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| | MAF1 | Highly conserved negative regulator of RNA polymerase III; involved in tRNA processing and stability; inhibits tRNA degradation via rapid tRNA decay (RTD) pathway; binds N-terminal domain of Rpc160p subunit of Pol III to prevent closed-complex formation; regulated by phosphorylation mediated by TORC1, protein kinase A, Sch9p, casein kinase 2; localizes to cytoplasm during vegetative growth and translocates to nucleus and nucleolus under stress conditions. (395 aa) |
| | RPO26 | RNA polymerase subunit ABC23; common to RNA polymerases I, II, and III; part of central core; similar to bacterial omega subunit; Belongs to the archaeal RpoK/eukaryotic RPB6 RNA polymerase subunit family. (155 aa) |
| | JIP5 | WD repeat-containing protein JIP5; Protein required for biogenesis of the large ribosomal subunit; required for biogenesis of the large ribosomal subunit; interacts with proteins involved in RNA processing, ribosome biogenesis, ubiquitination and demethylation; similar to WDR55, a human WD repeat protein; essential gene. (492 aa) |
| | NOC4 | Nucleolar protein; forms a complex with Nop14p that mediates maturation and nuclear export of 40S ribosomal subunits; relocalizes to the cytosol in response to hypoxia. (552 aa) |
| | RRP15 | Nucleolar protein; constituent of pre-60S ribosomal particles; required for proper processing of the 27S pre-rRNA at the A3 and B1 sites to yield mature 5.8S and 25S rRNAs; Belongs to the RRP15 family. (250 aa) |
| | RRP9 | Ribosomal RNA-processing protein 9; Protein involved in pre-rRNA processing; associated with U3 snRNP; component of small ribosomal subunit (SSU) processosome; ortholog of the human U3-55k protein; Belongs to the WD repeat RRP9 family. (573 aa) |
| | MRD1 | Multiple RNA-binding domain-containing protein 1; Essential conserved small ribosomal subunit (40s) synthesis factor; component of the 90S preribosome; required for production of 18S rRNA and small ribosomal subunit; contains five consensus RNA-binding domains and binds to the pre-rRNA at two sites within the 18S region. (887 aa) |
| | RPC40 | RNA polymerase subunit AC40; common to RNA polymerase I and III; predominant determinant targeting Ty1 integration upstream of Pol III-transcribed genes. (335 aa) |
| | FHL1 | Pre-rRNA-processing protein FHL1; Regulator of ribosomal protein (RP) transcription; has forkhead associated domain that binds phosphorylated proteins; recruits coactivator Ifh1p or corepressor Crf1p to RP gene promoters; also has forkhead DNA-binding domain though in vitro DNA binding assays give inconsistent results; computational analyses suggest it binds DNA directly at highly active RP genes and indirectly through Rap1p motifs at others; suppresses RNA pol III and splicing factor prp4 mutants. (936 aa) |
| | TIF6 | Eukaryotic translation initiation factor 6; Constituent of 66S pre-ribosomal particles; has similarity to human translation initiation factor 6 (eIF6); may be involved in the biogenesis and or stability of 60S ribosomal subunits; Belongs to the eIF-6 family. (245 aa) |
| | RPA135 | RNA polymerase I second largest subunit A135. (1203 aa) |
| | DIM1 | Essential 18S rRNA dimethylase (dimethyladenosine transferase); responsible for conserved m6(2)Am6(2)A dimethylation in 3'-terminal loop of 18S rRNA, part of 90S and 40S pre-particles in nucleolus, involved in pre-ribosomal RNA processing; human homolog DIMT1 complements yeast dim1 mutant. (318 aa) |
| | SRP68 | Core component of the signal recognition particle (SRP) complex; SRP complex functions in targeting nascent secretory proteins to the endoplasmic reticulum (ER) membrane; relocalizes from cytoplasm to the nuclear periphery upon DNA replication stress; Belongs to the SRP68 family. (599 aa) |
