and regulating plus-end MT dynamics; regulated by Cdc5 kinase. (622 aa) |
| | TEM1 | GTP-binding protein of the Ras superfamily; involved in termination of M-phase; controls actomyosin and septin dynamics during cytokinesis. (245 aa) |
| | ORC1 | Largest subunit of the origin recognition complex; involved in directing DNA replication by binding to replication origins; also involved in transcriptional silencing; exhibits ATPase activity; ORC1 has a paralog, SIR3, that arose from the whole genome duplication. (914 aa) |
| | CCR4 | Glucose-repressible alcohol dehydrogenase transcriptional effector; Component of the CCR4-NOT transcriptional complex; CCR4-NOT is involved in regulation of gene expression; component of the major cytoplasmic deadenylase, which is involved in mRNA poly(A) tail shortening; Belongs to the CCR4/nocturin family. (837 aa) |
| | TPD3 | Regulatory subunit A of the heterotrimeric PP2A complex; the heterotrimeric protein phosphatase 2A (PP2A) complex also contains regulatory subunit Cdc55p and either catalytic subunit Pph21p or Pph22p; required for cell morphogenesis and transcription by RNA polymerase III. (635 aa) |
| | FPR3 | FK506-binding nuclear protein; Nucleolar peptidyl-prolyl cis-trans isomerase (PPIase); FK506 binding protein; affects expression of multiple genes via its role in nucleosome assembly; phosphorylated by casein kinase II (Cka1p-Cka2p-Ckb1p-Ckb2p) and dephosphorylated by Ptp1p; PPIase domain acts as a transcriptional repressor when tethered to DNA by lexA, and repressor activity is dependent on PPIase activity; FPR3 has a paralog, FPR4, that arose from the whole genome duplication; Belongs to the FKBP-type PPIase family. FKBP3/4 subfamily. (411 aa) |
| | CSM3 | Chromosome segregation in meiosis protein 3; Replication fork associated factor; required for stable replication fork pausing; component of the DNA replication checkpoint pathway; required for accurate chromosome segregation during meiosis; forms nuclear foci upon DNA replication stress. (317 aa) |
| | BUB2 | Mitotic check point protein BUB2; Mitotic exit network regulator; forms GTPase-activating Bfa1p-Bub2p complex that binds Tem1p and spindle pole bodies, blocks cell cycle progression before anaphase in response to spindle and kinetochore damage; Belongs to the BUB2 family. (306 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) |
| | SGS1 | ATP-dependent helicase SGS1; RecQ family nucleolar DNA helicase; role in genome integrity maintenance, chromosome synapsis, meiotic joint molecule/crossover formation; stimulates activity of Top3p; rapidly lost in response to rapamycin in Rrd1p-dependent manner; forms nuclear foci upon DNA replication stress; yeast SGS1 complements mutations in human homolog BLM implicated in Bloom syndrome; also similar to human WRN implicated in Werner syndrome; human BLM and WRN can each complement yeast null mutant; Belongs to the helicase family. RecQ subfamily. (1447 aa) |
| | MRE11 | Double-strand break repair protein MRE11; Nuclease subunit of the MRX complex with Rad50p and Xrs2p; complex functions in repair of DNA double-strand breaks and in telomere stability; Mre11p associates with Ser/Thr-rich ORFs in premeiotic phase; nuclease activity required for MRX function; widely conserved; forms nuclear foci upon DNA replication stress; Belongs to the MRE11/RAD32 family. (692 aa) |
| | DMA2 | Ubiquitin-protein ligase (E3); controls septin dynamics and spindle position checkpoint (SPOC) with ligase Dma1p by regulating recruitment of Elm1p to bud neck; regulates levels of eIF2 subunit Gcd11p, as well as abundance, localization, and ubiquitination of Cdk inhibitory kinase Swe1p; ortholog of human RNF8, similar to human Chfr; contains FHA and RING finger domains; DMA2 has a paralog, DMA1, that arose from the whole genome duplication. (522 aa) |
