STRINGSTRING
POL30 POL30 MMS4 MMS4 PRI2 PRI2 IXR1 IXR1 CCE1 CCE1 MGM101 MGM101 RFC2 RFC2 POL32 POL32 RAD26 RAD26 POL31 POL31 SWE1 SWE1 RFA3 RFA3 ASF1 ASF1 GSH1 GSH1 SRS2 SRS2 DPB11 DPB11 MAD2 MAD2 RNR2 RNR2 MPS3 MPS3 MAD3 MAD3 MPH1 MPH1 MCM10 MCM10 REV7 REV7 MET18 MET18 RNR3 RNR3 ULP2 ULP2 DNA2 DNA2 MSH1 MSH1 YHR097C YHR097C RRM3 RRM3 SOD2 SOD2 RAD2 RAD2 TRX2 TRX2 BUB1 BUB1 RNR4 RNR4 ESP1 ESP1 COX4 COX4 SAE2 SAE2 RAD54 RAD54 LIF1 LIF1 MMS2 MMS2 MAD1 MAD1 AFT1 AFT1 RAD6 RAD6 ECO1 ECO1 DMC1 DMC1 RAD3 RAD3 BEM2 BEM2 RAD51 RAD51 RNR1 RNR1 YEN1 YEN1 CAN1 CAN1 URA3 URA3 MMS21 MMS21 RAD30 RAD30 SIZ1 SIZ1 MUS81 MUS81 RSC3 RSC3 MHR1 MHR1 DIN7 DIN7 RAD9 RAD9 DPB4 DPB4 PDS1 PDS1 FOB1 FOB1 MSH6 MSH6 UBC13 UBC13 RLI1 RLI1 RAD55 RAD55 PPH3 PPH3 RAD57 RAD57 NTH1 NTH1 PTP1 PTP1 CDC9 CDC9 CYK3 CYK3 POL3 POL3 DUN1 DUN1 UBC9 UBC9 RAD59 RAD59 MBP1 MBP1 SIR2 SIR2 MSH3 MSH3 RAD18 RAD18 MRC1 MRC1 DPB3 DPB3 SLX1 SLX1 CDC28 CDC28 MEC1 MEC1 IML3 IML3 Q0092 Q0092 NTG1 NTG1 RFA1 RFA1 APN2 APN2 MCM2 MCM2 POL12 POL12 TEL1 TEL1 ATP1 ATP1 HHF1 HHF1 HHT1 HHT1 CHS2 CHS2 ORC2 ORC2 RDH54 RDH54 HDA3 HDA3 DPB2 DPB2 TAZ1 TAZ1 CTF4 CTF4 CLB5 CLB5 HOS1 HOS1 TAH18 TAH18 YME1 YME1 HSP82 HSP82 IPL1 IPL1 DDC1 DDC1 REV3 REV3 MLH3 MLH3 RAD53 RAD53 HOS3 HOS3 GLR1 GLR1 GRX5 GRX5 RMI1 RMI1 RAD1 RAD1 ULP1 ULP1 RAD17 RAD17 REV1 REV1 HNT3 HNT3 WTM1 WTM1 RFC1 RFC1 ELG1 ELG1 CAT5 CAT5 RAS1 RAS1 SGO1 SGO1 EXO1 EXO1 BUB3 BUB3 DNL4 DNL4 HMI1 HMI1 MSH2 MSH2 NTG2 NTG2 TOP1 TOP1 SIN3 SIN3 NRM1 NRM1 RFA2 RFA2 STB1 STB1 TOF1 TOF1 POL2 POL2 ORC5 ORC5 RAD50 RAD50 NAR1 NAR1 RTT106 RTT106 PSY2 PSY2 UBP10 UBP10 RPS3 RPS3 INN1 INN1 POL1 POL1 TOP2 TOP2 PMS1 PMS1 HHT2 HHT2 HHF2 HHF2 YKU70 YKU70 ZDS1 ZDS1 SGS1 SGS1 HSC82 HSC82 MLH1 MLH1 SAS2 SAS2 YKU80 YKU80 CTF18 CTF18 ABF2 ABF2 CSM3 CSM3 CDC5 CDC5 RAD10 RAD10 OGG1 OGG1 SML1 SML1 RAD52 RAD52 UNG1 UNG1 DIF1 DIF1 ILV5 ILV5 ACO1 ACO1 NEJ1 NEJ1 SWI6 SWI6 SLX4 SLX4 HOG1 HOG1 CDC45 CDC45 TRX1 TRX1 RAD5 RAD5 DRE2 DRE2 APN1 APN1 RAD27 RAD27 DEF1 DEF1 CSE4 CSE4
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POL30Proliferating cell nuclear antigen (PCNA); functions as the sliding replication clamp for DNA polymerase delta; may function as a docking site for other proteins required for mitotic and meiotic chromosomal DNA replication and for DNA repair; PCNA ubiquitination at K164 plays a crucial role during Okazaki fragment processing. (258 aa)
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)
PRI2Subunit of DNA primase; DNA primase is required for DNA synthesis and double-strand break repair; Belongs to the eukaryotic-type primase large subunit family. (528 aa)
IXR1Intrastrand cross-link recognition protein; Transcriptional repressor that regulates hypoxic genes during normoxia; involved in the aerobic repression of genes such as COX5b, TIR1, and HEM13; binds DNA intrastrand cross-links formed by cisplatin; HMG (high mobility group box) domain containing protein which binds and bends cisplatin-modified DNA, blocking excision repair; IXR1 has a paralog, ABF2, that arose from the whole genome duplication. (597 aa)
CCE1Mitochondrial cruciform cutting endonuclease; cleaves Holliday junctions formed during recombination of mitochondrial DNA; CCE1 has a paralog, MRS1, that arose from the whole genome duplication. (353 aa)
MGM101Mitochondrial genome maintenance protein MGM101; Protein with a role in mitochondrial DNA recombinational repair; also involved in interstrand cross-link repair; binds to and catalyzes the annealing of single-stranded mtDNA; oligomerizes to form rings and filaments; related to Rad52-type recombination proteins, with limited overall similarity but sharing conserved functionally important residues; component of the mitochondrial nucleoid, required for the repair of oxidative mtDNA damage and mitochondrial genome maintenance; Belongs to the MGM101 family. (269 aa)
RFC2Subunit of heteropentameric Replication factor C (RF-C); RF-C is a DNA binding protein and ATPase that acts as a clamp loader of the proliferating cell nuclear antigen (PCNA) processivity factor for DNA polymerases delta and epsilon. (353 aa)
POL32Third subunit of DNA polymerase delta; involved in chromosomal DNA replication; required for error-prone DNA synthesis in the presence of DNA damage and processivity; forms a complex with Rev3p, Rev7p and Pol31p; interacts with Hys2p, PCNA (Pol30p), and Pol1p. (350 aa)
RAD26DNA repair and recombination protein RAD26; Protein involved in transcription-coupled nucleotide excision repair; repairs UV-induced DNA lesions; recruitment to DNA lesions is dependent on an elongating RNA polymerase II; homolog of human CSB protein; Belongs to the SNF2/RAD54 helicase family. (1085 aa)
POL31Subunit of DNA polymerase delta (polymerase III); essential for cell viability; involved in DNA replication and DNA repair; forms a complex with Rev3p, Rev7p and Pol32p; relocalizes to the cytosol in response to hypoxia. (487 aa)
SWE1Mitosis 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)
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)
ASF1Histone chaperone ASF1; Nucleosome assembly factor; involved in chromatin assembly, disassembly; required for recovery after DSB repair; role in H3K56 acetylation required for expression homeostasis, buffering mRNA synthesis rate against gene dosage changes in S phase; anti-silencing protein, derepresses silent loci when overexpressed; role in regulating Ty1 transposition; relocalizes to cytosol under hypoxia; growth defect of asf1 null is functionally complemented by either human ASF1A or ASF1B. (279 aa)
GSH1Glutamate--cysteine ligase; Gamma glutamylcysteine synthetase; catalyzes the first step in glutathione (GSH) biosynthesis; expression induced by oxidants, cadmium, and mercury; protein abundance increases in response to DNA replication stress; Belongs to the glutamate--cysteine ligase type 3 family. (678 aa)
SRS2DNA helicase and DNA-dependent ATPase; involved in DNA repair and checkpoint recovery, needed for proper timing of commitment to meiotic recombination and transition from Meiosis I to II; blocks trinucleotide repeat expansion; affects genome stability; disassembles Rad51p nucleoprotein filaments during meiotic recombination; stimulates activity of the Mus81p-Mms4p endonuclease, independently of Srs2p catalytic activity; functional homolog of human RTEL1. (1174 aa)
