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| | 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) |
| | LTE1 | Guanine nucleotide exchange factor LTE1; Protein similar to GDP/GTP exchange factors; without detectable GEF activity; required for asymmetric localization of Bfa1p at daughter-directed spindle pole bodies and for mitotic exit at low temperatures. (1435 aa) |
| | CLN3 | G1/S-specific cyclin CLN3; G1 cyclin involved in cell cycle progression; activates Cdc28p kinase to promote G1 to S phase transition; plays a role in regulating transcription of other G1 cyclins, CLN1 and CLN2; regulated by phosphorylation and proteolysis; acetyl-CoA induces CLN3 transcription in response to nutrient repletion to promote cell-cycle entry; cell cycle arrest phenotype of the cln1 cln2 cln3 triple null mutant is complemented by any of human cyclins CCNA2, CCNB1, CCNC, CCND1, or CCNE1. (580 aa) |
| | CDC15 | Cell division control protein 15; Protein kinase of the Mitotic Exit Network; localized to the spindle pole bodies at late anaphase; promotes mitotic exit by directly switching on the kinase activity of Dbf2p; required for spindle disassembly after meiosis II; relocalizes to the cytoplasm upon DNA replication stress. (974 aa) |
| | PHO11 | One of three repressible acid phosphatases; glycoprotein that is transported to the cell surface by the secretory pathway; induced by phosphate starvation and coordinately regulated by PHO4 and PHO2; PHO11 has a paralog, PHO12, that arose from a segmental duplication. (467 aa) |
| | FUS3 | Mitogen-activated serine/threonine protein kinase involved in mating; phosphoactivated by Ste7p; substrates include Ste12p, Far1p, Bni1p, Sst2p; inhibits invasive growth during mating by phosphorylating Tec1p, promoting its; inhibits recruitment of Ste5p, Cdc42p-mediated asymmetry and mating morphogenesis. (353 aa) |
| | MCM2 | Protein involved in DNA replication; component of the Mcm2-7 hexameric helicase complex that binds chromatin as a part of the pre-replicative complex; relative distribution to the nucleus increases upon DNA replication stress. (868 aa) |
| | CDC27 | 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. (758 aa) |
| | BRN1 | Subunit of the condensin complex; required for chromosome condensation and for clustering of tRNA genes at the nucleolus; may influence multiple aspects of chromosome transmission. (754 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) |
| | PHO5 | Repressible acid phosphatase; 1 of 3 repressible acid phosphatases that also mediates extracellular nucleotide-derived phosphate hydrolysis; secretory pathway derived cell surface glycoprotein; induced by phosphate starvation and coordinately regulated by PHO4 and PHO2; Belongs to the histidine acid phosphatase family. (467 aa) |
| | CYC8 | General transcriptional co-repressor; acts together with Tup1p; also acts as part of a transcriptional co-activator complex that recruits the SWI/SNF and SAGA complexes to promoters; can form the prion [OCT+]. (966 aa) |
| | HSL7 | Protein arginine N-methyltransferase; exhibits septin and Hsl1p-dependent localization to the bud neck in budded cells and periodic Hsl1p-dependent phosphorylation; required with Hsl1p, and Elm1p for the mother-bud neck recruitment, phosphorylation, and degradation of Swe1p; interacts directly with Swe1p; relocalizes away from bud neck upon DNA replication stress; human homolog PRMT5 can complement yeast hsl7 mutant. (827 aa) |
| | CKS1 | Cyclin-dependent protein kinase regulatory subunit and adaptor; interacts with Cdc28p (aka Cdk1p); required for G1/S and G2/M phase transitions and budding; mediates phosphorylation and degradation of Sic1p; modulates proteolysis of M-phase targets through interactions with the proteasome; role in transcriptional regulation, recruiting proteasomal subunits to target gene promoters; human homologs CKS1B and CKS2 can each complement yeast cks1 null mutant. (150 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) |
| | CDC28 | Cyclin-dependent kinase (CDK) catalytic subunit; master regulator of mitotic and meiotic cell cycles; alternately associates with G1, S, G2/M phase cyclins, which provide substrate specificity; regulates metabolism, basal transcription, chromosome dynamics, growth and morphogenesis; transcript induction in osmostress involves antisense RNA; human homologs CDK1, CDK2, CDK3 can complement yeast conditional cdc28 mutants; human CDK1, CDK2 can complement yeast cdc28 null mutant. (298 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) |