| | BMS1 | Ribosome biogenesis protein BMS1; GTPase required for ribosomal subunit synthesis and rRNA processing; required for synthesis of 40S ribosomal subunits and for processing the 35S pre-rRNA at sites A0, A1, and A2; interacts with Rcl1p, which stimulates its GTPase and U3 snoRNA binding activities; has similarity to Tsr1p; Belongs to the TRAFAC class translation factor GTPase superfamily. Bms1-like GTPase family. BMS1 subfamily. (1183 aa) |
| | NIP7 | Nucleolar protein required for 60S ribosome subunit biogenesis; constituent of 66S pre-ribosomal particles; physically interacts with Nop8p and the exosome subunit Rrp43p. (181 aa) |
| | SRP72 | Core component of the signal recognition particle (SRP); the SRP is a ribonucleoprotein (RNP) complex that functions in targeting nascent secretory proteins to the endoplasmic reticulum (ER) membrane. (640 aa) |
| | HRR25 | Conserved 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) |
| | RPL7B | Ribosomal 60S subunit protein L7B; required for processing of 27SA3 pre-rRNA to 27SB pre-rRNA during assembly of large ribosomal subunit; depletion leads to a turnover of pre-rRNA; contains a conserved C-terminal Nucleic acid Binding Domain (NDB2); binds to Domain II of 25S and 5.8S rRNAs; homologous to mammalian ribosomal protein L7 and bacterial L30; RPL7B has a paralog, RPL7A, that arose from the whole genome duplication. (244 aa) |
| | TGS1 | Trimethyl guanosine synthase, conserved nucleolar methyl transferase; converts the m(7)G cap structure of snRNAs, snoRNAs, and telomerase TLC1 RNA to m(2,2,7)G; also required for nucleolar assembly and splicing of meiotic pre-mRNAs; interacts with Swm2p, which may confer substrate specificity on Tgs1p. (315 aa) |
| | NOP53 | Nucleolar protein; involved in biogenesis of the 60S subunit of the ribosome; interacts with rRNA processing factors Cbf5p and Nop2p and with the nucleolar proteins Nop17p and Nip7p; null mutant is viable but growth is severely impaired; Belongs to the NOP53 family. (455 aa) |
| | NAN1 | NET1-associated nuclear protein 1; U3 snoRNP protein; component of the small (ribosomal) subunit (SSU) processosome containing U3 snoRNA; required for the biogenesis of18S rRNA. (896 aa) |
| | NOG1 | Nucleolar GTP-binding protein 1; Putative GTPase; associates with free 60S ribosomal subunits in the nucleolus and is required for 60S ribosomal subunit biogenesis; constituent of 66S pre-ribosomal particles; member of the ODN family of nucleolar G-proteins. (647 aa) |
| | RPS6A | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S6, no bacterial homolog; phosphorylated on S233 by Ypk3p in a TORC1-dependent manner, and on S232 in a TORC1/2-dependent manner by Ypk1/2/3p; RPS6A has a paralog, RPS6B, that arose from the whole genome duplication. (236 aa) |
| | RPS9A | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S9 and bacterial S4; RPS9A has a paralog, RPS9B, that arose from the whole genome duplication. (197 aa) |
| | NOP4 | Nucleolar protein; essential for processing and maturation of 27S pre-rRNA and large ribosomal subunit biogenesis; constituent of 66S pre-ribosomal particles; contains four RNA recognition motifs (RRMs). (685 aa) |
| | ULP1 | Ubiquitin-like-specific protease 1; Protease that specifically cleaves Smt3p protein conjugates; required for cell cycle progression; associates with nucleoporins and may interact with septin rings during telophase; sequestered to the nucleolus under stress conditions. (621 aa) |
| | RRP12 | Protein required for export of the ribosomal subunits; associates with the RNA components of the pre-ribosomes; has a role in nuclear import in association with Pse1p; also plays a role in the cell cycle and the DNA damage response; contains HEAT-repeats. (1228 aa) |
| | RPA190 | RNA polymerase I largest subunit A190; Belongs to the RNA polymerase beta' chain family. (1664 aa) |
| | RPA43 | RNA polymerase I subunit A43; Belongs to the eukaryotic RPA43 RNA polymerase subunit family. (326 aa) |
