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| RFA1 | Subunit of heterotrimeric Replication Protein A (RPA); RPA is a highly conserved single-stranded DNA binding protein involved in DNA replication, repair, and recombination; RPA protects against inappropriate telomere recombination, and upon telomere uncapping, prevents cell proliferation by a checkpoint-independent pathway; role in DNA catenation/decatenation pathway of chromosome disentangling; relocalizes to the cytosol in response to hypoxia. (621 aa) | ||||
| RDH54 | DNA-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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| MSH5 | Protein 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) | ||||
| MHF2 | Inner 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) | ||||
| HO | Homothallic switching endonuclease; Site-specific endonuclease; required for gene conversion at the MAT locus (homothallic switching) through the generation of a ds DNA break; expression restricted to mother cells in late G1 as controlled by Swi4p-Swi6p, Swi5p, and Ash1p. (586 aa) | ||||
| RAD55 | DNA 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) | ||||
| ZIP1 | Transverse filament protein of the synaptonemal complex; required for normal levels of meiotic recombination and pairing between homologous chromosome during meiosis; required for meiotic recombination between non-allelc sites; potential Cdc28p substrate. (875 aa) | ||||
| MCM21 | Inner kinetochore subunit MCM21; Component of the kinetochore sub-complex COMA; COMA (Ctf19p, Okp1p, Mcm21p, Ame1p) bridges kinetochore subunits in contact with centromeric DNA with subunits bound to microtubules during kinetochore assembly; involved in minichromosome maintenance; modified by sumoylation; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-O and fission yeast mal2; Belongs to the CENP-O/MCM21 family. (368 aa) | ||||
| 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) | ||||
| ECM11 | Meiosis-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) | ||||
| RAD51 | DNA 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) | ||||
| 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) | ||||
| DMC1 | Meiotic 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) | ||||
| MSH4 | MutS 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) | ||||
| HOP2 | Homologous-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) | ||||
| SOH1 | Subunit of the RNA polymerase II mediator complex; associates with core polymerase subunits to form the RNA polymerase II holoenzyme; involved in telomere maintenance; conserved with other metazoan MED31 subunits. (127 aa) | ||||
| RAD54 | DNA 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) | ||||
| HUR1 | Putative uncharacterized protein HUR1; Protein of unknown function; reported null mutant phenotype of hydroxyurea sensitivity may be due to effects on overlapping PMR1 gene. (110 aa) | ||||
| SAE2 | DNA endonuclease SAE2; Endonuclease required for telomere elongation; required for telomeric 5' C-rich strand resection; involved in ds-break repair and processing hairpin DNA structures with the MRX complex; function requires sumoylation and phosphorylation; exists as inactive oligomers that are transiently released into smaller active units by phosphorylation; DNA damage triggers Sae2p removal, so active Sae2p is present only transiently; sequence and functional similarity with human CtIP/RBBP8; Belongs to the COM1/SAE2/CtIP family. (345 aa) | ||||
| MND1 | Protein 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) | ||||
| SKI8 | Antiviral 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) | ||||
| ZIP2 | Meiosis-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) | ||||
| RMR1 | Protein 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) | ||||
| HFM1 | Meiosis 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) | ||||
| SPO11 | Meiosis-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) | ||||
| RIM4 | Meiotic activator RIM4; Putative RNA-binding protein; required for the expression of early and middle sporulation genes. (713 aa) | ||||
| MSC7 | Putative 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) | ||||
| SAE3 | Pachytene 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) | ||||
| REC104 | Protein 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) | ||||
| MND2 | Subunit 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) | ||||
| 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) | ||||
| RFA3 | Subunit of heterotrimeric Replication Protein A (RPA); RPA is a highly conserved single-stranded DNA binding protein complex involved in DNA replication, repair, recombination; RPA protects against inappropriate telomere recombination, and upon telomere uncapping, prevents cell proliferation by a checkpoint-independent pathway; with Sgs1p-Top2p-Rmi1p, stimulates DNA catenation/decatenation activity of Top3p; protein abundance increases in response to DNA replication stress. (122 aa) | ||||