| | 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) |
| | 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) |
| | TUP1 | General transcriptional corepressor TUP1; General repressor of transcription; forms complex with Cyc8p, involved in the establishment of repressive chromatin structure through interactions with histones H3 and H4, appears to enhance expression of some genes. (713 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) |
| | APC11 | Catalytic core subunit, Anaphase-Promoting Complex/Cyclosome (APC/C); which is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; contains a RING-H2 domain that is required for activity. (165 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) |
| | MBP1 | Transcription factor; involved in regulation of cell cycle progression from G1 to S phase, forms a complex with Swi6p that binds to MluI cell cycle box regulatory element in promoters of DNA synthesis genes. (833 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) |
| | PHO2 | Regulatory protein PHO2; Homeobox transcription factor; regulatory targets include genes involved in phosphate metabolism; binds cooperatively with Pho4p to the PHO5 promoter; phosphorylation of Pho2p facilitates interaction with Pho4p; relocalizes to the cytosol in response to hypoxia. (559 aa) |
| | PCL2 | PHO85 cyclin-2; Cyclin, interacts with cyclin-dependent kinase Pho85p; member of the Pcl1,2-like subfamily, involved in the regulation of polarized growth and morphogenesis and progression through the cell cycle; localizes to sites of polarized cell growth; PCL2 has a paralog, PCL9, that arose from the whole genome duplication. (308 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) |
| | PPH21 | Catalytic subunit of protein phosphatase 2A (PP2A); functionally redundant with Pph22p; methylated at C terminus; forms alternate complexes with several regulatory subunits; involved in signal transduction and regulation of mitosis; forms nuclear foci upon DNA replication stress; PPH21 has a paralog, PPH22, that arose from the whole genome duplication; Belongs to the PPP phosphatase family. PP-2A subfamily. (369 aa) |
| | CLB3 | G2/mitotic-specific cyclin-3; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the G2/M transition; may be involved in DNA replication and spindle assembly; accumulates during S phase and G2, then targeted for ubiquitin-mediated degradation; relative distribution to the nucleus increases upon DNA replication stress; CLB3 has a paralog, CLB4, that arose from the whole genome duplication. (427 aa) |
| | PCL9 | PHO85 cyclin-9; Cyclin; forms a functional kinase complex with Pho85p cyclin-dependent kinase (Cdk), expressed in late M/early G1 phase, activated by Swi5p; PCL9 has a paralog, PCL2, that arose from the whole genome duplication; Belongs to the cyclin family. PCL1,2 subfamily. (304 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) |
| | 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) |
| | FOB1 | Nucleolar protein that binds the rDNA replication fork barrier site; required for replication fork blocking, recombinational hotspot activity, condensin recruitment to replication fork barrier (RFB), and rDNA repeat segregation; related to retroviral integrases. (566 aa) |
| | PDS1 | Securin; inhibits anaphase by binding separin Esp1p; blocks cyclin destruction and mitotic exit, essential for meiotic progression and mitotic cell cycle arrest; localization is cell-cycle dependent and regulated by Cdc28p phosphorylation. (373 aa) |
| | APC4 | 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; component of the platform domain of the APC/C, based on structural analysis; relative distribution to the nucleus increases upon DNA replication stress. (652 aa) |
| | SWI5 | Transcriptional factor SWI5; Transcription factor that recruits Mediator and Swi/Snf complexes; activates transcription of genes expressed at the M/G1 phase boundary and in G1 phase; required for expression of the HO gene controlling mating type switching; localization to nucleus occurs during G1 and appears to be regulated by phosphorylation by Cdc28p kinase; SWI5 has a paralog, ACE2, that arose from the whole genome duplication. (709 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) |
