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RFA1 RFA1 RDH54 RDH54 MMS4 MMS4 IML3 IML3 MEC1 MEC1 CDC28 CDC28 PCH2 PCH2 CSM1 CSM1 CDC7 CDC7 MSH5 MSH5 CLB3 CLB3 MHF2 MHF2 HED1 HED1 RAD55 RAD55 PDS1 PDS1 NUM1 NUM1 CHL4 CHL4 ZIP1 ZIP1 MCM21 MCM21 YCG1 YCG1 MUS81 MUS81 LRS4 LRS4 ECM11 ECM11 CIN8 CIN8 RAD51 RAD51 MAM1 MAM1 RAD24 RAD24 DMC1 DMC1 MSH4 MSH4 TUB2 TUB2 ECO1 ECO1 CDC14 CDC14 SMC2 SMC2 HOP2 HOP2 AFT1 AFT1 MPS2 MPS2 RAD54 RAD54 MND1 MND1 CDC55 CDC55 SKI8 SKI8 KIP3 KIP3 ZIP2 ZIP2 RMR1 RMR1 HFM1 HFM1 ESP1 ESP1 BUB1 BUB1 AMA1 AMA1 SPO11 SPO11 RIM4 RIM4 SPO13 SPO13 MSC7 MSC7 SAE3 SAE3 SPO12 SPO12 SPO16 SPO16 REC104 REC104 HOP1 HOP1 CSM2 CSM2 DSN1 DSN1 MND2 MND2 MAD3 MAD3 MPS3 MPS3 SMC3 SMC3 SCP160 SCP160 RFA3 RFA3 REC107 REC107 MIF2 MIF2 MLH2 MLH2 SMC4 SMC4 SLX4 SLX4 MSC3 MSC3 TOP3 TOP3 RED1 RED1 YCS4 YCS4 MMS22 MMS22 REC102 REC102 CST9 CST9 GMC2 GMC2 RAD52 RAD52 TUB1 TUB1 TUB3 TUB3 MSC1 MSC1 CDC5 CDC5 CSM3 CSM3 PDS5 PDS5 REC114 REC114 MLH1 MLH1 SGS1 SGS1 MRE11 MRE11 TOP2 TOP2 MER1 MER1 RAD50 RAD50 RFA2 RFA2 NUF2 NUF2 MHF1 MHF1 NDJ1 NDJ1 RTS1 RTS1 SGO1 SGO1 MSC6 MSC6 RAD17 RAD17 NUD1 NUD1 IRC15 IRC15 RAD1 RAD1 MEI5 MEI5 MLH3 MLH3 CSM4 CSM4 HRR25 HRR25 IPL1 IPL1 REC8 REC8
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RFA1Subunit 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)
RDH54DNA-dependent ATPase; DNA recombination/repair translocase, supercoils DNA and promotes DNA strand opening; stimulates strand exchange by modifying dsDNA topology; involved in recombinational repair of DNA double-strand breaks (DSBs) during mitosis and meiosis; phosphorylated in Mec1p-, Rad53p-dependent way in response to one DSB; contributes to remodelling of nucleosomes; proposed to be involved in crossover interference; interacts with Dmc1p; stimulates Dmc1p and Rad51p. (958 aa)
MMS4Subunit 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)
IML3Inner 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)
MEC1Serine/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)
CDC28Cyclin-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)
PCH2Pachytene 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)
CSM1Monopolin 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)
CDC7Cell 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)
MSH5Protein of the MutS family; forms a dimer with Msh4p that facilitates crossovers between homologs during meiosis; msh5-Y823H mutation confers tolerance to DNA alkylating agents; homologs present in C. elegans and humans. (901 aa)
CLB3G2/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)
MHF2Inner kinetochore subunit MHF2; Component of the heterotetrameric MHF histone-fold complex; in humans the MHF complex interacts with both DNA and Mph1p ortholog FANCM to stabilize and remodel blocked replication forks and repair damaged DNA; mhf2 srs2 double mutants are MMS hypersensitive; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-X, also known as MHF2. (80 aa)
HED1Meiosis-specific protein; down-regulates Rad51p-mediated mitotic recombination when the meiotic recombination machinery is impaired; promotes synapsis and required for the normal morphogenesis of synaptonemal complex; prevents the recruitment of Rad54p to site-specific DNA double-strand breaks in vivo; early meiotic gene, transcribed specifically during meiotic prophase. (162 aa)
RAD55DNA repair protein RAD55; Protein that stimulates strand exchange; stimulates strand exchange by stabilizing the binding of Rad51p to single-stranded DNA; involved in the recombinational repair of double-strand breaks in DNA during vegetative growth and meiosis; forms heterodimer with Rad57p; Belongs to the RecA family. RAD55 subfamily. (406 aa)
PDS1Securin; 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)
NUM1Protein required for nuclear migration; component of the mitochondria-ER-cortex-ancor (MECA); required for the association of mitochondria with the cell cortex and for accurate distribution of mitochondrial network; interacts with Mdm36p to link the ER and mitochondria at the cortex; localizes to the mother cell cortex and the bud tip; may mediate interactions of dynein and cytoplasmic microtubules with the cell cortex. (2748 aa)
CHL4Inner 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)
ZIP1Transverse 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)
MCM21Inner 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)
YCG1Subunit 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)
MUS81Subunit 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)
