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| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| MSS4 | Phosphatidylinositol-4-phosphate 5-kinase; involved in actin cytoskeleton organization and cell morphogenesis; multicopy suppressor of stt4 mutation. (779 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| 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) | ||||
| GAS4 | 1,3-beta-glucanosyltransferase; involved with Gas2p in spore wall assembly; has similarity to Gas1p; localizes to the cell wall. (471 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) | ||||
| 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) | ||||
| 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) | ||||