| | 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) |
| | SWM1 | Subunit of the anaphase-promoting complex (APC); APC is an E3 ubiquitin ligase that regulates the metaphase-anaphase transition and exit from mitosis; required for activation of the daughter-specific gene expression and spore wall maturation; Belongs to the APC13 family. (170 aa) |
| | YCG1 | Subunit of the condensin complex; required for establishment and maintenance of chromosome condensation, chromosome segregation and chromatin binding by the complex; required for tRNA genes clustering at the nucleolus; required for replication slow zone breakage following Mec1p inactivation; transcription is cell cycle regulated, peaking in mitosis and declining in G1; protein is constitutively degraded by the proteasome; rate limiting for condensin recruitment to chromatin. (1035 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) |
| | YHP1 | Homeobox protein YHP1; Homeobox transcriptional repressor; binds Mcm1p and early cell cycle box (ECB) elements of cell cycle regulated genes, thereby restricting ECB-mediated transcription to the M/G1 interval; YHP1 has a paralog, YOX1, that arose from the whole genome duplication. (353 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) |
| | 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) |
| | SWI4 | Regulatory protein SWI4; DNA binding component of the SBF complex (Swi4p-Swi6p); a transcriptional activator that in concert with MBF (Mbp1-Swi6p) regulates late G1-specific transcription of targets including cyclins and genes required for DNA synthesis and repair; Slt2p-independent regulator of cold growth; acetylation at two sites, K1016 and K1066, regulates interaction with Swi6p. (1093 aa) |
| | SCC4 | MAU2 chromatid cohesion factor homolog; Subunit of cohesin loading factor (Scc2p-Scc4p); complex is required for the loading of cohesin complexes onto chromosomes; involved in establishing sister chromatid cohesion during double-strand break repair via phosphorylated histone H2AX. (624 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) |
| | SMC1 | Structural maintenance of chromosomes protein 1; Subunit of the multiprotein cohesin complex; essential protein involved in chromosome segregation and in double-strand DNA break repair; SMC chromosomal ATPase family member, binds DNA with a preference for DNA with secondary structure; Belongs to the SMC family. SMC1 subfamily. (1225 aa) |
| | CDC4 | Cell division control protein 4; F-box protein required for both the G1/S and G2/M phase transitions; modular substrate specificity factor which associates with core SCF (Cdc53p, Skp1p and Hrt1p/Rbx1p) to form the SCFCdc4 complex; SCFCdc4 acts as a ubiquitin-protein ligase directing ubiquitination of cyclin-dependent kinase (CDK) phosphorylated substrates, such as: Sic1p, Far1p, Cdc6p, Clb6p, and Cln3p. (779 aa) |
| | CAK1 | Serine/threonine-protein kinase CAK1; Cyclin-dependent kinase-activating kinase; required for passage through the cell cycle; phosphorylates and activates Cdc28p; nucleotide-binding pocket differs significantly from those of most other protein kinases. (368 aa) |
| | CDC14 | Tyrosine-protein phosphatase CDC14; Protein phosphatase required for mitotic exit; required for rDNA segregation, cytokinesis, meiosis I spindle disassembly, environmental stress response; held in nucleolus by Cdc55p in early meiosis, liberated by FEAR and Mitotic Exit Network in anaphase, enabling it to effect a decrease in CDK/B-cyclin activity and mitotic exit; sequestered in metaphase II, released upon entry into anaphase II; human homolog CDC14A can complement thermosensitivity of yeast cdc14-1 mutant. (551 aa) |
| | SMC2 | Structural maintenance of chromosomes protein 2; Subunit of the condensin complex; condensin reorganizes chromosomes during both mitosis and meiosis; essential SMC chromosomal ATPase family member that forms a subcomplex with Smc2p that has ATP-hydrolyzing and DNA-binding activity, but other condensin subunits are required for chromatin binding; required for clustering of tRNA genes at the nucleolus. (1170 aa) |