LRS4Monopolin 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)
ECM11Meiosis-specific protein; component of the Synaptonemal Complex (SC) along with Gmc2p; required for efficient crossover formation and for the efficient loading of the SC transverse filament protein, Zip1p; is SUMOlytaed in a Gmc2p manner, and SUMOylation is required for its function in meiosis; GFP fusion protein is present in discrete clusters in the nucleus throughout mitosis; may be involved in maintaining chromatin structure. (302 aa)
CIN8Kinesin-like protein CIN8; Kinesin motor protein; involved in mitotic spindle assembly and chromosome segregation. (1000 aa)
RAD51DNA repair protein RAD51; Strand exchange protein; forms a helical filament with DNA that searches for homology; involved in the recombinational repair of double-strand breaks in DNA during vegetative growth and meiosis; homolog of Dmc1p and bacterial RecA protein. (400 aa)
MAM1Monopolin 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)
RAD24Checkpoint 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)
DMC1Meiotic recombination protein DMC1; Meiosis-specific recombinase required for double-strand break repair; also required for pairing between homologous chromosomes; required for the normal morphogenesis of synaptonemal complex; homolog of Rad51p and the bacterial RecA protein; binds ssDNA and dsDNA, forms helical filaments; stimulated by Rdh54p. (334 aa)
MSH4MutS protein homolog 4; Protein involved in meiotic recombination; required for normal levels of crossing over, colocalizes with Zip2p to discrete foci on meiotic chromosomes, has homology to bacterial MutS protein; Belongs to the DNA mismatch repair MutS family. (878 aa)
TUB2Beta-tubulin; associates with alpha-tubulin (Tub1p and Tub3p) to form tubulin dimer, which polymerizes to form microtubules; mutation in human ortholog is associated with congenital fibrosis of the extraocular muscles (CFEOM) with polymicrogyria. (457 aa)
ECO1N-acetyltransferase ECO1; Acetyltransferase; required for establishment of sister chromatid cohesion; acetylates Mps3p to regulate nuclear organization; modifies Smc3p at replication forks and Mcd1p in response to dsDNA breaks; phosphorylated by three kinases (Cdc28p, Cdc7p, Mck1p) to generate pair of phosphates spaced precisely for recognition by ubiquitin ligase SCF-Cdc4; mutations in human homolog ESCO2 cause Roberts syndrome; relative distribution to nucleus increases upon DNA replication stress. (281 aa)
CDC14Tyrosine-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)
SMC2Structural 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)
HOP2Homologous-pairing protein 2; Meiosis-specific protein that localizes to chromosomes; prevents synapsis between nonhomologous chromosomes and ensures synapsis between homologs; complexes with Mnd1p to promote homolog pairing and meiotic double-strand break repair; heterodimer of Hop2p-Mnd1p stimulates the Dmc1p-mediated strand invasion. (214 aa)
AFT1Iron-regulated transcriptional activator AFT1; Transcription factor involved in iron utilization and homeostasis; binds consensus site PyPuCACCCPu and activates transcription in response to changes in iron availability; in iron-replete conditions localization is regulated by Grx3p, Grx4p, and Fra2p, and promoter binding is negatively regulated via Grx3p-Grx4p binding; AFT1 has a paralog, AFT2, that arose from the whole genome duplication; relative distribution to the nucleus increases upon DNA replication stress. (690 aa)
MPS2Monopolar 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)
RAD54DNA repair and recombination protein RAD54; DNA-dependent ATPase that stimulates strand exchange; modifies the topology of double-stranded DNA; involved in the recombinational repair of double-strand breaks in DNA during vegetative growth and meiosis; member of the SWI/SNF family of DNA translocases; forms nuclear foci upon DNA replication stress. (898 aa)
MND1Protein required for recombination and meiotic nuclear division; forms a complex with Hop2p, which is involved in chromosome pairing and repair of meiotic double-strand breaks. (219 aa)
CDC55Regulatory 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)
SKI8Antiviral protein SKI8; Ski complex component and WD-repeat protein; mediates 3'-5' RNA degradation by the cytoplasmic exosome; also required for meiotic double-strand break recombination; null mutants have superkiller phenotype. (397 aa)
