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| | SPO77 | Sporulation-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) |
| | LDS1 | Protein 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) |
| | MYO4 | Myosin-4; Type V myosin motor involved in actin-based transport of cargos; required for mRNA transport, including ASH1 mRNA, and facilitating the growth and movement of ER tubules into the growing bud along with She3p; MYO4 has a paralog, MYO2, that arose from the whole genome duplication. (1471 aa) |
| | GPB2 | Guanine 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) |
| | PAU8 | Seripauperin-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) |
| | SHP1 | UBX 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) |
| | HHF1 | Histone 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) |
| | HHT1 | Histone 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) |
| | CHS3 | Chitin 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) |
| | FIG1 | Factor-induced gene 1 protein; Integral membrane protein required for efficient mating; may participate in or regulate the low affinity Ca2+ influx system, which affects intracellular signaling and cell-cell fusion during mating. (298 aa) |
| | QDR3 | Quinidine 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) |
| | GIP1 | GLC7-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) |
| | IML3 | Inner 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) |
| | SHE3 | SWI5-dependent HO expression protein 3; Protein adaptor between Myo4p and the She2p-mRNA complex; part of the mRNA localization machinery that restricts accumulation of certain proteins to the bud; also required for cortical ER inheritance. (425 aa) |
| | YSW1 | Spore-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) |
| | DTR1 | Putative 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) |
| | BEM1 | Bud emergence protein 1; Protein containing SH3-domains; involved in establishing cell polarity and morphogenesis; functions as a scaffold protein for complexes that include Cdc24p, Ste5p, Ste20p, and Rsr1p. (551 aa) |
| | SPS22 | Protein 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) |
| | HMLALPHA1 | Silenced copy of ALPHA1 at HML; ALPHA1 encodes a transcriptional coactivator involved in the regulation of mating-type alpha-specific gene expression. (175 aa) |
| | MATALPHA1 | Transcriptional co-activator that regulates mating-type-specific genes; targets the transcription factor Mcm1p to the promoters of alpha-specific genes; one of two genes encoded by the MATalpha mating type cassette. (175 aa) |
| | RRT12 | Subtilase-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) |
| | FIG2 | Factor-induced gene 2 protein; Cell wall adhesin, expressed specifically during mating; may be involved in maintenance of cell wall integrity during mating; FIG2 has a paralog, AGA1, that arose from the whole genome duplication. (1609 aa) |
| | HMRA1 | Silenced copy of a1 at HMR; homeobox corepressor that interacts with Alpha2p to repress haploid-specific gene transcription in diploid cells. (126 aa) |
| | ARP2 | Actin-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) |
| | FMP45 | SUR7 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) |
| | HBT1 | Shmoo tip protein, substrate of Hub1p ubiquitin-like protein; mutants are defective for mating projection formation, thereby implicating Hbt1p in polarized cell morphogenesis; HBT1 has a paralog, YNL195C, that arose from the whole genome duplication. (1046 aa) |
| | HO | Homothallic switching endonuclease; Site-specific endonuclease; required for gene conversion at the MAT locus (homothallic switching) through the generation of a ds DNA break; expression restricted to mother cells in late G1 as controlled by Swi4p-Swi6p, Swi5p, and Ash1p. (586 aa) |
| | ADY3 | Accumulates 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) |
| | VPS54 | Vacuolar 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) |
| | AFR1 | Protein required for pheromone-induced projection (shmoo) formation; regulates septin architecture during mating; has an RVXF motif that mediates targeting of Glc7p to mating projections; interacts with Cdc12p; AFR1 has a paralog, YER158C, that arose from the whole genome duplication; To yeast YER158C. (620 aa) |
| | BMH2 | 14-3-3 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) |
| | SPO71 | Sporulation-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) |
| | CPR1 | Cytoplasmic 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) |
| | MSS4 | Phosphatidylinositol-4-phosphate 5-kinase; involved in actin cytoskeleton organization and cell morphogenesis; multicopy suppressor of stt4 mutation. (779 aa) |
| | SPR28 | Sporulation-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) |
| | SWM1 | Subunit of the anaphase-promoting complex (APC); APC is an E3 ubiquitin ligase that regulates the metaphase-anaphase transition and exit from mitosis; required for activation of the daughter-specific gene expression and spore wall maturation; Belongs to the APC13 family. (170 aa) |
| | DON1 | Donuts 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) |