| | PHO4 | Phosphate system positive regulatory protein PHO4; Basic helix-loop-helix (bHLH) transcription factor of the myc-family; activates transcription cooperatively with Pho2p in response to phosphate limitation; binding to 'CACGTG' motif is regulated by chromatin restriction, competitive binding of Cbf1p to the same DNA binding motif and cooperation with Pho2p; function is regulated by phosphorylation at multiple sites and by phosphate availability. (312 aa) |
| | CDC26 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); which is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; relocalizes to the cytosol in response to hypoxia; Belongs to the CDC26 family. (124 aa) |
| | CDH1 | Activator of anaphase-promoting complex/cyclosome (APC/C); antagonist of the spindle assembly checkpoint; directs ubiquitination of cyclins resulting in mitotic exit; targets the APC/C to specific substrates including: Cdc20p, Ase1p, Cin8p, Fin1p and Clb5p; partially active in metaphase, and fully active in anaphase; cell-cycle regulated; Belongs to the WD repeat CDC20/Fizzy family. (566 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) |
| | 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) |
| | 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) |
| | MCM6 | DNA replication licensing factor MCM6; Protein involved in DNA replication; component of the Mcm2-7 hexameric helicase complex that binds chromatin as a part of the pre-replicative complex; forms a subcomplex with Mcm4p and Mcm7p. (1017 aa) |
| | DOC1 | Anaphase-promoting complex subunit DOC1; Processivity factor; required for the ubiquitination activity of the anaphase promoting complex (APC), mediates the activity of the APC by contributing to substrate recognition; involved in cyclin proteolysis; contains a conserved DOC1 homology domain; Belongs to the APC10 family. (250 aa) |
| | KSS1 | Mitogen-activated protein kinase (MAPK); involved in signal transduction pathways that control filamentous growth and pheromone response; regulates septum assembly, and may directly phosphorylate Bni4p; the KSS1 gene is nonfunctional in S288C strains and functional in W303 strains. (368 aa) |
| | DBF2 | Cell cycle protein kinase DBF2; Ser/Thr kinase involved in transcription and stress response; functions as part of a network of genes in exit from mitosis; localization is cell cycle regulated; activated by Cdc15p during the exit from mitosis; also plays a role in regulating the stability of SWI5 and CLB2 mRNAs; phosphorylates Chs2p to regulate primary septum formation and Hof1p to regulate cytokinesis; DBF2 has a paralog, DBF20, that arose from the whole genome duplication. (572 aa) |
| | ESP1 | Separin; Separase, a caspase-like cysteine protease; promotes sister chromatid separation by mediating dissociation of the cohesin Scc1p from chromatin; inhibits protein phosphatase 2A-Cdc55p to promote mitotic exit; inhibited by Pds1p; relative distribution to the nucleus increases upon DNA replication stress. (1630 aa) |
| | CLB1 | G2/mitotic-specific cyclin-1; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the transition from G2 to M phase; accumulates during G2 and M, then targeted via a destruction box motif for ubiquitin-mediated degradation by the proteasome; CLB1 has a paralog, CLB2, that arose from the whole genome duplication. (471 aa) |
| | CLB6 | S-phase entry cyclin-6; B-type cyclin involved in DNA replication during S phase; activates Cdc28p to promote initiation of DNA synthesis; functions in formation of mitotic spindles along with Clb3p and Clb4p; most abundant during late G1; CLB6 has a paralog, CLB5, that arose from the whole genome duplication. (380 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) |
| | 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) |
| | PHO81 | Phosphate system positive regulatory protein PHO81; Cyclin-dependent kinase (CDK) inhibitor; regulates Pho80p-Pho85p and Pcl7p-Pho85p cyclin-CDK complexes in response to phosphate levels; inhibitory activity for Pho80p-Pho85p requires myo-D-inositol heptakisphosphate (IP7) generated by Vip1p; relative distribution to the nucleus increases upon DNA replication stress. (1178 aa) |
| | ORC6 | Subunit of the origin recognition complex (ORC); ORC directs DNA replication by binding to replication origins and is also involved in transcriptional silencing; phosphorylated by Cdc28p; mutation in the human Orc6p is linked to Meier-Gorlin syndrome. (435 aa) |
