STRINGSTRING
SPO19 SPO19 AQY1 AQY1 CLB5 CLB5 SMK1 SMK1 SPO24 SPO24 REC8 REC8 SSO1 SSO1 IPL1 IPL1 HRR25 HRR25 CSM4 CSM4 DDC1 DDC1 MLH3 MLH3 MEI5 MEI5 LGE1 LGE1 SMA1 SMA1 RAD1 RAD1 IRC15 IRC15 NUD1 NUD1 RAD17 RAD17 MSC6 MSC6 MEK1 MEK1 YOR338W YOR338W SPS4 SPS4 MUM3 MUM3 OSW1 OSW1 SSP2 SSP2 SPR1 SPR1 MPC54 MPC54 SGO1 SGO1 EXO1 EXO1 RTS1 RTS1 GAS4 GAS4 PAP2 PAP2 NDJ1 NDJ1 SPO21 SPO21 MSH2 MSH2 MHF1 MHF1 SDH5 SDH5 NUF2 NUF2 RRT8 RRT8 LDS2 LDS2 MSO1 MSO1 RFA2 RFA2 MCK1 MCK1 RIM21 RIM21 SEC2 SEC2 RAD50 RAD50 MER1 MER1 SPS18 SPS18 SPS19 SPS19 PSY2 PSY2 YNL194C YNL194C TEP1 TEP1 POL1 POL1 RAS2 RAS2 TOP2 TOP2 END3 END3 PMS1 PMS1 HHT2 HHT2 HHF2 HHF2 SPO1 SPO1 FKS3 FKS3 MRE11 MRE11 SGS1 SGS1 SSO2 SSO2 MLH1 MLH1 OSW5 OSW5 RIM11 RIM11 REC114 REC114 SEC14 SEC14 PDS5 PDS5 RIM9 RIM9 CSM3 CSM3 SPO20 SPO20 CDC5 CDC5 MSC1 MSC1 TUB3 TUB3 RAD10 RAD10 TUB1 TUB1 FPR3 FPR3 SMA2 SMA2 SUR7 SUR7 RAD52 RAD52 GMC2 GMC2 CST9 CST9 GAS2 GAS2 SPO77 SPO77 CHS5 CHS5 REC102 REC102 MMS22 MMS22 CDA2 CDA2 CDA1 CDA1 MEC3 MEC3 YCS4 YCS4 RED1 RED1 TOP3 TOP3 ADY4 ADY4 MSC3 MSC3 CRR1 CRR1 TUB4 TUB4 ATG26 ATG26 SLX4 SLX4 SMC4 SMC4 OSW2 OSW2 MLH2 MLH2 VPS13 VPS13 IRC19 IRC19 SPO75 SPO75 SPO14 SPO14 CWP1 CWP1 MIF2 MIF2 REC107 REC107 RFA3 RFA3 SET2 SET2 SCP160 SCP160 SMC3 SMC3 LOH1 LOH1 IRC18 IRC18 MPS3 MPS3 MAD3 MAD3 YVH1 YVH1 MND2 MND2 DSN1 DSN1 CSM2 CSM2 QDR1 QDR1 HOP1 HOP1 VID28 VID28 PFS1 PFS1 SSP1 SSP1 DNA2 DNA2 REC104 REC104 SPO16 SPO16 SPO12 SPO12 SPS100 SPS100 SET1 SET1 SAE3 SAE3 MSC7 MSC7 SPO13 SPO13 RIM101 RIM101 RIM4 RIM4 SPO11 SPO11 AMA1 AMA1 BUB1 BUB1 CLB6 CLB6 CLB1 CLB1 ESP1 ESP1 SPR3 SPR3 GSC2 GSC2 HFM1 HFM1 RMR1 RMR1 ZIP2 ZIP2 RTF1 RTF1 SHE10 SHE10 KIP3 KIP3 SKI8 SKI8 IME4 IME4 CDC55 CDC55 MND1 MND1 SAE2 SAE2 SPO74 SPO74 RAD54 RAD54 SOH1 SOH1 MPS2 MPS2 AFT1 AFT1 RAD6 RAD6 ERV14 ERV14 HOP2 HOP2 OSW7 OSW7 SMC2 SMC2 CDC14 CDC14 ECO1 ECO1 ACT1 ACT1 TUB2 TUB2 SEC4 SEC4 MSH4 MSH4 ISC10 ISC10 DMC1 DMC1 BMH1 BMH1 RAD24 RAD24 GLC7 GLC7 PMD1 PMD1 SPR6 SPR6 MAM1 MAM1 SHC1 SHC1 RAD51 RAD51 SPO73 SPO73 MEI4 MEI4 GPA2 GPA2 CIN8 CIN8 SPS1 SPS1 SPS2 SPS2 EMI2 EMI2 EMI1 EMI1 ECM11 ECM11 DOT1 DOT1 LRS4 LRS4 DIT1 DIT1 DIT2 DIT2 MUS81 MUS81 XRS2 XRS2 YCG1 YCG1 MCM21 MCM21 ZIP1 ZIP1 DON1 DON1 SWM1 SWM1 CHL4 CHL4 SPR28 SPR28 MSS4 MSS4 CPR1 CPR1 NUM1 NUM1 PDS1 PDS1 SPO71 SPO71 BMH2 BMH2 MSH6 MSH6 RAD55 RAD55 PPH3 PPH3 DBF4 DBF4 VPS54 VPS54 HED1 HED1 RAD61 RAD61 RAD57 RAD57 ADY3 ADY3 FMP45 FMP45 MHF2 MHF2 CLB3 CLB3 MSH5 MSH5 DUN1 DUN1 BRE1 BRE1 ARP2 ARP2 MPS1 MPS1 CDC7 CDC7 MSH3 MSH3 CSM1 CSM1 RRT12 RRT12 SPS22 SPS22 PCH2 PCH2 DTR1 DTR1 CDC28 CDC28 YSW1 YSW1 MEC1 MEC1 IML3 IML3 MMS4 MMS4 POL30 POL30 RDH54 RDH54 MUM2 MUM2 GIP1 GIP1 QDR3 QDR3 CHS3 CHS3 HHT1 HHT1 HHF1 HHF1 SHP1 SHP1 CDC15 CDC15 RFA1 RFA1 PAU8 PAU8 GPB2 GPB2 LDS1 LDS1
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SPO19Sporulation-specific protein 19; Meiosis-specific prospore protein; required to produce bending force necessary for proper assembly of the prospore membrane during sporulation; identified as a weak high-copy suppressor of the spo1-1 ts mutation; SPO19 has a paralog, YOR214C, that arose from the whole genome duplication. (223 aa)
AQY1Aquaporin-1; Spore-specific water channel; mediates the transport of water across cell membranes, developmentally controlled; may play a role in spore maturation, probably by allowing water outflow, may be involved in freeze tolerance. (305 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)
SMK1Middle sporulation-specific mitogen-activated protein kinase (MAPK); required for production of the outer spore wall layers; negatively regulates activity of the glucan synthase subunit Gsc2p. (388 aa)
SPO24Sporulation protein 24; Small (67 amino acids) protein involved in sporulation; localizes to the prospore membrane; phosphorylated during meiosis; a longer, 5'-extended mRNA is also transcribed beginning in mid-meiosis, regulated by two MSEs (middle sporulation elements), and includes an uORF of 15 codons in its 5'-UTR; evidence transcription is regulated by Pdr1p. (67 aa)
REC8Meiotic recombination protein REC8; Meiosis-specific component of the sister chromatid cohesion complex; alpha-kleisin family member that maintains cohesion between sister chromatids during meiosis I; maintains cohesion between centromeres of sister chromatids until meiosis II; independent of its role in sister chromatid cohesion, Rec8p promotes allelic collisions and prevents nonspecific chromosome interactions; homolog of S. pombe Rec8p. (680 aa)
SSO1Protein SSO1; Plasma membrane t-SNARE; involved in fusion of secretory vesicles at the plasma membrane and in vesicle fusion during sporulation; forms a complex with Sec9p that binds v-SNARE Snc2p; syntaxin homolog; functionally redundant with Sso2p; SSO1 has a paralog, SSO2, that arose from the whole genome duplication. (290 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)
HRR25Conserved casein kinase; regulates diverse events including: vesicular traffic, DNA repair, the CVT pathway, monopolar attachment of sister kinetochores at meiosis I, and ribosomal subunit biogenesis; monopolin subunit; binds the RNAPII CTD; phosphorylates COPII coat subunits; interacts with Sit4p phosphatase; antagonizes calcineurin signaling, reducing nuclear accumulation of Crz1p; phosphorylates Dsn1p, the kinetochore receptor for monopolin; homolog of mammalian CK1delta. (494 aa)
CSM4Protein required for accurate chromosome segregation during meiosis; involved in meiotic telomere clustering (bouquet formation) and telomere-led rapid prophase movements; functions with meiosis-specific telomere-binding protein Ndj1p; CSM4 has a paralog, MPS2, that arose from the whole genome duplication. (156 aa)
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)
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)
MEI5Meiosis-specific protein involved in meiotic recombination; involved in DMC1-dependent meiotic recombination; forms heterodimer with Sae3p; proposed to be an assembly factor for Dmc1p; Belongs to the SFR1/MEI5 family. (222 aa)
LGE1Transcriptional regulatory protein LGE1; Protein of unknown function; null mutant forms abnormally large cells, and homozygous diploid null mutant displays delayed premeiotic DNA synthesis and reduced efficiency of meiotic nuclear division. (332 aa)
SMA1Protein of unknown function involved in prospore membrane assembly; involved in the assembly of the prospore membrane during sporulation; interacts with Spo14p. (245 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)
IRC15Increased recombination centers protein 15; Microtubule associated protein; regulates microtubule dynamics; required for accurate meiotic chromosome segregation; null mutant displays large budded cells due to delayed mitotic progression, increased levels of spontaneous Rad52 foci; IRC15 has a paralog, LPD1, that arose from the whole genome duplication; Belongs to the class-I pyridine nucleotide-disulfide oxidoreductase family. (499 aa)
NUD1Protein NUD1; Component of the spindle pole body outer plaque; acts through the mitotic exit network to specify asymmetric spindle pole body inheritance. (851 aa)
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)
MSC6Multicopy suppressor of HER2 involved in mitochondrial translation; mutant is defective in directing meiotic recombination events to homologous chromatids. (692 aa)
MEK1Meiosis-specific serine/threonine protein kinase; functions in meiotic checkpoint, promotes recombination between homologous chromosomes by suppressing double strand break repair between sister chromatids; stabilizes Hop1-Thr318 phosphorylation to promote interhomolog recombination and checkpoint responses during meiosis. (497 aa)
YOR338WPutative protein of unknown function; YOR338W transcription is regulated by Azf1p and its transcript is a specific target of the G protein effector Scp160p; identified as being required for sporulation in a high-throughput mutant screen; YOR338W has a paralog, FUN19, that arose from the whole genome duplication. (363 aa)
SPS4Sporulation-specific protein 4; Protein whose expression is induced during sporulation; not required for sporulation; heterologous expression in E. coli induces the SOS response that senses DNA damage. (338 aa)
MUM3Protein of unknown function involved in outer spore wall organization; has similarity to the tafazzins superfamily of acyltransferases. (479 aa)
OSW1Outer spore wall protein 1; Protein involved in sporulation; required for the construction of the outer spore wall layers; required for proper localization of Spo14p. (278 aa)
SSP2Sporulation-specific protein 2; Sporulation specific protein that localizes to the spore wall; required for sporulation at a point after meiosis II and during spore wall formation; expression controlled by a tightly regulated middle-meiotic promoter that is activated by Ndt80p; translation of SSP2 mRNA is delayed, such that the mRNA is present as nuclear divisions are taking place but is not engaged by ribosomes until relatively late in meiotic development. (371 aa)