| | FAR11 | Factor arrest protein 11; Protein involved in recovery from cell cycle arrest; acts in response to pheromone; also involved in regulation of intra-S DNA damage checkpoint and autophagy; is essential for dephosphorylation of Atg13p; interacts with Far3p, Far7p, Far8p, Far9p, Far10p and with the phosphatases Pph21p, Pph22p and Pph3p; has similarity to the N- and C-termini of N. crassa HAM-2; similar to human Fam40A and Fam40B; Belongs to the FAR11 family. (953 aa) |
| | IBD2 | Protein IBD2; Component of the BUB2-dependent spindle checkpoint pathway; interacts with Bfa1p and functions upstream of Bub2p and Bfa1p. (351 aa) |
| | PSY2 | Subunit of protein phosphatase PP4 complex; Pph3p and Psy2p form the active complex, Psy4p may provide additional substrate specificity; regulates recovery from the DNA damage checkpoint, the gene conversion- and single-strand annealing-mediated pathways of meiotic double-strand break repair and efficient Non-Homologous End-Joining (NHEJ) pathway; Pph3p and Psy2p localize to foci on meiotic chromosomes; putative homolog of mammalian R3. (858 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) |
| | POL2 | Catalytic subunit of DNA polymerase (II) epsilon; a chromosomal DNA replication polymerase that exhibits processivity and proofreading exonuclease activity; participates in leading-strand synthesis during DNA replication; also involved in DNA synthesis during DNA repair; interacts extensively with Mrc1p. (2222 aa) |
| | TOF1 | Topoisomerase 1-associated factor 1; Subunit of a replication-pausing checkpoint complex; Tof1p-Mrc1p-Csm3p acts at the stalled replication fork to promote sister chromatid cohesion after DNA damage, facilitating gap repair of damaged DNA; interacts with the MCM helicase; relocalizes to the cytosol in response to hypoxia. (1238 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) |
| | 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) |
| | EXO1 | Exodeoxyribonuclease 1; 5'-3' exonuclease and flap-endonuclease; involved in recombination, double-strand break repair, MMS2 error-free branch of the post replication (PRR) pathway and DNA mismatch repair; role in telomere maintenance; member of the Rad2p nuclease family, with conserved N and I nuclease domains; relative distribution to the nucleus increases upon DNA replication stress; EXO1 has a paralog, DIN7, that arose from the whole genome duplication. (702 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) |
| | GAC1 | Regulatory subunit for Glc7p type-1 protein phosphatase (PP1); tethers Glc7p to Gsy2p glycogen synthase, binds Hsf1p heat shock transcription factor, required for induction of some HSF-regulated genes under heat shock; GAC1 has a paralog, PIG1, that arose from the whole genome duplication. (793 aa) |
| | KIN4 | Serine/threonine-protein kinase KIN4; Serine/threonine protein kinase; inhibits the mitotic exit network (MEN) when the spindle position checkpoint is activated; localized asymmetrically to mother cell cortex, spindle pole body and bud neck; KIN4 has a paralog, FRK1, that arose from the whole genome duplication. (800 aa) |
| | MEK1 | Meiosis-specific serine/threonine protein kinase; functions in meiotic checkpoint, promotes recombination between homologous chromosomes by suppressing double strand break repair between sister chromatids; stabilizes Hop1-Thr318 phosphorylation to promote interhomolog recombination and checkpoint responses during meiosis. (497 aa) |
| | RAD17 | Checkpoint protein; involved in the activation of the DNA damage and meiotic pachytene checkpoints; with Mec3p and Ddc1p, forms a clamp that is loaded onto partial duplex DNA; homolog of human and S. pombe Rad1 and U. maydis Rec1 proteins. (401 aa) |
| | CHL1 | Probable DNA helicase; involved in sister-chromatid cohesion and genome integrity and interstrand cross-link repair; interacts with ECO1 and CTF18; mutants are defective in silencing, rDNA recombination, aging and the heat shock response; FANCJ-like helicase family member; mutations in the human homolog, DDX11/ChLR1, cause Warsaw breakage syndrome; Belongs to the DEAD box helicase family. DEAH subfamily. DDX11/CHL1 sub-subfamily. (861 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) |