| | SPO12 | Sporulation-specific protein 12; Nucleolar protein of unknown function; positive regulator of mitotic exit; involved in regulating release of Cdc14p from the nucleolus in early anaphase, may play similar role in meiosis; SPO12 has a paralog, BNS1, that arose from the whole genome duplication. (173 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) |
| | PHO12 | One of three repressible acid phosphatases; glycoprotein that is transported to the cell surface by the secretory pathway; pregulated by phosphate starvation; PHO12 has a paralog, PHO11, that arose from a segmental duplication. (467 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) |
| | MET30 | F-box protein containing five copies of the WD40 motif; controls cell cycle function, sulfur metabolism, and methionine biosynthesis as part of the ubiquitin ligase complex; interacts with and regulates Met4p, localizes within the nucleus; dissociation of Met30p from SCF complex in response to cadmium stress is regulated by Cdc48p. (640 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) |
| | 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) |
| | SMC3 | Structural maintenance of chromosomes protein 3; Subunit of the multiprotein cohesin complex; required for sister chromatid cohesion in mitotic cells; also required, with Rec8p, for cohesion and recombination during meiosis; phylogenetically conserved SMC chromosomal ATPase family member. (1230 aa) |
| | NET1 | Nucleolar protein NET1; Core subunit of the RENT complex; involved in nucleolar silencing and telophase exit; stimulates transcription by RNA polymerase I and regulates nucleolar structure; NET1 has a paralog, TOF2, that arose from the whole genome duplication; To yeast YKR010c. (1189 aa) |
| | FAR1 | CDK inhibitor and nuclear anchor; during the cell cycle Far1p sequesters the GEF Cdc24p in the nucleus; phosphorylation by Cdc28p-Cln results in SCFCdc4 complex-mediated ubiquitin-dependent degradation, releasing Cdc24p for export and activation of GTPase Cdc42p; in response to pheromone, phosphorylation of Far1p by MAPK Fus3p results in association with, and inhibition of Cdc28p-Cln, as well as Msn5p mediated nuclear export of Far1p-Cdc24p, targeting Cdc24p to polarity sites. (830 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) |
| | TAH11 | Cell division cycle protein CDT1; DNA replication licensing factor; required for pre-replication complex assembly; Belongs to the Cdt1 family. (604 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) |
| | GRR1 | F-box protein component of an SCF ubiquitin-ligase complex; modular substrate specificity factor which associates with core SCF (Cdc53p, Skp1p and Hrt1p/Rbx1p) to form the SCF(Grr1) complex; SCF(Grr1) acts as a ubiquitin-protein ligase directing ubiquitination of substrates such as: Gic2p, Mks1p, Mth1p, Cln1p, Cln2p and Cln3p; involved in carbon catabolite repression, glucose-dependent divalent cation transport, glucose transport, morphogenesis, and sulfite detoxification. (1151 aa) |
| | CDC16 | Subunit of the anaphase-promoting complex/cyclosome (APC/C); which is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; required for sporulation; relocalizes to the cytosol in response to hypoxia. (840 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) |
| | ORC3 | Subunit of the origin recognition complex (ORC); ORC directs DNA replication by binding to replication origins and is also involved in transcriptional silencing; Belongs to the ORC3 family. (616 aa) |
| | SIC1 | Protein SIC1; Cyclin-dependent kinase inhibitor (CKI); inhibitor of Cdc28-Clb kinase complexes that controls G1/S phase transition, preventing premature S phase and ensuring genomic integrity; phosphorylated by Clb5/6-Cdk1 and Cln1/2-Cdk1 kinase which regulate timing of Sic1p degradation; phosphorylation targets Sic1p for SCF(CDC4)-dependent turnover; functional homolog of mammalian Kip1. (284 aa) |
| | SMC4 | Structural maintenance of chromosomes protein 4; Subunit of the condensin complex; condensin reorganizes chromosomes during both mitosis and meiosis; forms a subcomplex with Smc2p that has ATP-hydrolyzing and DNA-binding activity, but other condensin subunits are required for chromatin binding; required for tRNA gene clustering at the nucleolus; potential Cdc28p substrate; Belongs to the SMC family. SMC4 subfamily. (1418 aa) |