| | NOP58 | Nucleolar protein 58; Protein involved in producing mature rRNAs and snoRNAs; involved in pre-rRNA processing, 18S rRNA synthesis, and snoRNA synthesis; component of the small subunit processome complex, which is required for processing of pre-18S rRNA. (511 aa) |
| | UAF30 | Subunit of UAF (upstream activation factor) complex; UAF is an RNA polymerase I specific transcription stimulatory factor composed of Uaf30p, Rrn5p, Rrn9p, Rrn10p, histones H3 and H4; targeting factor for the UAF that facilitates activation of many rDNA genes; deletion decreases cellular growth rate; UAF30 has a paralog, TRI1, that arose from the whole genome duplication. (228 aa) |
| | RRS1 | Regulator of ribosome biosynthesis; Essential protein that binds ribosomal protein L11; required for nuclear export of the 60S pre-ribosomal subunit during ribosome biogenesis; localizes to the nucleolus and in foci along nuclear periphery; cooperates with Ebp2p and Mps3p to mediate telomere clustering by binding Sir4p, but is not involved in telomere tethering; mouse homolog shows altered expression in Huntington's disease model mice; Belongs to the RRS1 family. (203 aa) |
| | RRP36 | rRNA biogenesis protein RRP36; Component of 90S preribosomes; involved in early cleavages of the 35S pre-rRNA and in production of the 40S ribosomal subunit; Belongs to the RRP36 family. (300 aa) |
| | MOD5 | Delta 2-isopentenyl pyrophosphate:tRNA isopentenyl transferase; required for biosynthesis of isopentenyladenosine in mitochondrial and cytoplasmic tRNAs; also has a role in tRNA gene-mediated silencing; gene encodes two isozymic forms; converts to a prion form, prion conversion contributes to azole antifungal resistance by upregulating ergosterol biosynthesis; homolog of human TRIT1, a mutation in which is associated with severe combined respiratory chain defects. (428 aa) |
| | YTM1 | Ribosome biogenesis protein YTM1; Constituent of 66S pre-ribosomal particles; forms a complex with Nop7p and Erb1p that is required for maturation of the large ribosomal subunit; has seven C-terminal WD repeats. (460 aa) |
| | RPB8 | RNA polymerase subunit ABC14.5; common to RNA polymerases I, II, and III. (146 aa) |
| | RPB10 | RNA polymerase subunit ABC10-beta; common to RNA polymerases I, II, and III. (70 aa) |
| | NOC2 | Nucleolar complex protein 2; Protein involved in ribosome biogenesis; forms a nucleolar complex with Mak21p that binds to 90S and 66S pre-ribosomes; forms a nuclear complex with Noc3p that binds to 66S pre-ribosomes; both complexes mediate intranuclear transport of ribosomal precursors; acts as part of a Mak21p-Noc2p-Rrp5p module that associates with nascent pre-rRNA during transcription and has a role in bigenesis of the large ribosomal subunit. (710 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) |
| | PNO1 | Pre-rRNA-processing protein PNO1; Essential nucleolar protein required for pre-18S rRNA processing; interacts with Dim1p, an 18S rRNA dimethyltransferase, and also with Nob1p, which is involved in proteasome biogenesis; contains a KH domain. (274 aa) |
| | RIO1 | Serine/threonine-protein kinase RIO1; Serine kinase involved in cell cycle regulation and rDNA integrity; associated with late pre-40S particles via its conserved C-terminal domain and participates in late 40S biogenesis; association with pre-40S particles regulated by its catalytic ATPase site and likely occurs after the release of Rio2p from these particles; involved in cell cycle progression and processing of the 20S pre-rRNA into mature 18S rRNA; phosphorylates Rpa43p in anaphase to remove Pol I from rDNA. (484 aa) |
| | RPS7A | Protein component of the small (40S) ribosomal subunit; interacts with Kti11p; deletion causes hypersensitivity to zymocin; homologous to mammalian ribosomal protein S7, no bacterial homolog; RPS7A has a paralog, RPS7B, that arose from the whole genome duplication. (190 aa) |