| | DIT2 | Cytochrome 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) |
| | DIT1 | Sporulation-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) |
| | EMI1 | Early 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) |
| | EMI2 | Putative 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) |
| | SPS2 | Protein 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) |
| | SPS1 | Sporulation-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) |
| | GPA2 | Guanine 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) |
| | SPO73 | Sporulation-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) |
| | SHC1 | Protein 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) |
| | SPR6 | Protein of unknown function; expressed during sporulation; not required for sporulation, but gene exhibits genetic interactions with other genes required for sporulation. (191 aa) |
| | PMD1 | Protein 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) |
| | GLC7 | Serine/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) |
| | BMH1 | 14-3-3 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) |
| | ISC10 | Meiosis-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) |
| | SEC4 | Ras-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) |
| | ACT1 | Actin; structural protein involved in cell polarization, endocytosis, and other cytoskeletal functions. (375 aa) |
| | OSW7 | Protein 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) |
| | ERV14 | ER-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) |
| | SPO74 | Sporulation-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) |
| | SAE2 | DNA endonuclease SAE2; Endonuclease required for telomere elongation; required for telomeric 5' C-rich strand resection; involved in ds-break repair and processing hairpin DNA structures with the MRX complex; function requires sumoylation and phosphorylation; exists as inactive oligomers that are transiently released into smaller active units by phosphorylation; DNA damage triggers Sae2p removal, so active Sae2p is present only transiently; sequence and functional similarity with human CtIP/RBBP8; Belongs to the COM1/SAE2/CtIP family. (345 aa) |
| | SHE10 | Protein 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) |
| | GSC2 | Catalytic 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) |
| | SPR3 | Sporulation-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) |
| | AMA1 | Meiosis-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) |
| | RIM101 | pH-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) |
| | SET1 | Histone-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) |
| | SPS100 | Protein 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) |
| | SPO16 | Sporulation-specific protein 16; Meiosis-specific protein involved in synaptonemal complex assembly; implicated in regulation of crossover formation; required for sporulation. (198 aa) |
| | SSP1 | Sporulation-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) |
| | PFS1 | Sporulation 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) |
| | VID28 | Vacuolar 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) |
| | CKA1 | Alpha catalytic subunit of casein kinase 2 (CK2); a Ser/Thr protein kinase with roles in cell growth and proliferation; CK2, comprised of CKA1, CKA2, CKB1 and CKB2, has many substrates including transcription factors and all RNA polymerases; regulates Fkh1p-mediated donor preference during mating-type switching. (372 aa) |
| | QDR1 | Quinidine 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) |
| | FKH1 | Fork head protein homolog 1; Forkhead family transcription factor; rate-limiting replication origin activator; evolutionarily conserved lifespan regulator; binds multiple chromosomal elements with distinct specificities, cell cycle dynamics; regulates transcription elongation, chromatin silencing at mating loci, expression of G2/M phase genes; facilitates clustering, activation of early-firing replication origins; binds HML recombination enhancer, regulates donor preference during mating-type switching. (484 aa) |
| | MPH1 | 3'-5' DNA helicase involved in error-free bypass of DNA lesions; binds flap DNA, stimulates activity of Rad27p and Dna2p; prevents crossovers between ectopic sequences by removing substrates for Mus81-Mms4 or Rad1-Rad10 cleavage; homolog of human FANCM Fanconi anemia protein that is involved in stabilizing and remodeling blocked replication forks; member of SF2 DExD/H superfamily of helicases; nonsense or missense mutations in FANCM can make people more likely to get cancer. (993 aa) |
| | YVH1 | Tyrosine-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) |
| | IRC18 | Protein 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) |
| | LOH1 | Protein 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) |
| | DPB11 | DNA replication initiation protein; loads DNA pol epsilon onto pre-replication complexes at origins; checkpoint sensor recruited to stalled replication forks by the checkpoint clamp complex where it activates Mec1p; along with Rfa1p, binds to ultrafine anaphase bridges in mitotic cells and prevents accumulation of chromatin bridges by stimulating the Mec1p kinase and suppressing homologous recombination; ortholog of human TopBP1; forms nuclear foci upon DNA replication stress. (764 aa) |