DPB11DNA replication initiation protein; loads DNA pol epsilon onto pre-replication complexes at origins; checkpoint sensor recruited to stalled replication forks by the checkpoint clamp complex where it activates Mec1p; along with Rfa1p, binds to ultrafine anaphase bridges in mitotic cells and prevents accumulation of chromatin bridges by stimulating the Mec1p kinase and suppressing homologous recombination; ortholog of human TopBP1; forms nuclear foci upon DNA replication stress. (764 aa)
MAD2Component 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)
RNR2Ribonucleoside-diphosphate reductase small chain 1; Ribonucleotide-diphosphate reductase (RNR), small subunit; the RNR complex catalyzes the rate-limiting step in dNTP synthesis and is regulated by DNA replication and DNA damage checkpoint pathways via localization of the small subunits; RNR2 has a paralog, RNR4, that arose from the whole genome duplication; Belongs to the ribonucleoside diphosphate reductase small chain family. (399 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)
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)
MPH13'-5' DNA helicase involved in error-free bypass of DNA lesions; binds flap DNA, stimulates activity of Rad27p and Dna2p; prevents crossovers between ectopic sequences by removing substrates for Mus81-Mms4 or Rad1-Rad10 cleavage; homolog of human FANCM Fanconi anemia protein that is involved in stabilizing and remodeling blocked replication forks; member of SF2 DExD/H superfamily of helicases; nonsense or missense mutations in FANCM can make people more likely to get cancer. (993 aa)
MCM10Minichromosome maintenance protein 10; Essential chromatin-associated protein; involved in initiation of DNA replication; required for association of MCM2-7 complex with replication origins; required to stabilize catalytic subunit of DNA polymerase-alpha; self-associates through its N-terminal domain. (571 aa)
REV7Accessory subunit of DNA polymerase zeta; involved in translesion synthesis during post-replication repair; required for mutagenesis induced by DNA damage; involved in double-strand break repair; forms a complex with Rev3p, Pol31p and Pol32p; Belongs to the MAD2 family. (245 aa)
MET18DNA repair/transcription protein MET18/MMS19; Component of cytosolic iron-sulfur protein assembly (CIA) machinery; acts at a late step of Fe-S cluster assembly; forms the CIA targeting complex with Cia1p and Cia2p that directs Fe-S cluster incorporation into a subset of proteins involved in methionine biosynthesis, DNA replication and repair, transcription, and telomere maintenance; ortholog of human MMS19; Belongs to the MET18/MMS19 family. (1032 aa)
RNR3Ribonucleoside-diphosphate reductase large chain 2; Minor isoform of large subunit of ribonucleotide-diphosphate reductase; the RNR complex catalyzes rate-limiting step in dNTP synthesis, regulated by DNA replication and DNA damage checkpoint pathways via localization of small subunits; RNR3 has a paralog, RNR1, that arose from the whole genome duplication. (869 aa)
ULP2Ubiquitin-like-specific protease 2; Peptidase that deconjugates Smt3/SUMO-1 peptides from proteins; plays a role in chromosome cohesion at centromeric regions and recovery from checkpoint arrest induced by DNA damage or DNA replication defects; potential Cdc28p substrate; human homolog PML implicated in promyelocytic leukemia can partially complement yeast null mutant; Belongs to the peptidase C48 family. (1034 aa)
DNA2Tripartite DNA replication factor; single-stranded DNA-dependent ATPase, ATP-dependent nuclease, helicase; tracking protein for flap cleavage during Okazaki fragment maturation; involved in DNA repair/processing of meiotic DNA double strand breaks; component of telomeric chromatin with cell-cycle dependent localization; required for telomerase-dependent telomere synthesis; forms nuclear foci upon DNA replication stress; human homolog DNA2 complements yeast dna2 mutant. (1522 aa)
MSH1DNA-binding protein of the mitochondria; involved in repair of mitochondrial DNA; has ATPase activity and binds to DNA mismatches; has homology to E. coli MutS; transcription is induced during meiosis. (959 aa)
YHR097CUncharacterized protein YHR097C; Putative protein of unknown function; green fluorescent protein (GFP)-fusion protein localizes to the cytoplasm and the nucleus; YHR097C has a paralog, PAL1, that arose from the whole genome duplication. (366 aa)
RRM3DNA helicase involved in rDNA replication and Ty1 transposition; binds to and suppresses DNA damage at G4 motifs in vivo; relieves replication fork pauses at telomeric regions; structurally and functionally related to Pif1p; Belongs to the helicase family. (723 aa)
SOD2Mitochondrial manganese superoxide dismutase; protects cells against oxygen toxicity and oxidative stress; human mitochondrial SOD2 can complement a yeast null mutant and human cytoplasmic SOD1 can also complement when targeted to the mitochondrial matrix. (233 aa)
RAD2DNA repair protein RAD2; Single-stranded DNA endonuclease; cleaves single-stranded DNA during nucleotide excision repair to excise damaged DNA; subunit of Nucleotide Excision Repair Factor 3 (NEF3); homolog of human XPG protein. (1031 aa)
TRX2Thioredoxin-2; Cytoplasmic thioredoxin isoenzyme; part of thioredoxin system which protects cells against oxidative and reductive stress; forms LMA1 complex with Pbi2p; acts as a cofactor for Tsa1p; required for ER-Golgi transport and vacuole inheritance; with Trx1p, facilitates mitochondrial import of small Tims Tim9p, Tim10p, Tim13p by maintaining them in reduced form; abundance increases under DNA replication stress; TRX2 has a paralog, TRX1, that arose from the whole genome duplication. (104 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)
RNR4Ribonucleoside-diphosphate reductase small chain 2; Ribonucleotide-diphosphate reductase (RNR) small subunit; the RNR complex catalyzes the rate-limiting step in dNTP synthesis and is regulated by DNA replication and DNA damage checkpoint pathways via localization of the small subunits; relocalizes from nucleus to cytoplasm upon DNA replication stress; RNR4 has a paralog, RNR2, that arose from the whole genome duplication. (345 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)
COX4Subunit IV of cytochrome c oxidase; the terminal member of the mitochondrial inner membrane electron transport chain; precursor N-terminal 25 residues are cleaved during mitochondrial import; phosphorylated; spermidine enhances translation. (155 aa)
SAE2DNA 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)