| | BUD21 | Bud site selection protein 21; Component of small ribosomal subunit (SSU) processosome; this complex contains U3 snoRNA; required at post-transcriptional step for efficient retrotransposition; absence results in decreased Ty1 Gag:GFP protein levels; originally isolated as bud-site selection mutant that displays a random budding pattern; Belongs to the UTP16 family. (214 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) |
| | NOB1 | 20S-pre-rRNA D-site endonuclease NOB1; Protein involved in proteasomal and 40S ribosomal subunit biogenesis; required for cleavage of the 20S pre-rRNA to generate the mature 18S rRNA; cleavage is activated by Fun12p, a GTPase and translation initiation factor; relocalizes from nucleus to nucleolus upon DNA replication stress; Belongs to the NOB1 family. (459 aa) |
| | CKB2 | Beta' regulatory 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 polymerase. (258 aa) |
| | UTP23 | rRNA-processing protein UTP23; Component of the small subunit processome; involved in 40S ribosomal subunit biogenesis; interacts with snR30 and is required for dissociation of snR30 from large pre-ribosomal particles; has homology to PINc domain protein Fcf1p, although the PINc domain of Utp23p is not required for function; essential protein. (254 aa) |
| | RRP6 | Nuclear exosome exonuclease component; has 3'-5' exonuclease activity that is regulated by Lrp1p; involved in RNA processing, maturation, surveillance, degradation, tethering, and export; role in sn/snoRNAs precursor degradation; forms a stable heterodimer with Lrp1p; has similarity to E. coli RNase D and to human PM-Sc1 100 (EXOSC10); mutant displays reduced transcription elongation in the G-less-based. (733 aa) |
| | NOP8 | Nucleolar protein required for 60S ribosomal subunit biogenesis. (484 aa) |
| | RRP40 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; predicted to contain both S1 and KH RNA binding domains; mutations in the human homolog, EXOSC3, cause pontocerebellar hypoplasia with motor neuron degeneration. (240 aa) |
| | PAP2 | Non-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) |
| | REX4 | Putative RNA exonuclease; possibly involved in pre-rRNA processing and ribosome assembly. (289 aa) |
| | BRX1 | Ribosome biogenesis protein BRX1; Nucleolar protein; constituent of 66S pre-ribosomal particles; depletion leads to defects in rRNA processing and a block in the assembly of large ribosomal subunits; possesses a sigma(70)-like RNA-binding motif. (291 aa) |
| | NOP12 | Nucleolar protein involved in pre-25S rRNA processing; also involved in biogenesis of large 60S ribosomal subunit; contains an RNA recognition motif (RRM); binds to Ebp2; similar to Nop13p and Nsr1p; Belongs to the RRM RBM34 family. (459 aa) |
| | DIS3 | Exosome core complex catalytic subunit; has both endonuclease and 3'-5' exonuclease activity; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; role in degradation of tRNAs; similar to E. coli RNase R and to human DIS3, which partially complements dis3-81 heat sensitivity; mutations in Dis3p analogous to human mutations implicated in multiple myeloma impair exosome function; protein abundance increases under to DNA replication stress. (1001 aa) |
| | RCL1 | RNA 3'-terminal phosphate cyclase-like protein; Endonuclease that cleaves pre-rRNA at site A2 for 18S rRNA biogenesis; subunit of U3-containing 90S preribosome processome complex involved in small ribosomal subunit assembly; stimulates Bms1p GTPase and U3 binding activity; similar to RNA cyclase-like proteins but no cyclase activity detected. (367 aa) |
| | TOP1 | DNA topoisomerase 1; Topoisomerase I; nuclear enzyme that relieves torsional strain in DNA by cleaving and re-sealing the phosphodiester backbone; relaxes both positively and negatively supercoiled DNA; functions in replication, transcription, and recombination; role in processing ribonucleoside monophosphates in genomic DNA into irreversible single-strand breaks; enzymatic activity and interaction with Nsr1p are negatively regulated by polyphosphorylation. (769 aa) |