| | SET2 | Histone-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) |
| | CWP1 | Cell 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) |
| | RAD27 | Flap endonuclease 1; 5' to 3' exonuclease, 5' flap endonuclease; required for Okazaki fragment processing and maturation, for long-patch base-excision repair and large loop repair (LLR), ribonucleotide excision repair; member of the S. pombe RAD2/FEN1 family; relocalizes to the cytosol in response to hypoxia. (382 aa) |
| | SHE2 | SWI5-dependent HO expression protein 2; RNA-binding protein that binds specific mRNAs and interacts with She3p; part of the mRNA localization machinery that restricts accumulation of certain proteins to the bud; binds to ER-derived membranes and targets mRNAs to cortical ER. (246 aa) |
| | SPO14 | Phospholipase 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) |
| | SPO75 | Sporulation-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) |
| | SPA2 | Protein SPA2; Component of the polarisome; functions in actin cytoskeletal organization during polarized growth; acts as a scaffold for Mkk1p and Mpk1p cell wall integrity signaling components; potential Cdc28p substrate; coding sequence contains length polymorphisms in different strains; SPA2 has a paralog, SPH1, that arose from the whole genome duplication. (1466 aa) |
| | IRC19 | Increased 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) |
| | VPS13 | Vacuolar 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) |
| | OSW2 | Outer spore wall protein 2; Protein of unknown function reputedly involved in spore wall assembly. (724 aa) |
| | ATG26 | UDP-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) |
| | CRR1 | Probable 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) |
| | ADY4 | Accumulates 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) |
| | CDA1 | Chitin deacetylase; together with Cda2p involved in the biosynthesis ascospore wall component, chitosan; required for proper rigidity of the ascospore wall. (301 aa) |
| | CDA2 | Chitin deacetylase; together with Cda1p involved in the biosynthesis ascospore wall component, chitosan; required for proper rigidity of the ascospore wall. (312 aa) |
| | SPH1 | Protein involved in shmoo formation and bipolar bud site selection; localizes to sites of polarized growth in a cell cycle dependent- and Spa2p-dependent manner, interacts with MAPKKs Mkk1p, Mkk2p, and Ste7p; SPH1 has a paralog, SPA2, that arose from the whole genome duplication. (530 aa) |
| | CHS5 | Chitin 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) |
| | GAS2 | 1,3-beta-glucanosyltransferase; involved with Gas4p in spore wall assembly; has similarity to Gas1p; Belongs to the glycosyl hydrolase 72 family. (555 aa) |
| | SUR7 | Plasma 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) |
| | SMA2 | Meiosis-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) |
| | SPO20 | Sporulation-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) |
| | RIM9 | pH-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) |
| | SEC14 | SEC14 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) |
| | YKU80 | ATP-dependent DNA helicase II subunit 2; Subunit of telomeric Ku complex (Yku70p-Yku80p); involved in telomere length maintenance, structure and telomere position effect; required for localization of telomerase ribonucleoprotein via interaction with TLC1 guide RNA; relocates to sites of double-strand cleavage to promote nonhomologous end joining during DSB repair; colocalizes with quiescent cell telomere hyperclusters; Belongs to the ku80 family. (629 aa) |
| | RIM11 | Serine/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) |
| | FDO1 | Uncharacterized protein YMR144W; Protein involved in directionality of mating type switching; acts with Fkh1p to control which donor mating-type locus is inserted into MAT locus during mating type switching; localized to the nucleus; not an essential gene. (342 aa) |
| | OSW5 | Outer 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) |
| | SSO2 | Protein 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) |
| | SGS1 | ATP-dependent helicase SGS1; RecQ family nucleolar DNA helicase; role in genome integrity maintenance, chromosome synapsis, meiotic joint molecule/crossover formation; stimulates activity of Top3p; rapidly lost in response to rapamycin in Rrd1p-dependent manner; forms nuclear foci upon DNA replication stress; yeast SGS1 complements mutations in human homolog BLM implicated in Bloom syndrome; also similar to human WRN implicated in Werner syndrome; human BLM and WRN can each complement yeast null mutant; Belongs to the helicase family. RecQ subfamily. (1447 aa) |
| | MRE11 | Double-strand break repair protein MRE11; Nuclease subunit of the MRX complex with Rad50p and Xrs2p; complex functions in repair of DNA double-strand breaks and in telomere stability; Mre11p associates with Ser/Thr-rich ORFs in premeiotic phase; nuclease activity required for MRX function; widely conserved; forms nuclear foci upon DNA replication stress; Belongs to the MRE11/RAD32 family. (692 aa) |