SPR1Sporulation-specific exo-1,3-beta-glucanase; contributes to ascospore thermoresistance; SPR1 has a paralog, EXG1, that arose from the whole genome duplication; Belongs to the glycosyl hydrolase 5 (cellulase A) family. (445 aa)
MPC54Component of the meiotic outer plaque; a membrane-organizing center which is assembled on the cytoplasmic face of the spindle pole body during meiosis II and triggers the formation of the prospore membrane; potential Cdc28p substrate. (464 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)
RTS1Serine/threonine-protein phosphatase 2A 56 kDa regulatory subunit delta isoform; B-type regulatory subunit of protein phosphatase 2A (PP2A); Rts1p and Cdc55p are alternative regulatory subunits for PP2A catalytic subunits, Pph21p and Pph22p; PP2A-Rts1p protects cohesin when recruited by Sgo1p to the pericentromere; highly enriched at centromeres in the absence of Cdc55p; required for maintenance of septin ring organization during cytokinesis, for ring disassembly in G1 and for dephosphorylation of septin, Shs1p; homolog of the mammalian B' subunit of PP2A. (757 aa)
GAS41,3-beta-glucanosyltransferase; involved with Gas2p in spore wall assembly; has similarity to Gas1p; localizes to the cell wall. (471 aa)
PAP2Non-canonical poly(A) polymerase; involved in nuclear RNA degradation as a component of TRAMP; catalyzes polyadenylation of hypomodified tRNAs, and snoRNA and rRNA precursors; required for mRNA surveillance and maintenance of genome integrity, serving as a link between RNA and DNA metabolism; overlapping but non-redundant functions with Trf5p; relocalizes to cytosol in response to hypoxia. (584 aa)
NDJ1Non-disjunction protein 1; Protein that regulates meiotic SPB cohesion and telomere clustering; localizes to both spindle pole bodies (SPBs) and telomeres; required for bouquet formation, effective homolog pairing, ordered cross-over distribution, sister chromatid cohesion at meiotic telomeres, chromosomal segregation and telomere-led rapid prophase movement. (352 aa)
SPO21Sporulation-specific protein 21; Component of the meiotic outer plaque of the spindle pole body; involved in modifying the meiotic outer plaque that is required prior to prospore membrane formation; SPO21 has a paralog, YSW1, that arose from the whole genome duplication. (609 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)
MHF1Inner kinetochore subunit MHF1; Component of the heterotetrameric MHF histone-fold complex; in humans the MHF complex interacts with both DNA and Mph1p ortholog FANCM, a Fanconi anemia complementation group protein, to stabilize and remodel blocked replication forks and repair damaged DNA; mhf1 srs2 double mutants are MMS hypersensitive; ortholog of human centromere constitutive-associated network (CCAN) subunit CENP-S, also known as MHF1. (90 aa)
SDH5Succinate dehydrogenase assembly factor 2, mitochondrial; Protein required for flavinylation of Sdh1p; binds to Sdh1p and promotes FAD cofactor attachment, which is necessary for succinate dehydrogenase (SDH) complex assembly and activity; mutations in human ortholog PGL2 are associated with neuroendocrine tumors (paraganglioma). (162 aa)
NUF2Kinetochore protein NUF2; Component of the kinetochore-associated Ndc80 complex; involved in chromosome segregation, spindle checkpoint activity, and kinetochore clustering; evolutionarily conserved; other members include Ndc80p, Nuf2p, Spc24p, and Spc25p. (451 aa)
RRT8Protein involved in spore wall assembly; shares similarity with Lds1p and Lds2p and a strain mutant for all 3 genes exhibits reduced dityrosine fluorescence relative to the single mutants; identified in a screen for mutants with increased levels of rDNA transcription; green fluorescent protein (GFP)-fusion protein localizes to lipid particles; protein abundance increases in response to DNA replication stress. (342 aa)
LDS2Protein Involved in spore wall assembly; localizes to lipid droplets found on or outside of the prospore membrane; shares similarity with Lds1p and Rrt8p, and a strain mutant for all 3 genes exhibits reduced dityrosine fluorescence relative to the single mutants; green fluorescent protein (GFP)-fusion protein localizes to the cytoplasm in a punctate pattern. (356 aa)
MSO1Protein MSO1; Lipid-interacting protein in SNARE complex assembly machinery; acts at late step in secretion; interacts with membranes through two distinct binding sites; shows genetic and physical interactions with Sec1p; required for prospore membrane formation during sporulation; N-terminus closely associates with plasma membrane, C-terminus colocalizes with Sec4p on intracellular membranes; relocalizes from bud neck to nucleus upon DNA replication stress. (210 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)
MCK1Protein kinase MCK1; Dual-specificity ser/thr and tyrosine protein kinase; roles in chromosome segregation, meiotic entry, genome stability, phosphorylation-dependent protein degradation (Rcn1p and Cdc6p), inhibition of protein kinase A, transcriptional regulation, inhibition of RNA pol III, calcium stress and inhibition of Clb2p-Cdc28p after nuclear division; MCK1 has a paralog, YGK3, that arose from the whole genome duplication. (375 aa)
RIM21pH-response regulator protein palH/RIM21; pH sensor molecule, component of the RIM101 pathway; has a role in cell wall construction and alkaline pH response; is glycosylated and phosphorylated; interacts with Dfg16p and Rim9p to form a pH-sensing complex; localization to the plasma membrane is dependent on Dfg16p and Rim9p; has similarity to A. nidulans PalH; Belongs to the palH/RIM21 family. (533 aa)
SEC2Guanyl-nucleotide exchange factor for the small G-protein Sec4p; essential for post-Golgi vesicle transport and for autophagy; associates with the exocyst, via exocyst subunit Sec15p, on secretory vesicles; Belongs to the SEC2 family. (759 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)
MER1Meiotic recombination 1 protein; mRNA-binding protein required for meiosis-specific mRNA splicing; required for chromosome pairing and meiotic recombination; Mer1p regulon embraces four essential meiotic pre-mRNAs: REC107, HFM1, AMA1 and SPO22. (270 aa)
SPS18Sporulation protein SPS18; Protein of unknown function, contains a putative zinc-binding domain; expressed during sporulation; SPS18 has a paralog, GCS1, that arose from the whole genome duplication. (300 aa)
SPS19Peroxisomal 2,4-dienoyl-CoA reductase; auxiliary enzyme of fatty acid beta-oxidation; homodimeric enzyme required for growth and sporulation on petroselineate medium; expression induced during late sporulation and in the presence of oleate; Belongs to the short-chain dehydrogenases/reductases (SDR) family. (292 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)
YNL194CIntegral membrane protein; required for sporulation and plasma membrane sphingolipid content; similar to SUR7; GFP-fusion protein is induced in response to the DNA-damaging agent MMS; GFP-fusion protein is more abundant at MCCs (membrane compartment occupied by Can1) in the presence of glycerol and oleate; YNL194C has a paralog, FMP45, that arose from the whole genome duplication. (301 aa)
TEP1Probable phosphatidylinositol 3,4,5-trisphosphate 3-phosphatase TEP1; PTEN homolog with no demonstrated inositol lipid phosphatase activity; plays a role in normal sporulation; homolog of human tumor suppressor gene PTEN/MMAC1/TEP1 and fission yeast ptn1. (434 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)
RAS2Ras-like protein 2; GTP-binding protein; regulates nitrogen starvation response, sporulation, and filamentous growth; farnesylation and palmitoylation required for activity and localization to plasma membrane; homolog of mammalian Ras proto-oncogenes; RAS2 has a paralog, RAS1, that arose from the whole genome duplication; Belongs to the small GTPase superfamily. Ras family. (322 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)
END3Actin cytoskeleton-regulatory complex protein END3; EH domain-containing protein involved in endocytosis; actin cytoskeletal organization and cell wall morphogenesis; forms a complex with Sla1p and Pan1p. (349 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)
SPO1Putative meiotic phospholipase SPO1; Meiosis-specific prospore protein; required for meiotic spindle pole body duplication and separation; required to produce bending force necessary for proper prospore membrane assembly during sporulation; has similarity to phospholipase B. (631 aa)
FKS31,3-beta-glucan synthase component FKS3; Protein involved in spore wall assembly; has similarity to 1,3-beta-D-glucan synthase catalytic subunits Fks1p and Gsc2p; the authentic, non-tagged protein is detected in highly purified mitochondria in high-throughput studies; Belongs to the glycosyltransferase 48 family. (1785 aa)
MRE11Double-strand break repair protein MRE11; Nuclease subunit of the MRX complex with Rad50p and Xrs2p; complex functions in repair of DNA double-strand breaks and in telomere stability; Mre11p associates with Ser/Thr-rich ORFs in premeiotic phase; nuclease activity required for MRX function; widely conserved; forms nuclear foci upon DNA replication stress; Belongs to the MRE11/RAD32 family. (692 aa)