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)
LIF1Ligase-interacting factor 1; Component of the DNA ligase IV complex; this complex mediates nonhomologous end joining in DNA double-strand break repair; physically interacts with Dnl4p and Nej1p; homologous to mammalian XRCC4 protein. (421 aa)
MMS2Ubiquitin-conjugating enzyme variant; involved in error-free postreplication repair; forms a heteromeric complex with Ubc13p, an active ubiquitin-conjugating enzyme; cooperates with chromatin-associated RING finger proteins, Rad18p and Rad5p; protein abundance increases in response to DNA replication stress. (137 aa)
MAD1Coiled-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)
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)
RAD6Ubiquitin-conjugating enzyme (E2); involved in postreplication repair as a heterodimer with Rad18p, regulation of K63 polyubiquitination in response to oxidative stress, DSBR and checkpoint control as a heterodimer with Bre1p, ubiquitin-mediated N-end rule protein degradation as a heterodimer with Ubr1p, ERAD with Ubr1p in the absence of canonical ER membrane ligases, and Rpn4p turnover as part of proteasome homeostasis, in complex with Ubr2p and Mub1p. (172 aa)
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)
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)
RAD35' to 3' DNA helicase; involved in nucleotide excision repair and transcription; subunit of RNA polII initiation factor TFIIH and of Nucleotide Excision Repair Factor 3 (NEF3); homolog of human XPD protein; mutant has aneuploidy tolerance; protein abundance increases in response to DNA replication stress; Belongs to the helicase family. RAD3/XPD subfamily. (778 aa)
BEM2GTPase-activating protein BEM2/IPL2; Rho GTPase activating protein (RhoGAP); involved in the control of cytoskeleton organization and cellular morphogenesis; required for bud emergence; potential GAP for Rho4p. (2167 aa)
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)
RNR1Ribonucleoside-diphosphate reductase large chain 1; Major isoform of large subunit of ribonucleotide-diphosphate reductase; the RNR complex catalyzes rate-limiting step in dNTP synthesis, regulated by DNA replication and DNA damage checkpoint pathways via localization of small subunits; relative distribution to the nucleus increases upon DNA replication stress; RNR1 has a paralog, RNR3, that arose from the whole genome duplication. (888 aa)
YEN1Holliday junction resolvase; promotes template switching during break-induced replication (BIR), causing non-reciprocal translocations (NRTs); localization is cell-cycle dependent and regulated by Cdc28p phosphorylation; homolog of human GEN1; similar to S. cerevisiae endonuclease Rth1p. (759 aa)
CAN1Plasma membrane arginine permease; requires phosphatidyl ethanolamine (PE) for localization, exclusively associated with lipid rafts; mutation confers canavanine resistance; CAN1 has a paralog, ALP1, that arose from the whole genome duplication. (590 aa)
URA3Orotidine-5'-phosphate (OMP) decarboxylase; catalyzes the sixth enzymatic step in the de novo biosynthesis of pyrimidines, converting OMP into uridine monophosphate (UMP); converts 5-FOA into 5-fluorouracil, a toxic compound. (267 aa)
MMS21Highly conserved SUMO E3 ligase subunit of SMC5-SMC6 complex; required for anchoring dsDNA breaks to the nuclear periphery; SMC5-SMC6 plays a key role in removal of X-shaped DNA structures that arise between sister chromatids during DNA replication and repair; required for efficient sister chromatid cohesion; mutants are sensitive to MMS, show increased spontaneous mutation and mitotic recombination; SUMOylates and inhibits Snf1p function; supports nucleolar function; Belongs to the NSE2 family. (267 aa)
RAD30DNA polymerase eta; involved in translesion synthesis during post-replication repair; catalyzes the synthesis of DNA opposite cyclobutane pyrimidine dimers and other lesions; involved in formation of post-replicative damage-induced genome-wide cohesion; may also have a role in protection against mitochondrial mutagenesis; mutations in human pol eta are responsible for XPV. (632 aa)
SIZ1SUMO E3 ligase; promotes attachment of small ubiquitin-related modifier sumo (Smt3p) to primarily cytoplasmic proteins; regulates Rsp5p ubiquitin ligase activity and is in turn itself regulated by Rsp5p; required for sumoylation of septins and histone H3 variant Cse4p, a prerequisite for STUbL-mediated Ub-dependent degradation; localizes to the septin ring; acts as an adapter between E2, Ubc9p and substrates; tends to compensate for survival of DNA damage in absence of Nfi1p. (904 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)
RSC3Chromatin structure-remodeling complex protein RSC3; Component of the RSC chromatin remodeling complex; essential gene required for maintenance of proper ploidy and regulation of ribosomal protein genes and the cell wall/stress response; RSC3 has a paralog, RSC30, that arose from the whole genome duplication. (885 aa)
MHR1Mitochondrial ribosomal protein of the large subunit; also involved in homologous recombination in mitochondria; required for recombination-dependent mtDNA partitioning; involved in stimulation of mitochondrial DNA replication in response to oxidative stress. (226 aa)
DIN7DNA damage-inducible protein DIN7; Mitochondrial nuclease functioning in DNA repair and replication; modulates the stability of the mitochondrial genome, induced by exposure to mutagens, also induced during meiosis at a time nearly coincident with commitment to recombination; DIN7 has a paralog, EXO1, that arose from the whole genome duplication; Belongs to the XPG/RAD2 endonuclease family. (430 aa)
RAD9DNA 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)
DPB4Subunit of DNA pol epsilon and of ISW2 chromatin accessibility complex; involved in both chromosomal DNA replication and inheritance of telomeric silencing; stabilizes the interaction of Pol epsilon with primer-template DNA, positively affecting the processivity of the polymerase and exonuclease activities of Pol epsilon; interacts with extranucleosomal DNA and acts as anchor point for ISW2 complex that retains its position on DNA during nucleosome mobilization. (196 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)
FOB1Nucleolar 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)