KIP3Kinesin-like protein KIP3; Kinesin-related antiparallel sliding motor protein; involved in mitotic spindle positioning; sliding activity promotes bipolar spindle assembly and maintenance of genome stability; inhibits spindle elongation, destabilizing late anaphase spindle microtubules that polymerize beyond the midzone; Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family. Kinesin II subfamily. (805 aa)
ZIP2Meiosis-specific protein; involved in normal synaptonemal complex formation and pairing between homologous chromosomes during meiosis; relocalizes from mitochondrion to cytoplasm upon DNA replication stress; Belongs to the ZIP2 family. (704 aa)
RMR1Protein required for meiotic recombination and gene conversion; null mutant displays reduced PIS1 expression and growth defects on non-fermentable carbon sources and minimal media; GFP-fusion protein localizes to both cytoplasm and nucleus; Belongs to the RMR1 family. (241 aa)
HFM1Meiosis specific DNA helicase; involved in the conversion of double-stranded breaks to later recombination intermediates and in crossover control; catalyzes the unwinding of Holliday junctions; has ssDNA and dsDNA stimulated ATPase activity. (1187 aa)
ESP1Separin; 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)
BUB1Protein 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)
AMA1Meiosis-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)
SPO11Meiosis-specific protein that initiates meiotic recombination; initiates meiotic recombination by catalyzing the formation of double-strand breaks in DNA via a transesterification reaction; required for homologous chromosome pairing and synaptonemal complex formation; Belongs to the TOP6A family. (398 aa)
RIM4Meiotic activator RIM4; Putative RNA-binding protein; required for the expression of early and middle sporulation genes. (713 aa)
SPO13Meiosis-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)
MSC7Putative aldehyde dehydrogenase-like protein YHR039C; Protein of unknown function; green fluorescent protein (GFP)-fusion protein localizes to the endoplasmic reticulum; msc7 mutants are defective in directing meiotic recombination events to homologous chromatids. (644 aa)
SAE3Pachytene arrest protein SAE3; Meiosis-specific protein involved in meiotic recombination; involved in DMC1-dependent meiotic recombination; forms heterodimer with Mei5p; proposed to be an assembly factor for Dmc1p. (91 aa)
SPO12Sporulation-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)
SPO16Sporulation-specific protein 16; Meiosis-specific protein involved in synaptonemal complex assembly; implicated in regulation of crossover formation; required for sporulation. (198 aa)
REC104Protein involved in early stages of meiotic recombination; required for meiotic crossing over; forms a complex with Rec102p and Spo11p necessary during the initiation of recombination. (182 aa)
HOP1Meiosis-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)
CSM2Chromosome segregation in meiosis protein 2; Component of Shu complex (aka PCSS complex); Shu complex also includes Psy3, Shu1, Shu2, and promotes error-free DNA repair,; Shu complex mediates inhibition of Srs2p function; promotes formation of Rad51p filaments; Psy3p and Csm2p contain similar DNA-binding regions which work together to form a single DNA binding site; required for accurate chromosome segregation during meiosis. (213 aa)
DSN1Kinetochore-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)
MND2Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); necessary for maintaining sister chromatid cohesion in prophase I of meiosis by inhibiting premature ubiquitination and subsequent degradation of substrates by the APC(Ama1) ubiquitin ligase; Belongs to the APC15 family. (368 aa)
MAD3Subunit 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)
MPS3Spindle 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)
SMC3Structural 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)
SCP160Protein 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)
RFA3Subunit 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)
REC107Recombination protein 107; Protein involved in early stages of meiotic recombination; involved in coordination between the initiation of recombination and the first division of meiosis; part of a complex (Rec107p-Mei4p-Rec114p) required for ds break formation. (314 aa)
MIF2Inner 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)
MLH2Protein involved in mismatch repair and meiotic recombination; only certain frameshift intermediates are mismatch repair substrates; forms a complex with Mlh1p. (695 aa)