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)
SSO2Protein SSO2; Plasma membrane t-SNARE; involved in fusion of secretory vesicles at the plasma membrane; syntaxin homolog that is functionally redundant with Sso1p; SSO2 has a paralog, SSO1, that arose from the whole genome duplication. (295 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)
OSW5Outer spore wall protein 5; Protein of unknown function with possible role in spore wall assembly; predicted to contain an N-terminal transmembrane domain; osw5 null mutant spores exhibit increased spore wall permeability and sensitivity to beta-glucanase digestion; Belongs to the OSW5 family. (148 aa)
RIM11Serine/threonine-protein kinase RIM11/MSD1; Protein kinase; required for signal transduction during entry into meiosis; promotes the formation of the Ime1p-Ume6p complex by phosphorylating Ime1p and Ume6p; shares similarity with mammalian glycogen synthase kinase 3-beta; protein abundance increases in response to DNA replication stress; RIM11 has a paralog, MRK1, that arose from the whole genome duplication; Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. GSK-3 subfamily. (370 aa)
REC114Protein involved in early stages of meiotic recombination; possibly involved in the coordination of recombination and meiotic division; mutations lead to premature initiation of the first meiotic division. (428 aa)
SEC14SEC14 cytosolic factor; Phosphatidylinositol/phosphatidylcholine transfer protein; involved in regulating PtdIns, PtdCho, and ceramide metabolism, products of which regulate intracellular transport and UPR; has a role in localization of lipid raft proteins; functionally homologous to mammalian PITPs; SEC14 has a paralog, YKL091C, that arose from the whole genome duplication. (304 aa)
PDS5Cohesion maintenance factor; involved in sister chromatid condensation and cohesion; colocalizes with cohesin on chromosomes; performs its cohesin maintenance function in pre-anaphase cells by protecting the integrity of the cohesion complex; also required during meiosis; relocalizes to the cytosol in response to hypoxia. (1277 aa)
RIM9pH-response regulator protein palI/RIM9; Plasma membrane protein of unknown function; involved in the proteolytic activation of Rim101p in response to alkaline pH; interacts with Rim21p and Dfg16p to form a pH-sensing complex in the Rim101 pathway and is required to maintain Rim21p levels; has similarity to A. nidulans PalI;; Belongs to the palI/RIM9 family. (239 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)
SPO20Sporulation-specific protein 20; Meiosis-specific subunit of the t-SNARE complex; required for prospore membrane formation during sporulation; similar to but not functionally redundant with Sec9p; binds to phosphatidic acid; SNAP-25 homolog. (397 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)
MSC1Protein of unknown function; mutant is defective in directing meiotic recombination events to homologous chromatids; the authentic, non-tagged protein is detected in highly purified mitochondria and is phosphorylated. (513 aa)
TUB3Tubulin alpha-3 chain; Alpha-tubulin; associates with beta-tubulin (Tub2p) to form tubulin dimer, which polymerizes to form microtubules; expressed at lower level than Tub1p; TUB3 has a paralog, TUB1, that arose from the whole genome duplication. (445 aa)
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)
TUB1Tubulin alpha-1 chain; Alpha-tubulin; associates with beta-tubulin (Tub2p) to form tubulin dimer, which polymerizes to form microtubules; relative distribution to nuclear foci increases upon DNA replication stress; TUB1 has a paralog, TUB3, that arose from the whole genome duplication. (447 aa)
FPR3FK506-binding nuclear protein; Nucleolar peptidyl-prolyl cis-trans isomerase (PPIase); FK506 binding protein; affects expression of multiple genes via its role in nucleosome assembly; phosphorylated by casein kinase II (Cka1p-Cka2p-Ckb1p-Ckb2p) and dephosphorylated by Ptp1p; PPIase domain acts as a transcriptional repressor when tethered to DNA by lexA, and repressor activity is dependent on PPIase activity; FPR3 has a paralog, FPR4, that arose from the whole genome duplication; Belongs to the FKBP-type PPIase family. FKBP3/4 subfamily. (411 aa)
SMA2Meiosis-specific prospore membrane protein; required to produce bending force necessary for proper assembly of the prospore membrane during sporulation; Belongs to the SMA2 family. (369 aa)
SUR7Plasma membrane protein, component of eisosomes; long-lived protein that remains stable in eisosomes of mother cells while other eisosome proteins, Pil1p and Lsp1p, turn over; may function to anchor the eisosome in place; sporulation and plasma membrane sphingolipid content are altered in mutants; localizes to furrow-like invaginations (MCC patches). (302 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)
GMC2Grand meiotic recombination cluster protein 2; Protein involved in meiotic crossing over; component of the Synaptonemal Complex (SC) along with Ecm11p; required for the efficient loading of the SC transverse filament protein, Zip1p; promotes SUMOylation of Ecm11p; mutants are delayed in meiotic nuclear division and are defective in synaptonemal complex assembly; transcription is regulated by Ume6p and induced in response to alpha factor. (188 aa)
CST9Chromosome stability protein 9; SUMO E3 ligase; required for synaptonemal complex formation; localizes to synapsis initiation sites on meiotic chromosomes; associates with centromeres early in meiosis, then with chromosome axes and finally with double-strand break sites that are engaged in repair by crossovers; potential Cdc28p substrate. (482 aa)
GAS21,3-beta-glucanosyltransferase; involved with Gas4p in spore wall assembly; has similarity to Gas1p; Belongs to the glycosyl hydrolase 72 family. (555 aa)
SPO77Sporulation-specific protein 77; Meiosis-specific protein of unknown function; required for spore wall formation during sporulation and for timely prospore membrane closure along with SPS1; required with Sps1p for phosphorylation and turnover of Ssp1p; dispensable for both nuclear divisions during meiosis. (477 aa)
CHS5Chitin biosynthesis protein CHS5; Component of the exomer complex; the exomer which also contains Csh6p, Bch1p, Bch2p, and Bud7, is involved in the export of select proteins, such as chitin synthase Chs3p, from the Golgi to the plasma membrane; interacts selectively with the activated, GTP-bound form of Arf1p; Chs5p is the only protein with a BRCT domain that is not localized to the nucleus. (671 aa)
REC102Protein involved in early stages of meiotic recombination; required for chromosome synapsis; forms a complex with Rec104p and Spo11p necessary during the initiation of recombination. (264 aa)
MMS22E3 ubiquitin-protein ligase substrate receptor MMS22; Subunit of E3 ubiquitin ligase complex involved in replication repair; stabilizes protein components of the replication fork, such as the fork-pausing complex and leading strand polymerase, preventing fork collapse and promoting efficient recovery during replication stress; required for accurate meiotic chromosome segregation; Belongs to the MMS22 family. (1454 aa)
CDA2Chitin deacetylase; together with Cda1p involved in the biosynthesis ascospore wall component, chitosan; required for proper rigidity of the ascospore wall. (312 aa)
CDA1Chitin deacetylase; together with Cda2p involved in the biosynthesis ascospore wall component, chitosan; required for proper rigidity of the ascospore wall. (301 aa)
MEC3DNA damage and meiotic pachytene checkpoint protein; subunit of a heterotrimeric complex (Rad17p-Mec3p-Ddc1p) that forms a sliding clamp, loaded onto partial duplex DNA by a clamp loader complex; homolog of human and S. pombe Hus1. (474 aa)
YCS4Subunit of the condensin complex; required for establishment and maintenance of chromosome condensation, chromosome segregation, chromatin binding of condensin, tRNA gene clustering at the nucleolus, and silencing at the mating type locus; required for replication slow zone (RSZ) breakage following Mec1p inactivation; Belongs to the CND1 (condensin subunit 1) family. (1176 aa)
RED1Protein component of the synaptonemal complex axial elements; involved in chromosome segregation during the first meiotic division; critical for coupling checkpoint signaling to SC formation; promotes interhomolog recombination by phosphorylating Hop1p; also interacts with Mec3p and Ddc1p. (827 aa)
TOP3DNA Topoisomerase III; conserved protein that functions in a complex with Sgs1p and Rmi1p to relax single-stranded negatively-supercoiled DNA preferentially; DNA catenation/decatenation activity is stimulated by RPA and Sgs1p-Top3p-Rmi1p; involved in telomere stability and regulation of mitotic recombination. (656 aa)
ADY4Accumulates dyads protein 4; Structural component of the meiotic outer plaque; outer plaque is a membrane-organizing center that assembles on the cytoplasmic face of the spindle pole body during meiosis II and triggers the formation of the prospore membrane. (493 aa)