| | ESF2 | Pre-rRNA-processing protein ESF2; Essential nucleolar protein involved in pre-18S rRNA processing; binds to RNA and stimulates ATPase activity of Dbp8; involved in assembly of the small subunit (SSU) processome. (316 aa) |
| | NOG2 | Nucleolar GTP-binding protein 2; Putative GTPase; associates with pre-60S ribosomal subunits in the nucleolus and is required for their nuclear export and maturation; recruited by ribosomal proteins L17, L35, and L37 to assembling ribosomes after 27SB pre-rRNA is generated, immediately preceding removal of ITS2; Belongs to the TRAFAC class YlqF/YawG GTPase family. NOG2 subfamily. (486 aa) |
| | TRM112 | Multifunctional methyltransferase subunit TRM112; Protein involved in methylation of tRNA, rRNA, and translation factors; also involved in ribosome biogenesis; subunit of tRNA methyltransferase (MTase) complexes in combination with Trm9p and Trm11p; N7-methylates G1575 of 18S rRNA as complex with Bud23p; subunit of complex with Mtq2p that methylates Sup45p (eRF1) in the ternary complex eRF1-eRF3-GTP; relative distribution to the nucleus increases upon DNA replication stress; functional homolog of human TRMT112. (135 aa) |
| | DBP6 | ATP-dependent RNA helicase DBP6; Essential protein involved in ribosome biogenesis; putative ATP-dependent RNA helicase of the DEAD-box protein family; human homolog DDX51 complements yeast dbp6 mutant. (629 aa) |
| | SWM2 | Nucleolar protein SWM2; Protein with a role in snRNA and snoRNA cap trimethylation; interacts with Tgs1p and shows similar phenotypes; required for trimethylation of the caps of spliceosomal snRNAs and the U3 snoRNA, and for efficient 3' end processing of U3 snoRNA; may act as a specificity factor for Tgs1p; Belongs to the SWM2 family. (146 aa) |
| | KRI1 | Protein KRI1; Essential nucleolar protein required for 40S ribosome biogenesis; associate with snR30; physically and functionally interacts with Krr1p; Belongs to the KRI1 family. (591 aa) |
| | TRF5 | Non-canonical poly(A) polymerase; involved in nuclear RNA degradation as a component of the TRAMP complex; catalyzes polyadenylation of hypomodified tRNAs, and snoRNA and rRNA precursors; overlapping but non-redundant functions with Pap2p. (642 aa) |
| | POP3 | Ribonucleases P/MRP protein subunit POP3; Subunit of both RNase MRP and nuclear RNase P; RNase MRP cleaves pre-rRNA, while nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs; relocalizes to the cytosol in response to hypoxia. (195 aa) |
| | RPA49 | RNA polymerase I subunit A49; essential for nucleolar assembly and for high polymerase loading rate; required for nucleolar localization of Rpa34p. (415 aa) |
| | CSL4 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; predicted to contain an S1 RNA binding domain; human homolog EXOSC1 partially complements yeast csl4 null mutant, and can complement inviability of strain in which expression of CSL4 is repressed. (292 aa) |
| | JJJ1 | J protein JJJ1; Co-chaperone that stimulates the ATPase activity of Ssa1p; required for a late step of ribosome biogenesis; associated with the cytosolic large ribosomal subunit; contains a J-domain; mutation causes defects in fluid-phase endocytosis. (590 aa) |
| | SQS1 | Protein that stimulates the ATPase and helicase activities of Prp43p; acts with Prp43p to stimulate 18s rRNA maturation by Nob1p; overexpression antagonizes the suppression of splicing defects by spp382 mutants; component of pre-ribosomal particles; relocalizes from nucleus to nucleolus upon DNA replication stress; Belongs to the SQS1 family. (767 aa) |
| | POP1 | Ribonucleases P/MRP protein subunit POP1; Subunit of RNase MRP, nuclear RNase P and telomerase complexes; RNase MRP cleaves pre-rRNA, nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs, while telomerase replenishes telomeric DNA; binds to the RPR1 RNA subunit in RNase P. (875 aa) |