| REC107 | Recombination 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) | ||||
| RAD27 | Flap endonuclease 1; 5' to 3' exonuclease, 5' flap endonuclease; required for Okazaki fragment processing and maturation, for long-patch base-excision repair and large loop repair (LLR), ribonucleotide excision repair; member of the S. pombe RAD2/FEN1 family; relocalizes to the cytosol in response to hypoxia. (382 aa) | ||||
| MLH2 | Protein involved in mismatch repair and meiotic recombination; only certain frameshift intermediates are mismatch repair substrates; forms a complex with Mlh1p. (695 aa) | ||||
| SLX4 | Structure-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) | ||||
| MSC3 | Meiotic 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) | ||||
| TOP3 | DNA 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) | ||||
| 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) | ||||
| NEJ1 | Non-homologous end-joining protein 1; Protein involved in regulation of nonhomologous end joining; interacts with DNA ligase IV components Dnl4p and Lif1p; repressed by MAT heterozygosity; regulates cellular distribution of Lif1p. (342 aa) | ||||
| REC102 | Protein 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) | ||||
| CST9 | Chromosome stability protein 9; SUMO E3 ligase; required for synaptonemal complex formation; localizes to synapsis initiation sites on meiotic chromosomes; associates with centromeres early in meiosis, then with chromosome axes and finally with double-strand break sites that are engaged in repair by crossovers; potential Cdc28p substrate. (482 aa) | ||||
| GMC2 | Grand 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) | ||||
| RAD52 | DNA 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) | ||||
| MSC1 | Protein 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) | ||||
| 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) | ||||
| REC114 | Protein 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) | ||||
| MLH1 | Protein 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) | ||||
| 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) | ||||
| TOP2 | DNA 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) | ||||
| MER1 | Meiotic 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) | ||||
| 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) | ||||
| RFA2 | Subunit of heterotrimeric Replication Protein A (RPA); RPA is a highly conserved single-stranded DNA binding protein involved in DNA replication, repair, and recombination; RPA protects against inappropriate telomere recombination, and upon telomere uncapping, prevents cell proliferation by a checkpoint-independent pathway; in concert with Sgs1p-Top2p-Rmi1p, stimulates DNA catenation/decatenation activity of Top3p; protein abundance increases in response to DNA replication s. (273 aa) | ||||
| MHF1 | Inner 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) | ||||
| MSH2 | Protein that binds to DNA mismatches; forms heterodimers with Msh3p and Msh6p that bind to DNA mismatches to initiate the mismatch repair process; contains a Walker ATP-binding motif required for repair activity and involved in interstrand cross-link repair; Msh2p-Msh6p binds to and hydrolyzes ATP. (964 aa) | ||||
| NDJ1 | Non-disjunction protein 1; Protein that regulates meiotic SPB cohesion and telomere clustering; localizes to both spindle pole bodies (SPBs) and telomeres; required for bouquet formation, effective homolog pairing, ordered cross-over distribution, sister chromatid cohesion at meiotic telomeres, chromosomal segregation and telomere-led rapid prophase movement. (352 aa) | ||||
| 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) | ||||
| MSC6 | Multicopy suppressor of HER2 involved in mitochondrial translation; mutant is defective in directing meiotic recombination events to homologous chromatids. (692 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) | ||||
| RAD1 | Single-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) | ||||
| MEI5 | Meiosis-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) | ||||
| MLH3 | Protein 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) | ||||
| REC8 | Meiotic recombination protein REC8; Meiosis-specific component of the sister chromatid cohesion complex; alpha-kleisin family member that maintains cohesion between sister chromatids during meiosis I; maintains cohesion between centromeres of sister chromatids until meiosis II; independent of its role in sister chromatid cohesion, Rec8p promotes allelic collisions and prevents nonspecific chromosome interactions; homolog of S. pombe Rec8p. (680 aa) | ||||