MSC3Meiotic sister-chromatid recombination protein 3; Protein of unknown function; green fluorescent protein (GFP)-fusion protein localizes to the cell periphery; msc3 mutants are defective in directing meiotic recombination events to homologous chromatids; potential Cdc28p substrate; protein abundance increases in response to DNA replication stress. (728 aa)
CRR1Probable glycosidase CRR1; Putative glycoside hydrolase of the spore wall envelope; required for normal spore wall assembly, possibly for cross-linking between the glucan and chitosan layers; expressed during sporulation; Belongs to the glycosyl hydrolase 16 family. CRR1 subfamily. (422 aa)
TUB4Gamma-tubulin; involved in nucleating microtubules from both the cytoplasmic and nuclear faces of the spindle pole body; protein abundance increases in response to DNA replication stress. (473 aa)
ATG26UDP-glucose:sterol glucosyltransferase; conserved enzyme involved in synthesis of sterol glucoside membrane lipids; in contrast to ATG26 from P. pastoris, S. cerevisiae ATG26 is not involved in autophagy; Belongs to the glycosyltransferase 28 family. (1198 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)
SMC4Structural maintenance of chromosomes protein 4; Subunit of the condensin complex; condensin reorganizes chromosomes during both mitosis and meiosis; forms a subcomplex with Smc2p that has ATP-hydrolyzing and DNA-binding activity, but other condensin subunits are required for chromatin binding; required for tRNA gene clustering at the nucleolus; potential Cdc28p substrate; Belongs to the SMC family. SMC4 subfamily. (1418 aa)
OSW2Outer spore wall protein 2; Protein of unknown function reputedly involved in spore wall assembly. (724 aa)
MLH2Protein involved in mismatch repair and meiotic recombination; only certain frameshift intermediates are mismatch repair substrates; forms a complex with Mlh1p. (695 aa)
VPS13Vacuolar protein sorting-associated protein 13; Protein involved in prospore membrane morphogenesis; peripheral membrane protein that localizes to the prospore membrane and at numerous membrane contact sites; involved in sporulation, vacuolar protein sorting, prospore membrane formation during sporulation, and protein-Golgi retention; required for mitochondrial integrity; contains a PH-like domain; homologous to human CHAC and COH1 which are involved in Chorea-acanthocytosis and Cohen syndrome, respectively. (3144 aa)
IRC19Increased recombination centers protein 19; Protein of unknown function; YLL033W is not an essential gene but mutant is defective in spore formation; null mutant displays increased levels of spontaneous Rad52p foci. (230 aa)
SPO75Sporulation-specific protein 75; Meiosis-specific protein of unknown function; required for spore wall formation during sporulation; dispensable for both nuclear divisions during meiosis; Belongs to the CSC1 (TC 1.A.17) family. (868 aa)
SPO14Phospholipase D; catalyzes the hydrolysis of phosphatidylcholine, producing choline and phosphatidic acid; involved in Sec14p-independent secretion; required for meiosis and spore formation; differently regulated in secretion and meiosis; participates in transcription initiation and/or early elongation of specific genes; interacts with "foot domain" of RNA polymerase II; deletion results in abnormal CTD-Ser5 phosphorylation of RNA polymerase II at specific promoter regions. (1683 aa)
CWP1Cell wall mannoprotein that localizes to birth scars of daughter cells; linked to a beta-1,3- and beta-1,6-glucan heteropolymer through a phosphodiester bond; required for propionic acid resistance; Belongs to the SRP1/TIP1 family. (239 aa)
MIF2Inner kinetochore subunit MIF2; Protein required for structural integrity of elongating spindles; localizes to the kinetochore; interacts with histones H2A, H2B, and H4; phosphorylated by Ipl1p; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-C and fission yeast cnp3; Belongs to the CENP-C/MIF2 family. (549 aa)
REC107Recombination protein 107; Protein involved in early stages of meiotic recombination; involved in coordination between the initiation of recombination and the first division of meiosis; part of a complex (Rec107p-Mei4p-Rec114p) required for ds break formation. (314 aa)
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)
SET2Histone-lysine N-methyltransferase, H3 lysine-36 specific; Histone methyltransferase with a role in transcriptional elongation; methylates H3 lysine 36 (H3K36), which suppresses incorporation of acetylated histones and signals for the deacetylation of these histones within transcribed genes; associates with the C-terminal domain(CTD) of Rpo21p; H3K36me3 (trimethylation) requires Spt6p, proline 38 on H3, CTD of Rpo21p, Ctk1p, and C-terminal SRI domain of Ste2p; relocalizes to the cytosol in response to hypoxia. (733 aa)
SCP160Protein SCP160; Essential RNA-binding G protein effector of mating response pathway; ligand-activated RNA-binding protein that delivers RNAs involved in polarization and perpetualizing mating signal to shmoo tip during pheromone signaling; Scp160p-mediated RNA trafficking essential for chemotropism and successful mating; mainly associated with nuclear envelope and ER, interacts in mRNA-dependent manner with translating ribosomes via multiple KH domains, similar to vertebrate vigilins. (1222 aa)
SMC3Structural maintenance of chromosomes protein 3; Subunit of the multiprotein cohesin complex; required for sister chromatid cohesion in mitotic cells; also required, with Rec8p, for cohesion and recombination during meiosis; phylogenetically conserved SMC chromosomal ATPase family member. (1230 aa)
LOH1Protein involved in outer spore wall assembly; likely involved directly in dityrosine layer assembly; induced during sporulation; repressed during vegetative growth by Sum1p and Hst1p; sequence similar to adjacent ORF, IRC18/YJL037W, and the irc18 loh1 double mutant exhibits reduced dityrosine fluorescence relative to the single mutants; SWAT-GFP and mCherry fusion proteins localize to the cytosol; proposed role in maintenance of genome integrity. (219 aa)
IRC18Protein involved in outer spore wall assembly; possible role in assembly of the dityrosine layer; similar to adjacent ORF, LOH1; irc18 loh1 double mutant exhibits reduced dityrosine fluorescence relative to single mutants; SWAT-GFP fusion protein localizes to the ER and vacuole, while mCherry fusion localizes to the vacuole; expression induced in respiratory-deficient cells and carbon-limited chemostat culture; null mutant displays increased levels of spontaneous Rad52p foci; Belongs to the OSW4/6 family. (224 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)
YVH1Tyrosine-protein phosphatase YVH1; Dual specificity protein phosphatase; regulates growth, sporulation, and glycogen accumulation in a cAMP-dependent protein kinase cascade dependent manner; mutants are defective in 60S ribosome assembly; positively regulates pre-autophagosomal structure (PAS) formation upon nitrogen starvation or rapamycin treatment. (364 aa)
MND2Subunit of the Anaphase-Promoting Complex/Cyclosome (APC/C); necessary for maintaining sister chromatid cohesion in prophase I of meiosis by inhibiting premature ubiquitination and subsequent degradation of substrates by the APC(Ama1) ubiquitin ligase; Belongs to the APC15 family. (368 aa)
DSN1Kinetochore-associated protein DSN1; Essential component of the MIND kinetochore complex; joins kinetochore subunits contacting DNA to those contacting microtubules; Dsn1p phosphorylation promotes interaction between outer and inner kinetochore proteins; kinetochore receptor for monopolin, via interaction with subunit Csm1p; essential for meiotic but not mitotic chromosome segregation; MIND complex consists of Mtw1p, Nnf1p, Nsl1p and Dsn1p; modified by sumoylation; phosphorylated by monopolin subunit Hrr25p. (576 aa)
CSM2Chromosome segregation in meiosis protein 2; Component of Shu complex (aka PCSS complex); Shu complex also includes Psy3, Shu1, Shu2, and promotes error-free DNA repair,; Shu complex mediates inhibition of Srs2p function; promotes formation of Rad51p filaments; Psy3p and Csm2p contain similar DNA-binding regions which work together to form a single DNA binding site; required for accurate chromosome segregation during meiosis. (213 aa)
QDR1Quinidine resistance protein 1; Multidrug transporter of the major facilitator superfamily; member of the 12-spanner drug:H(+) antiporter DHA1 family; involved in spore wall assembly; sequence similarity to DTR1 and QDR3, and the triple mutant dtr1 qdr1 qdr3 exhibits reduced dityrosine fluorescence relative to the single mutants; required for resistance to quinidine, ketoconazole, fluconazole, and barban; QDR1 has a paralog, AQR1, that arose from the whole genome duplication. (563 aa)
HOP1Meiosis-specific protein required for chromosome synapsis; displays Red1p dependent localization to the unsynapsed axial-lateral elements of the synaptonemal complex; required for chiasma formation; in vitro, displays the ability to promote intra- and intermolecular synapsis between double-stranded DNA molecules and to fold DNA into rigid protein-DNA filaments. (605 aa)