| | SLZ1 | Sporulation-specific protein with a leucine zipper motif; subunit of the MIS complex which controls mRNA methylation during during the induction of sporulation. (298 aa) |
| | UBP10 | Ubiquitin carboxyl-terminal hydrolase 10; Ubiquitin-specific protease, deubiquitinates Ub-protein moieties; interacts with proteins that function in rRNA production and ribosome biogenesis via its intrinsically disordered regions; stabilizes Rpa190p by deubiquitination; controls PCNA deubiquitylation; may regulate silencing by acting on Sir4p; involved in posttranscriptionally regulating Gap1p, possibly other transporters; localized to the nucleolus; null mutant phenotypes are functionally complemented by human USP36; Belongs to the peptidase C19 family. (792 aa) |
| | NOP13 | Nucleolar protein found in preribosomal complexes; contains an RNA recognition motif (RRM); relative distribution to the nucleolus increases upon DNA replication stress. (403 aa) |
| | LSM7 | U6 snRNA-associated Sm-like protein LSm7; Lsm (Like Sm) protein; part of heteroheptameric complexes (Lsm2p-7p and either Lsm1p or 8p): cytoplasmic Lsm1p complex involved in mRNA decay; nuclear Lsm8p complex part of U6 snRNP and possibly involved in processing tRNA, snoRNA, and rRNA; protein abundance increases and forms cytoplasmic foci in response to DNA replication stress. (115 aa) |
| | KRE33 | RNA cytidine acetyltransferase; Protein required for biogenesis of the small ribosomal subunit; heterozygous mutant shows haploinsufficiency in K1 killer toxin resistance; essential gene; NAT10, the human homolog, implicated in several types of cancer and premature aging. (1056 aa) |
| | RPC19 | RNA polymerase subunit AC19; common to RNA polymerases I and III. (142 aa) |
| | NOP15 | Ribosome biogenesis protein 15; Constituent of 66S pre-ribosomal particles; involved in 60S ribosomal subunit biogenesis; localizes to both nucleolus and cytoplasm. (220 aa) |
| | RPS7B | Protein component of the small (40S) ribosomal subunit; interacts with Kti11p; deletion causes hypersensitivity to zymocin; homologous to mammalian ribosomal protein S7, no bacterial homolog; RPS7B has a paralog, RPS7A, that arose from the whole genome duplication; protein abundance increases in response to DNA replication stress. (190 aa) |
| | IMP4 | U3 small nucleolar ribonucleoprotein protein IMP4; Component of the SSU processome; SSU processome is required for pre-18S rRNA processing; interacts with Mpp10p; member of a superfamily of proteins that contain a sigma(70)-like motif and associate with RNAs. (290 aa) |
| | RPL16B | Ribosomal 60S subunit protein L16B; N-terminally acetylated, binds 5.8 S rRNA; transcriptionally regulated by Rap1p; homologous to mammalian ribosomal protein L13A and bacterial L13; RPL16B has a paralog, RPL16A, that arose from the whole genome duplication. (198 aa) |
| | NOP2 | rRNA m5C methyltransferase; methylates cytosine at position 2870 of 25S rRNA; has an essential function independent of rRNA methylation; contains seven beta-strand methyltransferase motif; essential for processing and maturation of 27S pre-rRNA and large ribosomal subunit biogenesis; localized to the nucleolus; constituent of 66S pre-ribosomal particles; rRNA methylation defect and lethality are functionally complemented by human NOP2, a gene upregulated in cancer; Belongs to the class I-like SAM-binding methyltransferase superfamily. RsmB/NOP family. (618 aa) |
| | RCM1 | rRNA m5C methyltransferase; methylates cytosine at position 2278 of 25S rRNA while Nop2p methylates cytosine at position 2870; contains seven beta-strand methyltransferase motif; localized to the nucleolus; interacts with Trm112p; homolog of NSUN5A, a human gene which is deleted in Williams-Beuren Syndrome. (490 aa) |
| | RLP7 | Ribosome biogenesis protein RLP7; Nucleolar protein similar to large ribosomal subunit L7 proteins; constituent of 66S pre-ribosomal particles; plays an essential role in processing of precursors to the large ribosomal subunit RNAs; binds junction of ITS2 and ITS2-proximal stem between the 3' end of 5.8S rRNA and the 5' end of 25S rRNA. (322 aa) |