MSH6Protein required for mismatch repair in mitosis and meiosis; forms a complex with Msh2p to repair both single-base & insertion-deletion mispairs; also involved in interstrand cross-link repair; potentially phosphorylated by Cdc28p. (1242 aa)
UBC13E2 ubiquitin-conjugating enzyme; involved in the error-free DNA postreplication repair pathway; interacts with Mms2p to assemble ubiquitin chains at the Ub Lys-63 residue; DNA damage triggers redistribution from the cytoplasm to the nucleus. (153 aa)
RLI1Translation initiation factor RLI1; Essential Fe-S protein; required for ribosome biogenesis, translation initiation/termination; facilitates binding of multifactor complex (MFC) of initiation factors to small ribosomal subunit; Dom34-Hbs1 complex and Rli1p work in dissociating inactive ribosomes, thereby facilitating translation restart; forms complex with Lto1p and Yae1p; dependency on ROS-labile FeS clusters, activity in nuclear ribosomal-subunit export impaired by mild oxidative stress. (608 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)
PPH3Catalytic subunit of protein phosphatase PP4 complex; Pph3p and Psy2p form active complex, Psy4p may provide substrate specificity; regulates recovery from the DNA damage checkpoint, the gene conversion- and single-strand annealing-mediated pathways of meiotic double-strand break repair and efficient Non-Homologous End-Joining (NHEJ) pathway; involved in activation of Gln3p to alleviate nitrogen catabolite repression; Pph3p and Psy2p localize to foci on meiotic chromosomes. (308 aa)
RAD57DNA repair protein RAD57; 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 Rad55p; Belongs to the RecA family. (460 aa)
NTH1Neutral trehalase, degrades trehalose; required for thermotolerance and may mediate resistance to other cellular stresses; phosphorylated and activated by Cdc28p at the G1/S phase transition to coordinately regulate carbohydrate metabolism and the cell cycle; inhibited by Dcs1p; NTH1 has a paralog, NTH2, that arose from the whole genome duplication; Belongs to the glycosyl hydrolase 37 family. (751 aa)
PTP1Tyrosine-protein phosphatase 1; Phosphotyrosine-specific protein phosphatase; dephosphorylates a broad range of substrates in vivo, including Fpr3p; localized to the cytoplasm and the mitochondria; proposed to be a negative regulator of filamentation; Belongs to the protein-tyrosine phosphatase family. Non- receptor class subfamily. (335 aa)
CDC9DNA ligase I found in nucleus and mitochondria; essential enzyme that joins Okazaki fragments during DNA replication; also acts in ribonucleotide excision repair, base excision repair, and recombination; DNA ligase I mutants trigger ubiquitination of PCNA at K107, facilitating Rad59p-mediated bypass of unligated Okazaki fragments; human homolog LIG1 can complement yeast cdc9 temperature-sensitive mutant at restrictive temperature. (755 aa)
CYK3SH3-domain protein located in the bud neck and cytokinetic actin ring; relocalizes from bud neck to nucleus upon DNA replication stress; activates the chitin synthase activity of Chs2p during cytokinesis; suppressor of growth and cytokinesis defects of chs2 phospho-mutants. (885 aa)
POL3Catalytic subunit of DNA polymerase delta; required for chromosomal DNA replication during mitosis and meiosis, intragenic recombination, repair of double strand DNA breaks, and DNA replication during nucleotide excision repair (NER). (1097 aa)
DUN1DNA 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)
UBC9SUMO-conjugating enzyme involved in the Smt3p conjugation pathway; nuclear protein required for S- and M-phase cyclin degradation and mitotic control; involved in proteolysis mediated by the anaphase-promoting complex cyclosome (APCC). (157 aa)
RAD59Protein involved DNA double-strand break repair; repairs breaks in DNA during vegetative growth via recombination and single-strand annealing; anneals complementary single-stranded DNA; forms nuclear foci upon DNA replication stress; required for loading of Rad52p to DSBs; regulates replication fork progression in DNA ligase I-deficient cells; paralog of Rad52p; Belongs to the RAD52 family. (238 aa)
MBP1Transcription 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)
SIR2Conserved NAD+ dependent histone deacetylase of the Sirtuin family; deacetylation targets are primarily nuclear proteins; required for telomere hypercluster formation in quiescent yeast cells; involved in regulation of lifespan; plays roles in silencing at HML, HMR, telomeres, and rDNA; negatively regulates initiation of DNA replication; functions as regulator of autophagy like mammalian homolog SIRT1, and also of mitophagy. (562 aa)
MSH3Mismatch repair protein; forms dimers with Msh2p that mediate repair of insertion or deletion mutations and removal of nonhomologous DNA ends, contains a PCNA (Pol30p) binding motif required for genome stability; Belongs to the DNA mismatch repair MutS family. MSH3 subfamily. (1018 aa)
RAD18Postreplication repair E3 ubiquitin-protein ligase RAD18; E3 ubiquitin ligase; forms heterodimer with Rad6p to monoubiquitinate PCNA-K164; heterodimer binds single-stranded DNA and has single-stranded DNA dependent ATPase activity; required for postreplication repair; SUMO-targeted ubiquitin ligase (STUbl) that contains a SUMO-interacting motif (SIM) which stimulates its ubiquitin ligase activity towards the sumoylated form of PCNA. (487 aa)
MRC1S-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)
DPB3Third-largest subunit of DNA polymerase II (DNA polymerase epsilon); required to maintain fidelity of chromosomal replication and also for inheritance of telomeric silencing; stabilizes the interaction of Pol epsilon with primer-template DNA, positively affecting the processivity of the polymerase and exonuclease activities of Pol epsilon; mRNA abundance peaks at the G1/S boundary of the cell cycle; DPB3 has a paralog, DLS1, that arose from the whole genome duplication. (201 aa)
SLX1Endonuclease involved in DNA recombination and repair; subunit of a complex, with Slx4p, that hydrolyzes 5' branches from duplex DNA in response to stalled or converging replication forks; function overlaps with that of Sgs1p-Top3p. (304 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)
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)
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)
Q0092Putative uncharacterized protein Q0092, mitochondrial; Dubious open reading frame; unlikely to encode a functional protein, based on available experimental and comparative sequence data. (46 aa)