VID28Vacuolar import and degradation protein 28; GID Complex subunit, serves as adaptor for regulatory subunit Vid24p; protein involved in proteasome-dependent catabolite degradation of fructose-1,6-bisphosphatase (FBPase); localized to the nucleus and the cytoplasm. (921 aa)
PFS1Sporulation protein required for prospore membrane formation; required for prospore membrane formation at selected spindle poles; ensures functionality of all four spindle pole bodies during meiosis II; not required for meiotic recombination or meiotic chromosome segregation. (237 aa)
SSP1Sporulation-specific protein 1; Protein involved in the control of meiotic nuclear division; involved in the coordination of meiosis with spore formation; subunit of the leading edge protein (LEP) complex (Ssp1-Ady3-Don1-Irc10) that forms a ring-like structure at the leading edge of the prospore membrane during meiosis II; required for assembly of the leading edge coat and both prospore membrane shaping and organization; transcription is induced midway through meiosis. (571 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)
REC104Protein involved in early stages of meiotic recombination; required for meiotic crossing over; forms a complex with Rec102p and Spo11p necessary during the initiation of recombination. (182 aa)
SPO16Sporulation-specific protein 16; Meiosis-specific protein involved in synaptonemal complex assembly; implicated in regulation of crossover formation; required for sporulation. (198 aa)
SPO12Sporulation-specific protein 12; Nucleolar protein of unknown function; positive regulator of mitotic exit; involved in regulating release of Cdc14p from the nucleolus in early anaphase, may play similar role in meiosis; SPO12 has a paralog, BNS1, that arose from the whole genome duplication. (173 aa)
SPS100Protein required for spore wall maturation; expressed during sporulation; may be a component of the spore wall; expression also induced in cells treated with the mycotoxin patulin; SPS100 has a paralog, YGP1, that arose from the whole genome duplication. (326 aa)
SET1Histone-lysine N-methyltransferase, H3 lysine-4 specific; Histone methyltransferase, subunit of the COMPASS (Set1C) complex; COMPASS methylates histone H3K4; Set1p-dependent H3K4 trimethylation recruits Nrd1p, allowing efficient termination of snoRNAs and cryptic unstable transcripts (CUTs) by Nrd1p-Nab3p-Sen1p pathway; modulates histone acetylation levels in promoter proximal regions to ensure efficient Nrd1p-dependent termination; required in transcriptional silencing near telomeres and at silent mating type loci; has a SET domain; Belongs to the class V-like SAM-binding methyltransf [...] (1080 aa)
SAE3Pachytene arrest protein SAE3; Meiosis-specific protein involved in meiotic recombination; involved in DMC1-dependent meiotic recombination; forms heterodimer with Mei5p; proposed to be an assembly factor for Dmc1p. (91 aa)
MSC7Putative aldehyde dehydrogenase-like protein YHR039C; Protein of unknown function; green fluorescent protein (GFP)-fusion protein localizes to the endoplasmic reticulum; msc7 mutants are defective in directing meiotic recombination events to homologous chromatids. (644 aa)
SPO13Meiosis-specific protein SPO13; Meiotic regulator; involved in maintaining sister chromatid cohesion during meiosis I as well as promoting proper attachment of kinetochores to the spindle during meiosis I and meiosis II; anaphase-promoting complex (APC) substrate that is degraded during anaphase I; expressed only in meiotic cells. (291 aa)
RIM101pH-response transcription factor pacC/RIM101; Cys2His2 zinc-finger transcriptional repressor; involved in alkaline responsive gene repression as part of adaptation to alkaline conditions; involved in cell wall assembly; required for alkaline pH-stimulated haploid invasive growth and sporulation; activated by alkaline-dependent proteolytic processing which results in removal of the C-terminal tail; similar to A. nidulans PacC; Belongs to the pacC/RIM101 family. (625 aa)
RIM4Meiotic activator RIM4; Putative RNA-binding protein; required for the expression of early and middle sporulation genes. (713 aa)
SPO11Meiosis-specific protein that initiates meiotic recombination; initiates meiotic recombination by catalyzing the formation of double-strand breaks in DNA via a transesterification reaction; required for homologous chromosome pairing and synaptonemal complex formation; Belongs to the TOP6A family. (398 aa)
AMA1Meiosis-specific APC/C activator protein AMA1; Activator of meiotic anaphase promoting complex (APC/C); Cdc20p family member; required for initiation of spore wall assembly; required for Clb1p degradation during meiosis; prevents premature assembly of the meiosis I spindle, required for DSB induced prophase I arrest; Belongs to the WD repeat CDC20/Fizzy family. (593 aa)
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)
CLB6S-phase entry cyclin-6; B-type cyclin involved in DNA replication during S phase; activates Cdc28p to promote initiation of DNA synthesis; functions in formation of mitotic spindles along with Clb3p and Clb4p; most abundant during late G1; CLB6 has a paralog, CLB5, that arose from the whole genome duplication. (380 aa)
CLB1G2/mitotic-specific cyclin-1; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the transition from G2 to M phase; accumulates during G2 and M, then targeted via a destruction box motif for ubiquitin-mediated degradation by the proteasome; CLB1 has a paralog, CLB2, that arose from the whole genome duplication. (471 aa)
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)
SPR3Sporulation-regulated protein 3; Sporulation-specific homolog of the CDC3/10/11/12 family of genes; septin protein involved in sporulation; regulated by ABFI; the yeast CDC3/10/11/12 family is a family of bud neck microfilament genes; Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. Septin GTPase family. (512 aa)
GSC2Catalytic subunit of 1,3-beta-glucan synthase; involved in formation of the inner layer of the spore wall; activity positively regulated by Rho1p and negatively by Smk1p; GSC2 has a paralog, FKS1, that arose from the whole genome duplication; Belongs to the glycosyltransferase 48 family. (1895 aa)
HFM1Meiosis specific DNA helicase; involved in the conversion of double-stranded breaks to later recombination intermediates and in crossover control; catalyzes the unwinding of Holliday junctions; has ssDNA and dsDNA stimulated ATPase activity. (1187 aa)
RMR1Protein required for meiotic recombination and gene conversion; null mutant displays reduced PIS1 expression and growth defects on non-fermentable carbon sources and minimal media; GFP-fusion protein localizes to both cytoplasm and nucleus; Belongs to the RMR1 family. (241 aa)
ZIP2Meiosis-specific protein; involved in normal synaptonemal complex formation and pairing between homologous chromosomes during meiosis; relocalizes from mitochondrion to cytoplasm upon DNA replication stress; Belongs to the ZIP2 family. (704 aa)
RTF1RNA polymerase-associated protein RTF1; Subunit of RNAPII-associated chromatin remodeling Paf1 complex; regulates gene expression by directing cotranscriptional histone modification, influences transcription and chromatin structure through several independent functional domains; directly or indirectly regulates DNA-binding properties of Spt15p and relative activities of different TATA elements; involved in transcription elongation as demonstrated by the G-less-based run-on (GLRO) assay. (558 aa)
SHE10Protein involved in outer spore wall assembly; likely involved directly in dityrosine layer assembly; putative GPI-anchored protein; overexpression causes growth arrest;; SWAT-GFP, seamless-GFP and mCherry fusion proteins localize to the endoplasmic reticulum; SHE10 has a paralog, OSW7/YFR039C, that arose from the whole genome duplication; paralogs are redundant for spore wall dityrosine assembly. (577 aa)
KIP3Kinesin-like protein KIP3; Kinesin-related antiparallel sliding motor protein; involved in mitotic spindle positioning; sliding activity promotes bipolar spindle assembly and maintenance of genome stability; inhibits spindle elongation, destabilizing late anaphase spindle microtubules that polymerize beyond the midzone; Belongs to the TRAFAC class myosin-kinesin ATPase superfamily. Kinesin family. Kinesin II subfamily. (805 aa)
SKI8Antiviral protein SKI8; Ski complex component and WD-repeat protein; mediates 3'-5' RNA degradation by the cytoplasmic exosome; also required for meiotic double-strand break recombination; null mutants have superkiller phenotype. (397 aa)
IME4N6-adenosine-methyltransferase IME4; mRNA N6-adenosine methyltransferase required for entry into meiosis; mediates N6-adenosine methylation of bulk mRNA during the induction of sporulation which includes the meiotic regulators IME1, IME2 and IME4 itself; repressed in haploids via production of antisense IME4 transcripts; transcribed in diploid cells where antisense transcription is repressed; orthologous to human METTL3 (MT-A70); Belongs to the MT-A70-like family. (600 aa)