| | APC9 | 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. (265 aa) |
| | CDC45 | Cell division control protein 45; DNA replication initiation factor; recruited to MCM pre-RC complexes at replication origins; promotes release of MCM from Mcm10p, recruits elongation machinery; binds tightly to ssDNA, which disrupts interaction with the MCM helicase and stalls it during replication stress; mutants in human homolog may cause velocardiofacial and DiGeorge syndromes. (650 aa) |
| | APC2 | Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); which is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; component of the catalytic core of the APC/C; has similarity to cullin Cdc53p. (853 aa) |
| | RFX1 | RFX-like DNA-binding protein RFX1; Major transcriptional repressor of DNA-damage-regulated genes; recruits repressors Tup1p and Cyc8p to their promoters; involved in DNA damage and replication checkpoint pathway; similar to a family of mammalian DNA binding RFX1-4 proteins. (811 aa) |
| | SWI6 | Regulatory protein SWI6; Transcription cofactor; forms complexes with Swi4p and Mbp1p to regulate transcription at the G1/S transition; involved in meiotic gene expression; also binds Stb1p to regulate transcription at START; cell wall stress induces phosphorylation by Mpk1p, which regulates Swi6p localization; required for the unfolded protein response, independently of its known transcriptional coactivators. (803 aa) |
| | CLB4 | G2/mitotic-specific cyclin-4; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the G2/M transition; may be involved in DNA replication and spindle assembly; accumulates during S phase and G2, then targeted for ubiquitin-mediated degradation; CLB4 has a paralog, CLB3, that arose from the whole genome duplication. (460 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) |
| | YOX1 | Homeobox protein YOX1; Homeobox transcriptional repressor; binds to Mcm1p and to early cell cycle boxes (ECBs) in the promoters of cell cycle-regulated genes expressed in M/G1 phase; expression is cell cycle-regulated; phosphorylated by Cdc28p; relocalizes from nucleus to cytoplasm upon DNA replication stress; YOX1 has a paralog, YHP1, that arose from the whole genome duplication. (385 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) |
| | CDC5 | Cell cycle serine/threonine-protein kinase CDC5/MSD2; Polo-like kinase; controls targeting and activation of Rho1p at cell division site via Rho1p guanine nucleotide exchange factors; regulates Spc72p; also functions in adaptation to DNA damage during meiosis; regulates the shape of the nucleus and expansion of the nuclear envelope during mitosis; similar to Xenopus Plx1 and S. pombe Plo1p; human homologs PLK1, PLK3 can each complement yeast cdc5 thermosensitive mutants; Belongs to the protein kinase superfamily. Ser/Thr protein kinase family. CDC5/Polo subfamily. (705 aa) |
| | MIH1 | M-phase inducer phosphatase; Protein tyrosine phosphatase involved in cell cycle control; regulates the phosphorylation state of Cdc28p; homolog of S. pombe cdc25; Belongs to the MPI phosphatase family. (554 aa) |
| | MCM1 | Transcription factor; involved in cell-type-specific transcription and pheromone response; plays a central role in the formation of both repressor and activator complexes; relocalizes to the cytosol in response to hypoxia. (286 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) |
| | CLN1 | G1/S-specific cyclin CLN1; G1 cyclin involved in regulation of the cell cycle; activates Cdc28p kinase to promote the G1 to S phase transition; late G1 specific expression depends on transcription factor complexes, MBF (Swi6p-Mbp1p) and SBF (Swi6p-Swi4p); CLN1 has a paralog, CLN2, that arose from the whole genome duplication; cell cycle arrest phenotype of the cln1 cln2 cln3 triple null mutant is complemented by any of human cyclins CCNA2, CCNB1, CCNC, CCND1, or CCNE1. (546 aa) |
| | APC1 | Largest subunit of the Anaphase-Promoting Complex/Cyclosome; APC/C is a ubiquitin-protein ligase required for degradation of anaphase inhibitors, including mitotic cyclins, during the metaphase/anaphase transition; component of the platform domain of the APC/C, based on structural analysis; localizes to nuclear foci that become diffuse upon DNA replication stress; Belongs to the APC1 family. (1748 aa) |