NTG1Endonuclease III homolog 1; DNA N-glycosylase and apurinic/apyrimidinic (AP) lyase; involved in base excision repair; acts in both nucleus and mitochondrion; creates a double-strand break at mtDNA origins that stimulates replication in response to oxidative stress; required for maintaining mitochondrial genome integrity; NTG1 has a paralog, NTG2, that arose from the whole genome duplication; Belongs to the Nth/MutY family. (399 aa)
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)
APN2DNA-(apurinic or apyrimidinic site) lyase 2; Class II abasic (AP) endonuclease involved in repair of DNA damage; homolog of human HAP1 and E. coli exoIII. (520 aa)
MCM2Protein 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)
POL12B subunit of DNA polymerase alpha-primase complex; required for initiation of DNA replication during mitotic and premeiotic DNA synthesis; also functions in telomere capping and length regulation. (705 aa)
TEL1Serine/threonine-protein kinase TEL1; Protein kinase primarily involved in telomere length regulation; contributes to cell cycle checkpoint control in response to DNA damage; acts with Red1p and Mec1p to promote interhomolog recombination by phosphorylation of Hop1; functionally redundant with Mec1p; regulates P-body formation induced by replication stress; homolog of human ataxia-telangiectasia mutated (ATM) gene. (2787 aa)
ATP1Alpha subunit of the F1 sector of mitochondrial F1F0 ATP synthase; which is a large, evolutionarily conserved enzyme complex required for ATP synthesis; F1 translationally regulates ATP6 and ATP8 expression to achieve a balanced output of ATP synthase genes encoded in nucleus and mitochondria; phosphorylated; N-terminally propionylated in vivo; Belongs to the ATPase alpha/beta chains family. (545 aa)
HHF1Histone H4; core histone protein required for chromatin assembly and chromosome function; one of two identical histone proteins (see also HHF2); contributes to telomeric silencing; N-terminal domain involved in maintaining genomic integrity. (103 aa)
HHT1Histone H3; core histone protein required for chromatin assembly, part of heterochromatin-mediated telomeric and HM silencing; one of two identical histone H3 proteins (see HHT2); regulated by acetylation, methylation, and phosphorylation; H3K14 acetylation plays an important role in the unfolding of strongly positioned nucleosomes during repair of UV damage. (136 aa)
CHS2Chitin synthase II; catalyzes transfer of N-acetylglucosamine (GlcNAc) to chitin upon activation of zymogenic form; required for chitin synthesis in the primary septum during cytokinesis; localization regulated by Cdk1p during mitosis; phosphorylation by Dbf2p kinase regulates its dynamics and chitin synthesis during cytokinesis. (963 aa)
ORC2Subunit 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)
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)
HDA3Subunit of the HDA1 histone deacetylase complex; possibly tetrameric trichostatin A-sensitive class II histone deacetylase complex contains Hda1p homodimer and an Hda2p-Hda3p heterodimer; required for the activity of the complex; relocalizes to the cytosol in response to hypoxia; similar to Hda2p. (655 aa)
DPB2Second largest subunit of DNA polymerase II (DNA polymerase epsilon); required for maintenance of fidelity of chromosomal replication; essential motif in C-terminus is required for formation of the four-subunit Pol epsilon; expression peaks at the G1/S phase boundary; Cdc28p substrate. (689 aa)
TAZ1Lyso-phosphatidylcholine acyltransferase; required for normal phospholipid content of mitochondrial membranes; major determinant of the final acyl chain composition of the mitochondrial-specific phospholipid cardiolipin; mutations in human ortholog tafazzin (TAZ) cause Barth syndrome, a rare X-linked disease characterized by skeletal and cardiomyopathy and bouts of cyclic neutropenia; a specific splice variant of human TAZ can complement yeast null mutant. (381 aa)
CTF4DNA polymerase alpha-binding protein; Chromatin-associated protein; required for sister chromatid cohesion; interacts with DNA polymerase alpha (Pol1p) and may link DNA synthesis to sister chromatid cohesion. (927 aa)
CLB5S-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)
HOS1Class I histone deacetylase (HDAC) family member; deacetylates Smc3p on lysine residues at anaphase onset; has sequence similarity to Hda1p, Rpd3p, Hos2p, and Hos3p; interacts with the Tup1p-Ssn6p corepressor complex. (470 aa)
TAH18NADPH-dependent diflavin oxidoreductase 1; Conserved NAPDH-dependent diflavin reductase; component of an early step in the cytosolic Fe-S protein assembly (CIA) machinery; transfers electrons from NADPH to the Fe-S cluster of Dre2p; plays a pro-death role under oxidative stress; Tah18p-dependent nitric oxide synthesis confers high-temperature stress tolerance; possible target for development of antifungal drugs; In the N-terminal section; belongs to the flavodoxin family. (623 aa)
YME1Catalytic subunit of i-AAA protease complex; complex is located in mitochondrial inner membrane; responsible for degradation of unfolded or misfolded mitochondrial gene products; serves as nonconventional translocation motor to pull PNPase into intermembrane space; also has role in intermembrane space protein folding; mutation causes elevated rate of mitochondrial turnover; human homolog YME1L1 can complement yeast null mutant; In the C-terminal section; belongs to the peptidase M41 family. (747 aa)
HSP82ATP-dependent molecular chaperone HSP82; Hsp90 chaperone; redundant in function with Hsc82p; required for pheromone signaling, negative regulation of Hsf1p; docks with Tom70p for mitochondrial preprotein delivery; promotes telomerase DNA binding, nucleotide addition; protein abundance increases in response to DNA replication stress; contains two acid-rich unstructured regions that promote solubility of chaperone-substrate complexes; HSP82 has a paralog, HSC82, that arose from the whole genome duplication. (709 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)
DDC1DNA 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)
REV3Catalytic subunit of DNA polymerase zeta; involved in translesion synthesis during post-replication repair; required for mutagenesis induced by DNA damage; involved in double-strand break repair; forms a complex with Rev7p, Pol31p and Pol32p; Belongs to the DNA polymerase type-B family. (1504 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)