CDC55Regulatory subunit B of protein phosphatase 2A (PP2A); Zds1p/2p-dependent localization to cytoplasm promotes mitotic entry; localization to nucleus prevents mitotic exit; required for correct nuclear division, chromosome segregation during achiasmate meiosis; maintains nucleolar sequestration of Cdc14p during early meiosis; limits formation of PP2A-Rts1p holocomplexes to ensure timely dissolution of sister chromosome cohesion; homolog of mammalian B55. (526 aa)
MND1Protein required for recombination and meiotic nuclear division; forms a complex with Hop2p, which is involved in chromosome pairing and repair of meiotic double-strand breaks. (219 aa)
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)
SPO74Sporulation-specific protein 74; Component of the meiotic outer plaque of the spindle pole body; involved in modifying the meiotic outer plaque that is required prior to prospore membrane formation. (413 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)
SOH1Subunit 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)
MPS2Monopolar spindle protein 2; Essential membrane protein localized at nuclear envelope and SPBs; required for insertion of the newly duplicated spindle pole body into the nuclear envelope; potentially phosphorylated by Cdc28p; MPS2 has a paralog, CSM4, that arose from the whole genome duplication. (387 aa)
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)
ERV14ER-derived vesicles protein ERV14; COPII-coated vesicle protein; involved in vesicle formation and incorporation of specific secretory cargo; required for the delivery of bud-site selection protein Axl2p and Nha1p antiporter to cell surface; related to Drosophila cornichon; ERV14 has a paralog, ERV15, that arose from the whole genome duplication. (138 aa)
HOP2Homologous-pairing protein 2; Meiosis-specific protein that localizes to chromosomes; prevents synapsis between nonhomologous chromosomes and ensures synapsis between homologs; complexes with Mnd1p to promote homolog pairing and meiotic double-strand break repair; heterodimer of Hop2p-Mnd1p stimulates the Dmc1p-mediated strand invasion. (214 aa)
OSW7Protein involved in outer spore wall assembly; likely involved directly in dityrosine layer assembly; may be involved in response to high salt and changes in carbon source; SWAT-GFP, seamless-GFP and mCherry fusion proteins localize to the endoplasmic reticulum; deletion mutant has decreased spore survival in Drosophila feces; OSW7 has a paralog, SHE10, that arose from the whole genome duplication; paralogs are redundant for spore wall dityrosine assembly. (510 aa)
SMC2Structural maintenance of chromosomes protein 2; Subunit of the condensin complex; condensin reorganizes chromosomes during both mitosis and meiosis; essential SMC chromosomal ATPase family member that forms a subcomplex with Smc2p that has ATP-hydrolyzing and DNA-binding activity, but other condensin subunits are required for chromatin binding; required for clustering of tRNA genes at the nucleolus. (1170 aa)
CDC14Tyrosine-protein phosphatase CDC14; Protein phosphatase required for mitotic exit; required for rDNA segregation, cytokinesis, meiosis I spindle disassembly, environmental stress response; held in nucleolus by Cdc55p in early meiosis, liberated by FEAR and Mitotic Exit Network in anaphase, enabling it to effect a decrease in CDK/B-cyclin activity and mitotic exit; sequestered in metaphase II, released upon entry into anaphase II; human homolog CDC14A can complement thermosensitivity of yeast cdc14-1 mutant. (551 aa)
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)
ACT1Actin; structural protein involved in cell polarization, endocytosis, and other cytoskeletal functions. (375 aa)
TUB2Beta-tubulin; associates with alpha-tubulin (Tub1p and Tub3p) to form tubulin dimer, which polymerizes to form microtubules; mutation in human ortholog is associated with congenital fibrosis of the extraocular muscles (CFEOM) with polymicrogyria. (457 aa)
SEC4Ras-related protein SEC4; Rab family GTPase; essential for vesicle-mediated exocytic secretion and autophagy; associates with the exocyst component Sec15p and may regulate polarized delivery of transport vesicles to the exocyst at the plasma membrane. (215 aa)
MSH4MutS protein homolog 4; Protein involved in meiotic recombination; required for normal levels of crossing over, colocalizes with Zip2p to discrete foci on meiotic chromosomes, has homology to bacterial MutS protein; Belongs to the DNA mismatch repair MutS family. (878 aa)
ISC10Meiosis-specific protein ISC10; Protein required for sporulation; transcript is induced 7.5 hours after induction of meiosis, expected to play significant role in the formation of reproductive cells. (267 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)
BMH114-3-3 protein, major isoform; controls proteome at post-transcriptional level, binds proteins and DNA, involved in regulation of exocytosis, vesicle transport, Ras/MAPK and rapamycin-sensitive signaling, aggresome formation, spindle position checkpoint; protein increases in abundance and relative distribution to the nucleus increases upon DNA replication stress; antiapoptotic gene similar to human 14-3-3; BMH1 has a paralog, BMH2, that arose from whole genome duplication. (267 aa)
RAD24Checkpoint protein; involved in the activation of the DNA damage and meiotic pachytene checkpoints; subunit of a clamp loader that loads Rad17p-Mec3p-Ddc1p onto DNA; homolog of human and S. pombe Rad17 protein. (659 aa)
GLC7Serine/threonine-protein phosphatase PP1-2; Type 1 S/T protein phosphatase (PP1) catalytic subunit; involved in glycogen metabolism, sporulation and mitotic progression; interacts with multiple regulatory subunits; regulates actomyosin ring formation; subunit of CPF; recruited to mating projections by Afr1p interaction; regulates nucleocytoplasmic shuttling of Hxk2p; import into the nucleus is inhibited during spindle assembly checkpoint arrest; involved in dephosphorylating Rps6a/b and Bnr1p. (312 aa)
PMD1Protein with an N-terminal kelch-like domain; putative negative regulator of early meiotic gene expression; required, with Mds3p, for growth under alkaline conditions; PMD1 has a paralog, MDS3, that arose from the whole genome duplication. (1753 aa)
SPR6Protein of unknown function; expressed during sporulation; not required for sporulation, but gene exhibits genetic interactions with other genes required for sporulation. (191 aa)
MAM1Monopolin complex subunit MAM1; Monopolin; meiosis-specific kinetochore-associated protein involved in monopolar attachment of sister kinetochores to the meiotic spindle; subunit of monopolin, a complex that prevents biorientation of sister kinetochores to ensure homolog biorientation during meiosis I; regulates the conformation, enzyme kinetics and substrate specificity of the Dsn1p kinase, Hrr1p; expressed only during the first meiotic division. (302 aa)
SHC1Protein SHC1; Sporulation-specific activator of Chs3p (chitin synthase III); required for the synthesis of the chitosan layer of ascospores; transcriptionally induced at alkaline pH; SHC1 has a paralog, SKT5, that arose from the whole genome duplication. (512 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)
SPO73Sporulation-specific protein 73; Meiosis-specific protein required for prospore membrane morphogenesis; required for the proper shape of the prospore membrane (PSM) and for spore wall formation; functions cooperatively with SPO71 in PSM elongation; physically interacts with Spo71p; genetically antagonistic to SPO1, similar to SPO71; localizes to the PSM; required for spore wall formation during sporulation; dispensable for both nuclear divisions during meiosis; dysferlin domain-only protein; Belongs to the SPO73 family. (143 aa)
MEI4Meiosis-specific protein involved in forming DSBs; involved in double-strand break (DSBs) formation during meiotic recombination; required for chromosome synapsis and production of viable spores. (408 aa)
GPA2Guanine nucleotide-binding protein alpha-2 subunit; Nucleotide binding alpha subunit of the heterotrimeric G protein; interacts with the receptor Gpr1p, has signaling role in response to nutrients; required for the recruitment of Ras-GTP at the plasma membrane and in the nucleus. (449 aa)
CIN8Kinesin-like protein CIN8; Kinesin motor protein; involved in mitotic spindle assembly and chromosome segregation. (1000 aa)
SPS1Sporulation-specific protein 1; Putative protein serine/threonine kinase; localizes to the nucleus and cytoplasm; required for efficient spore packaging, prospore membrane development and closure and localization of enzymes involved in spore wall synthesis; interacts with and required for Ssp1p phosphorylation and turnover; member of the GCKIII subfamily of STE20 kinases; multiply phosphorylated on S/T residues; interacts with 14-3-3 proteins, Bmh1p and Bmh2p; expressed at the end of meiosis. (490 aa)
SPS2Protein expressed during sporulation; SPS2 has a paralog, SPS22, that arose from the whole genome duplication; redundant with Sps22p for organization of the beta-glucan layer of the spore wall; S. pombe ortholog is a spore wall component. (502 aa)