SMC4Structural 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)
SLX4Structure-specific endonuclease subunit SLX4; Endonuclease involved in processing DNA; acts during recombination, repair; promotes template switching during break-induced replication (BIR), causing non-reciprocal translocations (NRTs); cleaves branched structures in complex with Slx1p; involved in interstrand cross-link repair, Rad1p/Rad10p-dependent removal of 3'-nonhomologous tails during DSBR via single-strand annealing; relative distribution to nuclear foci increases upon DNA replication stress; FANCP-related factor; Belongs to the SLX4 family. (748 aa)
MSC3Meiotic sister-chromatid recombination protein 3; Protein of unknown function; green fluorescent protein (GFP)-fusion protein localizes to the cell periphery; msc3 mutants are defective in directing meiotic recombination events to homologous chromatids; potential Cdc28p substrate; protein abundance increases in response to DNA replication stress. (728 aa)
TOP3DNA Topoisomerase III; conserved protein that functions in a complex with Sgs1p and Rmi1p to relax single-stranded negatively-supercoiled DNA preferentially; DNA catenation/decatenation activity is stimulated by RPA and Sgs1p-Top3p-Rmi1p; involved in telomere stability and regulation of mitotic recombination. (656 aa)
RED1Protein 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)
YCS4Subunit 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)
MMS22E3 ubiquitin-protein ligase substrate receptor MMS22; Subunit of E3 ubiquitin ligase complex involved in replication repair; stabilizes protein components of the replication fork, such as the fork-pausing complex and leading strand polymerase, preventing fork collapse and promoting efficient recovery during replication stress; required for accurate meiotic chromosome segregation; Belongs to the MMS22 family. (1454 aa)
REC102Protein involved in early stages of meiotic recombination; required for chromosome synapsis; forms a complex with Rec104p and Spo11p necessary during the initiation of recombination. (264 aa)
CST9Chromosome 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)
GMC2Grand meiotic recombination cluster protein 2; Protein involved in meiotic crossing over; component of the Synaptonemal Complex (SC) along with Ecm11p; required for the efficient loading of the SC transverse filament protein, Zip1p; promotes SUMOylation of Ecm11p; mutants are delayed in meiotic nuclear division and are defective in synaptonemal complex assembly; transcription is regulated by Ume6p and induced in response to alpha factor. (188 aa)
RAD52DNA repair and recombination protein RAD52; Protein that stimulates strand exchange; stimulates strand exchange by facilitating Rad51p binding to single-stranded DNA; anneals complementary single-stranded DNA; involved in the repair of double-strand breaks in DNA during vegetative growth and meiosis and UV induced sister chromatid recombination; Belongs to the RAD52 family. (471 aa)
TUB1Tubulin alpha-1 chain; Alpha-tubulin; associates with beta-tubulin (Tub2p) to form tubulin dimer, which polymerizes to form microtubules; relative distribution to nuclear foci increases upon DNA replication stress; TUB1 has a paralog, TUB3, that arose from the whole genome duplication. (447 aa)
TUB3Tubulin alpha-3 chain; Alpha-tubulin; associates with beta-tubulin (Tub2p) to form tubulin dimer, which polymerizes to form microtubules; expressed at lower level than Tub1p; TUB3 has a paralog, TUB1, that arose from the whole genome duplication. (445 aa)
MSC1Protein of unknown function; mutant is defective in directing meiotic recombination events to homologous chromatids; the authentic, non-tagged protein is detected in highly purified mitochondria and is phosphorylated. (513 aa)
CDC5Cell 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)
CSM3Chromosome 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)
PDS5Cohesion maintenance factor; involved in sister chromatid condensation and cohesion; colocalizes with cohesin on chromosomes; performs its cohesin maintenance function in pre-anaphase cells by protecting the integrity of the cohesion complex; also required during meiosis; relocalizes to the cytosol in response to hypoxia. (1277 aa)
REC114Protein involved in early stages of meiotic recombination; possibly involved in the coordination of recombination and meiotic division; mutations lead to premature initiation of the first meiotic division. (428 aa)