RAD53Serine/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)
HOS3Histone deacetylase HOS3; Trichostatin A-insensitive homodimeric histone deacetylase (HDAC); specificity in vitro for histones H3, H4, H2A, and H2B; similar to Hda1p, Rpd3p, Hos1p, and Hos2p; deletion results in increased histone acetylation at rDNA repeats. (697 aa)
GLR1Cytosolic and mitochondrial glutathione oxidoreductase; converts oxidized glutathione to reduced glutathione; cytosolic Glr1p is the main determinant of the glutathione redox state of the mitochondrial intermembrane space; mitochondrial Glr1p has a role in resistance to hyperoxia; protein abundance increases in response to DNA replication stress. (483 aa)
GRX5Monothiol glutaredoxin-5, mitochondrial; Glutathione-dependent oxidoreductase; mitochondrial matrix protein involved at an early step in the biogenesis of iron-sulfur centers along with Bol1p; hydroperoxide and superoxide-radical responsive; monothiol glutaredoxin subfamily member along with Grx3p and Grx4p. (150 aa)
RMI1RecQ-mediated genome instability protein 1; Subunit of the RecQ (Sgs1p) - Topo III (Top3p) complex; stimulates superhelical relaxing, DNA catenation/decatenation and ssDNA binding activities of Top3p; involved in response to DNA damage; functions in S phase-mediated cohesion establishment via a pathway involving the Ctf18-RFC complex and Mrc1p; stimulates Top3p DNA catenation/decatenation activity; null mutants display increased rates of recombination and delayed S phase. (241 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)
ULP1Ubiquitin-like-specific protease 1; Protease that specifically cleaves Smt3p protein conjugates; required for cell cycle progression; associates with nucleoporins and may interact with septin rings during telophase; sequestered to the nucleolus under stress conditions. (621 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)
REV1DNA repair protein REV1; Deoxycytidyl transferase; involved in repair of abasic sites and adducted guanines in damaged DNA by translesion synthesis (TLS); forms a complex with the subunits of DNA polymerase zeta, Rev3p and Rev7p; relocalizes from nucleus to cytoplasm upon DNA replication stress. (985 aa)
HNT3Aprataxin-like protein; DNA 5' AMP hydrolase involved in DNA repair; member of the histidine triad (HIT) superfamily of nucleotide-binding proteins; homolog of Aprataxin, a Hint related protein that is mutated in individuals with ataxia with oculomotor apraxia; relative distribution to nuclear foci decreases upon DNA replication stress. (217 aa)
WTM1Transcriptional modulator; involved in regulation of meiosis, silencing, and expression of RNR genes; required for nuclear localization of the ribonucleotide reductase small subunit Rnr2p and Rnr4p; contains WD repeats. (437 aa)
RFC1Subunit of heteropentameric Replication factor C (RF-C); RF-C is a DNA binding protein and ATPase that acts as a clamp loader of the proliferating cell nuclear antigen (PCNA) processivity factor for DNA polymerases delta and epsilon; Belongs to the activator 1 large subunit family. (861 aa)
ELG1Telomere length regulation protein ELG1; Subunit of an alternative replication factor C complex; important for DNA replication and genome integrity; suppresses spontaneous DNA damage; involved in homologous recombination-mediated repair and telomere homeostasis; required for PCNA (Pol30p) unloading during DNA replication; Belongs to the ELG1 family. (791 aa)
CAT55-demethoxyubiquinone hydroxylase, mitochondrial; Protein required for ubiquinone (Coenzyme Q) biosynthesis; localizes to the matrix face of the mitochondrial inner membrane in a large complex with ubiquinone biosynthetic enzymes; required for gluconeogenic gene activation; Belongs to the COQ7 family. (233 aa)
RAS1Ras-like protein 1; GTPase involved in G-protein signaling in adenylate cyclase activation; plays a role in cell proliferation; localized to the plasma membrane; homolog of mammalian RAS proto-oncogenes; relative distribution to the nucleus increases upon DNA replication stress; RAS1 has a paralog, RAS2, that arose from the whole genome duplication. (309 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)
EXO1Exodeoxyribonuclease 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)
BUB3Cell 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)
DNL4DNA ligase required for nonhomologous end-joining (NHEJ); forms stable heterodimer with required cofactor Lif1p, interacts with Nej1p; involved in meiosis, not essential for vegetative growth; mutations in human ortholog lead to ligase IV syndrome and Dubowitz syndrome; Belongs to the ATP-dependent DNA ligase family. (944 aa)
HMI1Mitochondrial inner membrane localized ATP-dependent DNA helicase; required for the maintenance of the mitochondrial genome; not required for mitochondrial transcription; has homology to E. coli helicase uvrD. (706 aa)
MSH2Protein 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)
NTG2Endonuclease III homolog 2; DNA N-glycosylase and apurinic/apyrimidinic (AP) lyase; involved in base excision repair, localizes to the nucleus; sumoylated; NTG2 has a paralog, NTG1, that arose from the whole genome duplication. (380 aa)
TOP1DNA topoisomerase 1; Topoisomerase I; nuclear enzyme that relieves torsional strain in DNA by cleaving and re-sealing the phosphodiester backbone; relaxes both positively and negatively supercoiled DNA; functions in replication, transcription, and recombination; role in processing ribonucleoside monophosphates in genomic DNA into irreversible single-strand breaks; enzymatic activity and interaction with Nsr1p are negatively regulated by polyphosphorylation. (769 aa)
SIN3Transcriptional regulatory protein SIN3; Component of both the Rpd3S and Rpd3L histone deacetylase complexes; involved in transcriptional repression and activation of diverse processes, including mating-type switching and meiosis; involved in the maintenance of chromosomal integrity. (1536 aa)
NRM1Transcription factor NRM1; Transcriptional co-repressor of MBF-regulated gene expression; Nrm1p associates stably with promoters via MCB binding factor (MBF) to repress transcription upon exit from G1 phase. (249 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)