EMI2Putative glucokinase-2; Non-essential protein of unknown function; required for transcriptional induction of the early meiotic-specific transcription factor IME1; required for sporulation; expression regulated by glucose-repression transcription factors Mig1/2p; EMI2 has a paralog, GLK1, that arose from the whole genome duplication; protein abundance increases in response to DNA replication stress; Belongs to the hexokinase family. (500 aa)
EMI1Early meiotic induction protein 1; Non-essential protein of unknown function; required for transcriptional induction of the early meiotic-specific transcription factor IME1, also required for sporulation; contains twin cysteine-x9-cysteine motifs; deletion affects mitochondrial morphology. (187 aa)
ECM11Meiosis-specific protein; component of the Synaptonemal Complex (SC) along with Gmc2p; required for efficient crossover formation and for the efficient loading of the SC transverse filament protein, Zip1p; is SUMOlytaed in a Gmc2p manner, and SUMOylation is required for its function in meiosis; GFP fusion protein is present in discrete clusters in the nucleus throughout mitosis; may be involved in maintaining chromatin structure. (302 aa)
DOT1Histone-lysine N-methyltransferase, H3 lysine-79 specific; Nucleosomal histone H3-Lys79 methylase; methylation is required for telomeric silencing, meiotic checkpoint control, and DNA damage response. (582 aa)
LRS4Monopolin complex subunit LRS4; Nucleolar protein that forms a complex with Csm1p; and then Mam1p at kinetochores during meiosis I to mediate accurate homolog segregation; required for condensin recruitment to the replication fork barrier site and rDNA repeat segregation. (347 aa)
DIT1Sporulation-specific enzyme required for spore wall maturation; involved in the production of a soluble LL-dityrosine-containing precursor of the spore wall; transcripts accumulate at the time of prospore enclosure. (536 aa)
DIT2Cytochrome P450-DIT2; N-formyltyrosine oxidase; sporulation-specific microsomal enzyme involved in the production of N,N-bisformyl dityrosine required for spore wall maturation, homologous to cytochrome P-450s; SWAT-GFP and mCherry fusion proteins localize to the endoplasmic reticulum and vacuole respectively. (489 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)
XRS2Protein required for DNA repair; component of the Mre11 complex, which is involved in double strand breaks, meiotic recombination, telomere maintenance, and checkpoint signaling. (854 aa)
YCG1Subunit of the condensin complex; required for establishment and maintenance of chromosome condensation, chromosome segregation and chromatin binding by the complex; required for tRNA genes clustering at the nucleolus; required for replication slow zone breakage following Mec1p inactivation; transcription is cell cycle regulated, peaking in mitosis and declining in G1; protein is constitutively degraded by the proteasome; rate limiting for condensin recruitment to chromatin. (1035 aa)
MCM21Inner kinetochore subunit MCM21; Component of the kinetochore sub-complex COMA; COMA (Ctf19p, Okp1p, Mcm21p, Ame1p) bridges kinetochore subunits in contact with centromeric DNA with subunits bound to microtubules during kinetochore assembly; involved in minichromosome maintenance; modified by sumoylation; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-O and fission yeast mal2; Belongs to the CENP-O/MCM21 family. (368 aa)
ZIP1Transverse filament protein of the synaptonemal complex; required for normal levels of meiotic recombination and pairing between homologous chromosome during meiosis; required for meiotic recombination between non-allelc sites; potential Cdc28p substrate. (875 aa)
DON1Donuts protein 1; Meiosis-specific component of the spindle pole body; subunit of the leading edge protein (LEP) complex (Ssp1-Ady3-Don1-Irc10) that forms a ring-like structure at the leading edge of the prospore membrane (PSM) during meiosis II; required for PSM growth and closure; DON1 has a paralog, CUE5, that arose from the whole genome. (365 aa)
SWM1Subunit of the anaphase-promoting complex (APC); APC is an E3 ubiquitin ligase that regulates the metaphase-anaphase transition and exit from mitosis; required for activation of the daughter-specific gene expression and spore wall maturation; Belongs to the APC13 family. (170 aa)
CHL4Inner kinetochore subunit CHL4; Outer kinetochore protein required for chromosome stability; involved in new kinetochore assembly and sister chromatid cohesion; forms a stable complex with Iml3p; peripheral component of the Ctf19 kinetochore complex that interacts with Ctf19p, Ctf3p, and Mif2p; required for the spindle assembly checkpoint; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-N and fission yeast mis15; Belongs to the CENP-N/CHL4 family. (458 aa)
SPR28Sporulation-regulated protein 28; Sporulation-specific homolog of the CDC3/10/11/12 family of genes; meiotic septin expressed at high levels during meiotic divisions and ascospore formation; the yeast CDC3/10/11/12 family is a family of bud neck microfilament genes; Belongs to the TRAFAC class TrmE-Era-EngA-EngB-Septin-like GTPase superfamily. Septin GTPase family. (423 aa)
MSS4Phosphatidylinositol-4-phosphate 5-kinase; involved in actin cytoskeleton organization and cell morphogenesis; multicopy suppressor of stt4 mutation. (779 aa)
CPR1Cytoplasmic peptidyl-prolyl cis-trans isomerase (cyclophilin); catalyzes the cis-trans isomerization of peptide bonds N-terminal to proline residues; binds the drug cyclosporin A; N-terminally propionylated in vivo; protein abundance increases in response to DNA replication stress; Belongs to the cyclophilin-type PPIase family. PPIase A subfamily. (162 aa)
NUM1Protein required for nuclear migration; component of the mitochondria-ER-cortex-ancor (MECA); required for the association of mitochondria with the cell cortex and for accurate distribution of mitochondrial network; interacts with Mdm36p to link the ER and mitochondria at the cortex; localizes to the mother cell cortex and the bud tip; may mediate interactions of dynein and cytoplasmic microtubules with the cell cortex. (2748 aa)
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)
SPO71Sporulation-specific protein 71; Meiosis-specific protein required for prospore membrane morphogenesis; localizes to the prospore membrane (PSM) during sporulation; required for PSM elongation and closure; genetically antagonistic to SPO1; recruits Vps13p to the PSM during sporulation; interacts and functions cooperatively with Spo73p; mutants have defects in the PSM, aberrant spore wall formation and reduced PtdIns-phosphate pools in the PSM; contains three PH-like domains. (1245 aa)
BMH214-3-3 protein, minor isoform; controls proteome at post-transcriptional level, binds proteins and DNA, involved in regulation of many processes including exocytosis, vesicle transport, Ras/MAPK signaling, and rapamycin-sensitive signaling; protein increases in abundance and relative distribution to the nucleus increases upon DNA replication stress; abundance relative to Bmh1p increases during sporulation. (273 aa)
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)
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)
DBF4Regulatory subunit of Cdc7p-Dbf4p kinase complex; required for Cdc7p kinase activity and initiation of DNA replication; phosphorylates the Mcm2-7 family of proteins; cell cycle regulated; relative distribution to the nucleus increases upon DNA replication stress; co-expression of human CDC7 and DBF4 complements single cdc7 or dbf4 null mutations or the cdc7 dbf4 double null mutation. (704 aa)
VPS54Vacuolar protein sorting-associated protein 54; Component of the GARP (Golgi-associated retrograde protein) complex; GARP is required for the recycling of proteins from endosomes to the late Golgi, and for mitosis after DNA damage induced checkpoint arrest; potentially phosphorylated by Cdc28p; members of the GARP complex are Vps51p-Vps52p-Vps53p-Vps54p. (889 aa)
HED1Meiosis-specific protein; down-regulates Rad51p-mediated mitotic recombination when the meiotic recombination machinery is impaired; promotes synapsis and required for the normal morphogenesis of synaptonemal complex; prevents the recruitment of Rad54p to site-specific DNA double-strand breaks in vivo; early meiotic gene, transcribed specifically during meiotic prophase. (162 aa)
RAD61Protein RAD61; Subunit of a complex that inhibits sister chromatid cohesion; also negatively regulates chromosome condensation; inhibited by Eco1p-acetylated cohesin subunits Smc3p and Mcd1p; binds Smc3p ATPase head of cohesin; related to the human Wapl protein that controls the association of cohesin with chromatin. (647 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)