MLH1Protein required for mismatch repair in mitosis and meiosis; also required for crossing over during meiosis; forms a complex with Pms1p and Msh2p-Msh3p during mismatch repair; human homolog is associated with hereditary non-polyposis colon cancer; Belongs to the DNA mismatch repair MutL/HexB family. (769 aa)
SGS1ATP-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)
MRE11Double-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)
TOP2DNA topoisomerase 2; Topoisomerase II; relieves torsional strain in DNA by cleaving and re-sealing phosphodiester backbone of both positively and negatively supercoiled DNA; cleaves complementary strands; localizes to axial cores in meiosis; required for replication slow zone (RSZ) breakage following Mec1p inactivation; human homolog TOP2A implicated in cancers, and can complement yeast null mutant; Belongs to the type II topoisomerase family. (1428 aa)
MER1Meiotic recombination 1 protein; mRNA-binding protein required for meiosis-specific mRNA splicing; required for chromosome pairing and meiotic recombination; Mer1p regulon embraces four essential meiotic pre-mRNAs: REC107, HFM1, AMA1 and SPO22. (270 aa)
RAD50DNA repair protein RAD50; Subunit of MRX complex with Mre11p and Xrs2p; complex is involved in processing double-strand DNA breaks in vegetative cells, initiation of meiotic DSBs, telomere maintenance, and nonhomologous end joining; forms nuclear foci upon DNA replication stress; Belongs to the SMC family. RAD50 subfamily. (1312 aa)
RFA2Subunit 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)
NUF2Kinetochore 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)
MHF1Inner kinetochore subunit MHF1; Component of the heterotetrameric MHF histone-fold complex; in humans the MHF complex interacts with both DNA and Mph1p ortholog FANCM, a Fanconi anemia complementation group protein, to stabilize and remodel blocked replication forks and repair damaged DNA; mhf1 srs2 double mutants are MMS hypersensitive; ortholog of human centromere constitutive-associated network (CCAN) subunit CENP-S, also known as MHF1. (90 aa)
NDJ1Non-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)
RTS1Serine/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)
SGO1Shugoshin; 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)
MSC6Multicopy suppressor of HER2 involved in mitochondrial translation; mutant is defective in directing meiotic recombination events to homologous chromatids. (692 aa)
RAD17Checkpoint 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)
NUD1Protein NUD1; Component of the spindle pole body outer plaque; acts through the mitotic exit network to specify asymmetric spindle pole body inheritance. (851 aa)
IRC15Increased recombination centers protein 15; Microtubule associated protein; regulates microtubule dynamics; required for accurate meiotic chromosome segregation; null mutant displays large budded cells due to delayed mitotic progression, increased levels of spontaneous Rad52 foci; IRC15 has a paralog, LPD1, that arose from the whole genome duplication; Belongs to the class-I pyridine nucleotide-disulfide oxidoreductase family. (499 aa)
RAD1Single-stranded DNA endonuclease (with Rad10p); cleaves single-stranded DNA during nucleotide excision repair and double-strand break repair; subunit of Nucleotide Excision Repair Factor 1 (NEF1); homolog of human XPF protein. (1100 aa)
MEI5Meiosis-specific protein involved in meiotic recombination; involved in DMC1-dependent meiotic recombination; forms heterodimer with Sae3p; proposed to be an assembly factor for Dmc1p; Belongs to the SFR1/MEI5 family. (222 aa)
MLH3Protein involved in DNA mismatch repair and meiotic recombination; involved in crossing-over during meiotic recombination; forms a complex with Mlh1p; mammalian homolog is implicated mammalian microsatellite instability; Belongs to the DNA mismatch repair MutL/HexB family. (715 aa)
CSM4Protein 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)
HRR25Conserved 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)
IPL1Spindle 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)
REC8Meiotic 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)
Your Current Organism:
Saccharomyces cerevisiae
NCBI taxonomy Id: 4932
Other names: ATCC 18824, Candida robusta, Mycoderma cerevisiae, NRRL Y-12632, S. cerevisiae, Saccharomyces capensis, Saccharomyces italicus, Saccharomyces oviformis, Saccharomyces uvarum var. melibiosus, yeast
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