STB1Protein with role in regulation of MBF-specific transcription at Start; phosphorylated by Cln-Cdc28p kinases in vitro; unphosphorylated form binds Swi6p, which is required for Stb1p function; expression is cell-cycle regulated; STB1 has a paralog, YOL131W, that arose from the whole genome duplication. (420 aa)
TOF1Topoisomerase 1-associated factor 1; Subunit of a replication-pausing checkpoint complex; Tof1p-Mrc1p-Csm3p acts at the stalled replication fork to promote sister chromatid cohesion after DNA damage, facilitating gap repair of damaged DNA; interacts with the MCM helicase; relocalizes to the cytosol in response to hypoxia. (1238 aa)
POL2Catalytic 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)
ORC5Subunit of the origin recognition complex (ORC); ORC directs DNA replication by binding to replication origins and is also involved in transcriptional silencing. (479 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)
NAR1Cytosolic Fe-S cluster assembly factor NAR1; Subunit of the cytosolic iron-sulfur (FeS) protein assembly machinery; required for maturation of cytosolic and nuclear FeS proteins and for normal resistance to oxidative stress; deficiency results in shortened lifespan and sensitivity to paraquat; homologous to human Narf. (491 aa)
RTT106Histone chaperone; involved in regulation of chromatin structure in both transcribed and silenced chromosomal regions; affects transcriptional elongation; has a role in regulation of Ty1 transposition; interacts physically and functionally with Chromatin Assembly Factor-1 (CAF-1); Belongs to the RTT106 family. (455 aa)
PSY2Subunit of protein phosphatase PP4 complex; Pph3p and Psy2p form the active complex, Psy4p may provide additional substrate specificity; regulates recovery from the DNA damage checkpoint, the gene conversion- and single-strand annealing-mediated pathways of meiotic double-strand break repair and efficient Non-Homologous End-Joining (NHEJ) pathway; Pph3p and Psy2p localize to foci on meiotic chromosomes; putative homolog of mammalian R3. (858 aa)
UBP10Ubiquitin carboxyl-terminal hydrolase 10; Ubiquitin-specific protease, deubiquitinates Ub-protein moieties; interacts with proteins that function in rRNA production and ribosome biogenesis via its intrinsically disordered regions; stabilizes Rpa190p by deubiquitination; controls PCNA deubiquitylation; may regulate silencing by acting on Sir4p; involved in posttranscriptionally regulating Gap1p, possibly other transporters; localized to the nucleolus; null mutant phenotypes are functionally complemented by human USP36; Belongs to the peptidase C19 family. (792 aa)
RPS3Protein component of the small (40S) ribosomal subunit; has apurinic/apyrimidinic (AP) endonuclease activity; essential for viability; nascent Rps3p is bound by specific chaperone Yar1p during translation; homologous to mammalian ribosomal protein S3 and bacterial S3. (240 aa)
INN1Ingression protein 1; Essential protein that associates with contractile actomyosin ring; required for ingression of the plasma membrane into the bud neck during cytokinesis; C2 domain, a membrane targeting module, is required for function; activates chitin synthase activity of Chs2p during cytokinesis. (409 aa)
POL1Catalytic subunit of the DNA polymerase I alpha-primase complex; required for the initiation of DNA replication during mitotic DNA synthesis and premeiotic DNA synthesis. (1468 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)
PMS1ATP-binding protein required for mismatch repair; required for both mitosis and meiosis; functions as a heterodimer with Mlh1p; binds double- and single-stranded DNA via its N-terminal domain, similar to E. coli MutL. (873 aa)
HHT2Histone H3; core histone protein required for chromatin assembly, part of heterochromatin-mediated telomeric and HM silencing; one of two identical histone H3 proteins (see HHT1); regulated by acetylation, methylation, and phosphorylation; H3K14 acetylation plays an important role in the unfolding of strongly positioned nucleosomes during repair of UV damage. (136 aa)
HHF2Histone H4; core histone protein required for chromatin assembly and chromosome function; one of two identical histone proteins (see also HHF1); contributes to telomeric silencing; N-terminal domain involved in maintaining genomic integrity. (103 aa)
YKU70ATP-dependent DNA helicase II subunit 1; Subunit of the telomeric Ku complex (Yku70p-Yku80p); involved in telomere length maintenance, structure and telomere position effect; required for localization of telomerase ribonucleoprotein to nucleus via interaction with the TLC1 guide RNA; relocates to sites of double-strand cleavage to promote nonhomologous end joining during DSB repair. (602 aa)
ZDS1Protein with a role in regulating Swe1p-dependent polarized growth; involved in maintaining Cdc55p in the cytoplasm where it promotes mitotic entry; involved in mitotic exit through Cdc14p regulation; interacts with silencing proteins at telomeres; has a role in Bcy1p localization; implicated in mRNA nuclear export; ZDS1 has a paralog, ZDS2, that arose from the whole genome duplication. (915 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)
HSC82ATP-dependent molecular chaperone HSC82; Cytoplasmic chaperone of the Hsp90 family; plays a role in determining prion variants; redundant in function and nearly identical with Hsp82p, and together they are essential; expressed constitutively at 10-fold higher basal levels than HSP82 and induced 2-3 fold by heat shock; contains two acid-rich unstructured regions that promote the solubility of chaperone-substrate complexes; HSC82 has a paralog, HSP82, that arose from the whole genome duplication. (705 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)
SAS2Histone acetyltransferase (HAT) catalytic subunit of the SAS complex; acetylates free histones and nucleosomes and regulates transcriptional silencing; member of the MYSTacetyltransferase family; other members are Sas4p and Sas5p. (338 aa)
YKU80ATP-dependent DNA helicase II subunit 2; Subunit of telomeric Ku complex (Yku70p-Yku80p); involved in telomere length maintenance, structure and telomere position effect; required for localization of telomerase ribonucleoprotein via interaction with TLC1 guide RNA; relocates to sites of double-strand cleavage to promote nonhomologous end joining during DSB repair; colocalizes with quiescent cell telomere hyperclusters; Belongs to the ku80 family. (629 aa)