ADY3Accumulates dyads protein 3; Protein required for spore wall formation; subunit of the leading edge protein (LEP) complex (Ssp1-Ady3-Don1-Irc10) that forms a ring-like structure at the leading edge of the prospore membrane during meiosis II; mediates assembly of the LEP complex, formation of the ring-like structure via interaction with spindle pole body components and prospore membrane maturation; potentially phosphorylated by Cdc28p; ADY3 has a paralog, CNM67, that arose from the whole. (790 aa)
FMP45SUR7 family protein FMP45; Integral membrane protein localized to mitochondria; required for sporulation and maintaining sphingolipid content; similar to SUR7; FMP45 has a paralog, YNL194C, that arose from the whole genome duplication. (309 aa)
MHF2Inner kinetochore subunit MHF2; Component of the heterotetrameric MHF histone-fold complex; in humans the MHF complex interacts with both DNA and Mph1p ortholog FANCM to stabilize and remodel blocked replication forks and repair damaged DNA; mhf2 srs2 double mutants are MMS hypersensitive; orthologous to human centromere constitutive-associated network (CCAN) subunit CENP-X, also known as MHF2. (80 aa)
CLB3G2/mitotic-specific cyclin-3; B-type cyclin involved in cell cycle progression; activates Cdc28p to promote the G2/M transition; may be involved in DNA replication and spindle assembly; accumulates during S phase and G2, then targeted for ubiquitin-mediated degradation; relative distribution to the nucleus increases upon DNA replication stress; CLB3 has a paralog, CLB4, that arose from the whole genome duplication. (427 aa)
MSH5Protein of the MutS family; forms a dimer with Msh4p that facilitates crossovers between homologs during meiosis; msh5-Y823H mutation confers tolerance to DNA alkylating agents; homologs present in C. elegans and humans. (901 aa)
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)
BRE1E3 ubiquitin ligase; forms heterodimer with Rad6p to regulate K63 polyubiquitination in response to oxidative stress and to monoubiquinate histone H2B-K123, which is required for the subsequent methylation of histone H3-K4 and H3-K79; required for DSBR, transcription, silencing, and checkpoint control; interacts with RNA-binding protein Npl3p, linking histone ubiquitination to mRNA processing; Bre1p-dependent histone ubiquitination promotes pre-mRNA splicing. (700 aa)
ARP2Actin-related protein 2; Essential component of the Arp2/3 complex; Arp2/3 is a highly conserved actin nucleation center required for the motility and integrity of actin patches; involved in endocytosis and membrane growth and polarity; required for efficient Golgi-to-ER trafficking in COPI mutants. (391 aa)
MPS1Serine/threonine-protein kinase MPS1; Dual-specificity kinase; autophosphorylation required for function; required for spindle pole body (SPB) duplication and spindle checkpoint function; contributes to bi-orientation by promoting formation of force-generating kinetochore-microtubule attachments in meiosis I; substrates include SPB proteins Spc42p, Spc110p, and Spc98p, mitotic exit network protein Mob1p, kinetochore protein Cnn1p, and checkpoint protein Mad1p; substrate of APCC(Cdh1); similar to human Mps1p. (764 aa)
CDC7Cell division control protein 7; DDK (Dbf4-dependent kinase) catalytic subunit; required for origin firing and replication fork progression in mitotic S phase through phosphorylation of Mcm2-7p complexes and Cdc45p; kinase activity correlates with cyclical DBF4 expression; required for pre-meiotic DNA replication, meiotic DSB formation, recruitment of monopolin complex to kinetochores during meiosis I, regulation of meiosis-specific Ndt80p; mutation complemented by human CDC7 and DBF4 co-expression. (507 aa)
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)
CSM1Monopolin complex subunit CSM1; Nucleolar protein that mediates homolog segregation during meiosis I; forms a complex with Lrs4p and then Mam1p at kinetochores; required for condensin recruitment to the replication fork barrier site and rDNA repeat segregation. (190 aa)
RRT12Subtilase-type proteinase RRT12; Probable subtilisin-family protease; role in formation of the dityrosine layer of spore walls; localizes to the spore wall and also the nuclear envelope and ER region in mature spores. (491 aa)
SPS22Protein of unknown function; SPS22 has a paralog, SPS2, that arose from the whole genome duplication; redundant with Sps2p for the organization of the beta-glucan layer of the spore wall; Belongs to the SPS2 family. (463 aa)
PCH2Pachytene checkpoint protein 2; Hexameric ring ATPase that remodels chromosome axis protein Hop1p; nucleolar component of the pachytene checkpoint, which prevents chromosome segregation when recombination and chromosome synapsis are defective; also represses meiotic interhomolog recombination in rDNA; required for meiotic double-stranded break formation. (564 aa)
DTR1Putative dityrosine transporter of the major facilitator superfamily; member of the 12-spanner drug:H(+) antiporter DHA1 family; required for spore wall synthesis; sequence similarity to QDR1 and QDR3, and the triple mutant dtr1 qdr1 qdr3 exhibits reduced dityrosine fluorescence relative to the single mutants; expressed during sporulation. (572 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)
YSW1Spore-specific protein YSW1; Protein required for normal prospore membrane formation; interacts with Gip1p, which is the meiosis-specific regulatory subunit of the Glc7p protein phosphatase; expressed specifically in spores and localizes to the prospore membrane; YSW1 has a paralog, SPO21, that arose from the whole genome duplication. (609 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)
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)
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)
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)
MUM2Protein essential for meiotic DNA replication and sporulation; cytoplasmic protein; subunit of the MIS complex which controls mRNA methylation during during the induction of sporulation; also interacts with Orc2p, which is a component of the origin recognition complex; Belongs to the fl(2)d family. (366 aa)
GIP1GLC7-interacting protein 1; Meiosis-specific regulatory subunit of the Glc7p protein phosphatase; regulates spore wall formation and septin organization, required for expression of some late meiotic genes and for normal localization of Glc7p. (639 aa)
QDR3Quinidine resistance protein 3; Multidrug transporter of the major facilitator superfamily; member of the 12-spanner drug:H(+) antiporter DHA1 family; has a role in polyamine homeostasis; involved in spore wall asembly; sequence similarity to DTR1 and QDR1, and the triple mutant dtr1 qdr1 qdr3 exhibits reduced dityrosine fluorescence relative to the single mutants; expression is upregulated under polyamine stress; required for resistance to quinidine, barban, cisplatin, and bleomycin. (689 aa)
CHS3Chitin synthase III; catalyzes the transfer of N-acetylglucosamine (GlcNAc) to chitin; required for synthesis of the majority of cell wall chitin, the chitin ring during bud emergence, and spore wall chitosan; contains overlapping di-leucine and di-acidic signals that mediate, respectively, intracellular trafficking by AP-1 and trafficking to plasma membrane by exomer complex; requires AP-3 complex for its intracellular retention. (1165 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)
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)
SHP1UBX domain-containing substrate adaptor for Cdc48p; ubiquitin regulatory X domain-containing protein that acts as a substrate recruiting cofactor for Cdc48p; positively regulates Glc7p PPase activity to promote growth and mitotic progression in complex with Cdc48p; ubiquitinated protein interactor involved in ER-associated degradation (ERAD); regulated by nuclear Ub-dependent degradation (INMAD pathway) independent of the Asi and Doa10 complexes; homolog of human p47 (NSFL1C). (423 aa)
CDC15Cell division control protein 15; Protein kinase of the Mitotic Exit Network; localized to the spindle pole bodies at late anaphase; promotes mitotic exit by directly switching on the kinase activity of Dbf2p; required for spindle disassembly after meiosis II; relocalizes to the cytoplasm upon DNA replication stress. (974 aa)
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)
PAU8Seripauperin-11; Protein of unknown function; member of the seripauperin multigene family encoded mainly in subtelomeric regions; Belongs to the SRP1/TIP1 family. Seripauperin subfamily. (120 aa)
GPB2Guanine nucleotide-binding protein subunit beta 2; Multistep regulator of cAMP-PKA signaling; inhibits PKA downstream of Gpa2p and Cyr1p, thereby increasing cAMP dependency; inhibits Ras activity through direct interactions with Ira1p/2p; regulated by G-alpha protein Gpa2p; GPB2 has a paralog, GPB1, that arose from the whole genome duplication. (880 aa)
LDS1Protein Involved in spore wall assembly; localizes to lipid droplets found on or outside of the prospore membrane; shares similarity with Lds2p and Rrt8p, and a strain mutant for all 3 genes exhibits reduced dityrosine fluorescence relative to the single mutants. (325 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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