| | FKS3 | 1,3-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) |
| | SPO1 | Putative 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) |
| | HHF2 | Histone 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) |
| | HHT2 | Histone 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) |
| | END3 | Actin 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) |
| | RAS2 | Ras-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) |
| | TEP1 | Probable 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) |
| | YNL194C | Integral 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) |
| | SPS19 | Peroxisomal 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) |
| | SPS18 | Sporulation 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) |
| | SEC2 | Guanyl-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) |
| | RIM21 | pH-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) |
| | MCK1 | Protein 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) |
| | HUB1 | Ubiquitin-like protein modifier; promotes alternative splicing of SRC1 pre-mRNA; binds non-covalently to the HIND domain of Snu66, may function in modification of Sph1p and Hbt1p, functionally complemented by the human or S. pombe ortholog; mechanism of Hub1p adduct formation not yet clear. (73 aa) |
| | MSO1 | Protein 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) |
| | LDS2 | Protein 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) |
| | RRT8 | Protein 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) |
| | SDH5 | Succinate 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) |
| | SPO21 | Sporulation-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) |
| | MDY2 | Ubiquitin-like protein MDY2; Protein involved in inserting tail-anchored proteins into ER membranes; forms a complex with Get4p; required for efficient mating; involved in shmoo formation and nuclear migration in the pre-zygote; associates with ribosomes. (212 aa) |
| | GAS4 | 1,3-beta-glucanosyltransferase; involved with Gas2p in spore wall assembly; has similarity to Gas1p; localizes to the cell wall. (471 aa) |
| | EXO1 | Exodeoxyribonuclease 1; 5'-3' exonuclease and flap-endonuclease; involved in recombination, double-strand break repair, MMS2 error-free branch of the post replication (PRR) pathway and DNA mismatch repair; role in telomere maintenance; member of the Rad2p nuclease family, with conserved N and I nuclease domains; relative distribution to the nucleus increases upon DNA replication stress; EXO1 has a paralog, DIN7, that arose from the whole genome duplication. (702 aa) |
| | SHE4 | SWI5-dependent HO expression protein 4; Protein containing a UCS (UNC-45/CRO1/SHE4) domain; binds to myosin motor domains to regulate myosin function; involved in endocytosis, polarization of the actin cytoskeleton, and asymmetric mRNA localization. (789 aa) |
| | CKA2 | Alpha' catalytic subunit of casein kinase 2 (CK2); CK2 is a Ser/Thr protein kinase with roles in cell growth and proliferation; CK2, comprised of CKA1, CKA2, CKB1 and CKB2, has many substrates including transcription factors and all RNA polymerases; protein abundance increases in response to DNA replication stress; regulates Fkh1p-mediated donor preference during mating-type switching. (339 aa) |
| | MPC54 | Component 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) |
| | SPR1 | Sporulation-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) |
| | ULS1 | ATP-dependent helicase ULS1; Swi2/Snf2-related translocase, SUMO-Targeted Ubiquitin Ligase (STUbL); required for maintenance of NHEJ inhibition at telomeres; functions at telomeres to translocate and ubiquitinylate poly-sumoylated Rap1p for proteosomal degradation; plays role in antagonizing silencing during mating-type switching; only known STUbL with a translocase activity; contains RING finger domain; relocalizes from nucleus to cytoplasm upon DNA replication stress. (1619 aa) |
| | SSP2 | Sporulation-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) |
| | OSW1 | Outer 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) |
| | MUM3 | Protein of unknown function involved in outer spore wall organization; has similarity to the tafazzins superfamily of acyltransferases. (479 aa) |
| | SPS4 | Sporulation-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) |
| | YOR338W | Putative 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) |
| | SMA1 | Protein of unknown function involved in prospore membrane assembly; involved in the assembly of the prospore membrane during sporulation; interacts with Spo14p. (245 aa) |
| | SPO19 | Sporulation-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) |
| | SSO1 | Protein 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) |
| | SPO24 | Sporulation 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) |
| | SMK1 | Middle 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) |
| | AQY1 | Aquaporin-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) |