| | RAD53 | Serine/threonine-protein kinase RAD53; DNA damage response protein kinase; required for cell-cycle arrest, regulation of copper genes in response to DNA damage; phosphorylates nuclear pores to counteract gene gating, preventing aberrant transitions at forks approaching transcribed genes; activates downstream kinase Dun1p; differentially senses mtDNA depletion, mitochondrial ROS; relocalizes to cytosol under hypoxia; human homolog CHEK2 implicated in breast cancer can complement yeast null mutant; Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. CHEK2 subfamily. (821 aa) |
| | DDC1 | DNA damage checkpoint protein; part of a PCNA-like complex required for DNA damage response, required for pachytene checkpoint to inhibit cell cycle in response to unrepaired recombination intermediates; potential Cdc28p substrate; forms nuclear foci upon DNA replication stress; Belongs to the DDC1 family. (612 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) |
| | 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) |
| | YPR015C | Putative zinc finger transcription factor; binds DNA in sequence-specific manner; overexpression causes a cell cycle delay or arrest. (247 aa) |
| | ALK2 | Serine/threonine-protein kinase Haspin homolog ALK2; Protein kinase; along with its paralog, ALK1, required for proper spindle positioning and nuclear segregation following mitotic arrest, proper organization of cell polarity factors in mitosis, proper localization of formins and polarity factors, and survival in cells that activate spindle assembly checkpoint; phosphorylated in response to DNA damage; ALK2 has a paralog, ALK1, that arose from the whole genome duplication; similar to mammalian haspins. (676 aa) |
| | PIN4 | RNA-binding protein PIN4; Protein involved in G2/M phase progression and response to DNA damage; interacts with Rad53p; contains an RNA recognition motif, a nuclear localization signal, and several SQ/TQ cluster domains; hyperphosphorylated in response to DNA damage. (668 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) |
| | MMS4 | Subunit of structure-specific Mms4p-Mus81p endonuclease; cleaves branched DNA; involved in recombination, DNA repair, and joint molecule formation/resolution during meiotic recombination; phosphorylation of the non-catalytic subunit Mms4p by Cdc28p and Cdc5p during mitotic cell cycle activates the function of Mms4p-Mus81p. (691 aa) |
| | MEC1 | Serine/threonine-protein kinase MEC1; Genome integrity checkpoint protein and PI kinase superfamily member; Mec1p and Dun1p function in same pathway to regulate dNTP pools and telomere length; signal transducer required for cell cycle arrest and transcriptional responses to damaged or unreplicated DNA; facilitates replication fork progression and regulates P-body formation under replication stress; promotes interhomolog recombination by phosphorylating Hop1p; associates with shortened, dysfunctional telomeres. (2368 aa) |
| | AMN1 | Antagonist of mitotic exit network protein 1; Protein required for daughter cell separation; multiple mitotic checkpoints, and chromosome stability; contains 12 degenerate leucine-rich repeat motifs; expression is induced by the Mitotic Exit Network (MEN); Belongs to the AMN1 family. (549 aa) |
| | PCH2 | Pachytene checkpoint protein 2; Hexameric ring ATPase that remodels chromosome axis protein Hop1p; nucleolar component of the pachytene checkpoint, which prevents chromosome segregation when recombination and chromosome synapsis are defective; also represses meiotic interhomolog recombination in rDNA; required for meiotic double-stranded break formation. (564 aa) |
| | CHK1 | Serine/threonine-protein kinase CHK1; Serine/threonine kinase and DNA damage checkpoint effector; mediates cell cycle arrest via phosphorylation of Pds1p; phosphorylated by checkpoint signal transducer Mec1p; homolog of S. pombe and mammalian Chk1 checkpoint kinase; Belongs to the protein kinase superfamily. CAMK Ser/Thr protein kinase family. NIM1 subfamily. (527 aa) |