CTF18Chromosome transmission fidelity protein 18; Subunit of a complex with Ctf8p; shares some subunits with Replication Factor C and is required for sister chromatid cohesion; may have overlapping functions with Rad24p in the DNA damage replication checkpoint; Belongs to the activator 1 small subunits family. CTF18 subfamily. (741 aa)
ABF2ARS-binding factor 2, mitochondrial; Mitochondrial DNA-binding protein; involved in mitochondrial DNA replication and recombination, member of HMG1 DNA-binding protein family; activity may be regulated by protein kinase A phosphorylation; ABF2 has a paralog, IXR1, that arose from the whole genome duplication; human homolog TFAM can complement yeast abf2 mutant, rescuing the loss-of-mitochondrial DNA phenotype in a yeast abf2 strain. (183 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)
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)
RAD10DNA repair protein RAD10; Single-stranded DNA endonuclease (with Rad1p); cleaves single-stranded DNA during nucleotide excision repair and double-strand break repair; subunit of Nucleotide Excision Repair Factor 1 (NEF1); homolog of human ERCC1 protein; Belongs to the ERCC1/RAD10/SWI10 family. (210 aa)
OGG1DNA-(apurinic or apyrimidinic site) lyase; Nuclear and mitochondrial glycosylase/lyase; specifically excises 7,8-dihydro-8-oxoguanine residues located opposite cytosine or thymine residues in DNA, repairs oxidative damage to mitochondrial DNA, contributes to UVA resistance; Belongs to the type-1 OGG1 family. (376 aa)
SML1Ribonucleotide reductase inhibitor; involved in regulating dNTP production; regulated by Mec1p and Rad53p during DNA damage and S phase; SML1 has a paralog, DIF1, that arose from the whole genome duplication. (104 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)
UNG1Uracil-DNA glycosylase; required for repair of uracil in DNA formed by spontaneous cytosine deamination; efficiently excises uracil from single-stranded DNA in vivo; not required for strand-specific mismatch repair; cell-cycle regulated, expressed in late G1; localizes to mitochondria and nucleus. (359 aa)
DIF1Damage-regulated import facilitator 1; Protein that regulates nuclear localization of Rnr2p and Rnr4p; phosphorylated by Dun1p in response to DNA damage and degraded; N-terminal half shows similarity to S. pombe Spd1 protein; DIF1 has a paralog, SML1, that arose from the whole genome duplication. (133 aa)
ILV5Ketol-acid reductoisomerase, mitochondrial; Acetohydroxyacid reductoisomerase and mtDNA binding protein; involved in branched-chain amino acid biosynthesis and maintenance of wild-type mitochondrial DNA; found in mitochondrial nucleoids. (395 aa)
ACO1Aconitate hydratase, mitochondrial; Aconitase; required for the tricarboxylic acid (TCA) cycle and also independently required for mitochondrial genome maintenance; component of the mitochondrial nucleoid; mutation leads to glutamate auxotrophy; human homolog ACO2 can complement yeast null mutant. (778 aa)
NEJ1Non-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)
SWI6Regulatory 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)
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)
HOG1Mitogen-activated protein kinase involved in osmoregulation; controls global reallocation of RNAPII during osmotic shock; mediates recruitment/activation of RNAPII at Hot1p-dependent promoters; binds calmodulin; stimulates antisense transcription to activate CDC28; defines novel S-phase checkpoint with Mrc1p that prevent replication/transcription conflicts; nuclear form represses pseudohyphal growth; autophosphorylates; protein abundance increases under DNA replication stress; Belongs to the protein kinase superfamily. Ser/Thr protein kinase family. MAP kinase subfamily. HOG1 sub-subfamily. (435 aa)
CDC45Cell 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)
TRX1Thioredoxin-1; Cytoplasmic thioredoxin isoenzyme; part of thioredoxin system which protects cells against oxidative and reductive stress; forms LMA1 complex with Pbi2p; acts as a cofactor for Tsa1p; required for ER-Golgi transport and vacuole inheritance; with Trx2p, facilitates mitochondrial import of small Tims Tim9p, Tim10p, Tim13p by maintaining them in reduced form; abundance increases iunder DNA replication stress; TRX1 has a paralog, TRX2, that arose from the whole genome duplication. (103 aa)
RAD5DNA repair protein RAD5; DNA helicase/Ubiquitin ligase; involved in error-free DNA damage tolerance (DDT), replication fork regression during postreplication repair by template switching, error-prone translesion synthesis; promotes synthesis of free and PCNA-bound polyubiquitin chains by Ubc13p-Mms2p; forms nuclear foci upon DNA replication stress; associates with native telomeres, cooperates with homologous recombination in senescent cells; human homolog HLTF can complement yeast null mutant. (1169 aa)
DRE2Component of the cytosolic Fe-S protein assembly (CIA) machinery; contains an Fe-S cluster that receives electrons from NADPH via the action of Tah18p in an early step in the CIA pathway; ortholog of human Ciapin1; protein abundance increases in response to DNA replication stress; inviability of the null mutant is functionally complemented by human CIAPIN1. (348 aa)
APN1DNA-(apurinic or apyrimidinic site) lyase 1; Major apurinic/apyrimidinic endonuclease; 3'-repair diesterase; involved in repair of DNA damage by oxidation and alkylating agents; also functions as a 3'-5' exonuclease to repair 7,8-dihydro-8-oxodeoxyguanosine; genetically interacts with NTG1 to maintain mitochondrial genome integrity; Belongs to the AP endonuclease 2 family. (367 aa)
RAD27Flap 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)
DEF1RNAPII degradation factor; forms a complex with Rad26p in chromatin, enables ubiquitination and proteolysis of RNAPII present in an elongation complex; mutant is deficient in Zip1p loading onto chromosomes during meiosis. (738 aa)
CSE4Histone H3-like centromere protein; associated with promoters, accessible chromatin, RNAPII-bound regions; phosphorylated Cse4p associates with centromeres; required for proper kinetochore function; levels regulated by E3 ubiquitin ligase Psh1p; phosphorylation may destabilize defective kinetochores to promote bi-orientation; ubiquitination of N-terminus regulates proteolysis for faithful chromosome segregation; yeast CSE4 can complement mutations in human homolog CENPA. (229 aa)
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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