| | ORC5 | Subunit of the origin recognition complex (ORC); ORC directs DNA replication by binding to replication origins and is also involved in transcriptional silencing. (479 aa) |
| | PCL1 | PHO85 cyclin-1; Cyclin, interacts with cyclin-dependent kinase Pho85p; member of the Pcl1,2-like subfamily, involved in the regulation of polarized growth and morphogenesis and progression through the cell cycle; is ubiquitinated by Dma1p; phosphorylation by Pho85p targets it for degradation; localizes to sites of polarized cell growth. (279 aa) |
| | PHO80 | PHO85 cyclin PHO80; Cyclin; interacts with cyclin-dependent kinase Pho85p; regulates the response to nutrient levels and environmental conditions, including the response to phosphate limitation and stress-dependent calcium signaling; Belongs to the cyclin family. PHO80 subfamily. (293 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) |
| | 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) |
| | WHI5 | G1-specific transcriptional repressor WHI5; Repressor of G1 transcription; binds to SCB binding factor (SBF) at SCB target promoters in early G1; dilution of Whi5p concentration during cell growth determines cell size; phosphorylation of Whi5p by the CDK, Cln3p/Cdc28p relieves repression and promoter binding by Whi5, and contributes to both the determination of critical cell size at START and cell fate; periodically expressed in G1; Belongs to the WHI5/NRM1 family. (295 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) |
| | APC5 | 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; component of the platform domain of the APC/C, based on structural analysis; relative distribution to nuclear foci decreases upon DNA replication stress; Belongs to the APC5 family. (685 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) |
| | PHO85 | Cyclin-dependent kinase; has ten cyclin partners; involved in regulating the cellular response to nutrient levels and environmental conditions and progression through the cell cycle; human lissencephaly-associated homolog CDK5 functionally complements null mutation; Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. CDC2/CDKX subfamily. (305 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) |
| | CLN2 | G1/S-specific cyclin CLN2; G1 cyclin involved in regulation of the cell cycle; activates Cdc28p kinase to promote the G1 to S phase transition; late G1 specific expression depends on transcription factor complexes, MBF (Swi6p-Mbp1p) and SBF (Swi6p-Swi4p); CLN2 has a paralog, CLN1, that arose from the whole genome duplication; cell cycle arrest phenotype of the cln1 cln2 cln3 triple null mutant is complemented by any of human cyclins CCNA2, CCNB1, CCNC, CCND1, or CCNE1. (545 aa) |
| | MCM4 | DNA replication licensing factor MCM4; Essential helicase component of heterohexameric MCM2-7 complexes; MCM2-7 complexes bind pre-replication complexes on DNA and melt DNA prior to replication; forms an Mcm4p-6p-7p subcomplex; shows nuclear accumulation in G1; homolog of S. pombe Cdc21p. (933 aa) |
| | DBF20 | Serine/threonine-protein kinase DBF20; Ser/Thr kinase involved in late nuclear division; one of the mitotic exit network (MEN) proteins; necessary for the execution of cytokinesis; also plays a role in regulating the stability of SWI5 and CLB2 mRNAs; DBF20 has a paralog, DBF2, that arose from the whole genome duplication. (564 aa) |
| | CLB2 | G2/mitotic-specific cyclin-2; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the transition from G2 to M phase; accumulates during G2 and M, then targeted via a destruction box motif for ubiquitin-mediated degradation by the proteasome; CLB2 has a paralog, CLB1, that arose from the whole genome duplication. (491 aa) |
| | CLB5 | S-phase entry cyclin-5; B-type cyclin involved in DNA replication during S phase; activates Cdc28p to promote initiation of DNA synthesis; functions in formation of mitotic spindles along with Clb3p and Clb4p; most abundant during late G1 phase; CLB5 has a paralog, CLB6, that arose from the whole genome duplication. (435 aa) |
| | ORC4 | Subunit of the origin recognition complex (ORC); ORC directs DNA replication by binding to replication origins and is also involved in transcriptional silencing; ORC4 has a paralog, RIF2, that arose from the whole genome duplication. (529 aa) |