| | HAS1 | ATP-dependent RNA helicase; involved in the biogenesis of 40S and 60S ribosome subunits; localizes to both the nuclear periphery and nucleolus; highly enriched in nuclear pore complex fractions; constituent of 66S pre-ribosomal particles. (505 aa) |
| | RRN9 | RNA polymerase I-specific transcription initiation factor RRN9; Protein involved in promoting high level transcription of rDNA; subunit of UAF (upstream activation factor) for RNA polymerase I. (365 aa) |
| | TMA23 | Nucleolar protein implicated in ribosome biogenesis; deletion extends chronological lifespan. (211 aa) |
| | RNT1 | Ribonuclease 3; Nuclear dsRNA-specific ribonuclease (RNase III); involved in rDNA transcription, rRNA processing and U2 snRNA 3' end formation by cleavage of a stem-loop structure at the 3' end of U2 snRNA; involved in polyadenylation-independent transcription termination; involved in the cell wall stress response, regulating the degradation of cell wall integrity and morphogenesis checkpoint genes. (471 aa) |
| | TRI1 | Protein TRI1; Non-essential sumoylated protein of unknown function; similar to components of human SWI/SNF complex including SMRD3; green fluorescent protein (GFP)-fusion protein localizes to the cytoplasm, nucleus and nucleolus; TRI1 has a paralog, UAF30, that arose from the whole genome duplication. (226 aa) |
| | RRP5 | rRNA biogenesis protein RRP5; RNA binding protein involved in synthesis of 18S and 5.8S rRNAs; component of ribosomal small subunit (SSU) processome and 90S preribosome; required for pre-rRNA packaging and compaction of processome into dense terminal balls; part of Mak21p-Noc2p-Rrp5p module that associates with nascent pre-rRNA during transcription with role in biogenesis of large ribosomal subunit; binds single stranded tracts of U's; relocalizes from nucleolus to nucleus upon DNA replication stress. (1729 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) |
| | RRB1 | Ribosome assembly protein RRB1; Specific chaperone for ribosomal protein Rpl3p; binds to nascent Rpl3p during translation; essential gene. (511 aa) |
| | ECM16 | Essential DEAH-box ATP-dependent RNA helicase specific to U3 snoRNP; predominantly nucleolar in distribution; required for 18S rRNA synthesis. (1267 aa) |
| | RPL15B | Ribosomal 60S subunit protein L15B; binds to 5.8 S rRNA; homologous to mammalian ribosomal protein L15, no bacterial homolog; RPL15B has a paralog, RPL15A, that arose from the whole genome duplication; relocalizes from nucleus to nucleolus upon DNA replication stress. (204 aa) |
| | UTP15 | Nucleolar protein; component of the small subunit (SSU) processome containing the U3 snoRNA that is involved in processing of pre-18S rRNA. (513 aa) |
| | ERB1 | Ribosome biogenesis protein ERB1; Constituent of 66S pre-ribosomal particles; forms a complex with Nop7p and Ytm1p that is required for maturation of the large ribosomal subunit; required for maturation of the 25S and 5.8S ribosomal RNAs; homologous to mammalian Bop1. (807 aa) |
| | BUD22 | Bud site selection protein 22; Protein required for rRNA maturation and ribosomal subunit biogenesis; required for 18S rRNA maturation; also required for small ribosomal subunit biogenesis; cosediments with pre-ribosomal particles; mutation decreases efficiency of +1 Ty1 frameshifting and transposition, and affects budding pattern. (519 aa) |
| | CTK3 | Gamma subunit of C-terminal domain kinase I; CTDK-I phosphorylates RNA polymerase II subunit Rpo21p to affect transcription and pre-mRNA 3' end processing, and also phosphorylates ribosomal protein Rps2p to increase translational fidelity; protein abundance increases in response to DNA replication stress. (296 aa) |
| | UTP14 | U3 small nucleolar RNA-associated protein 14; Subunit of U3-containing Small Subunit (SSU) processome complex; involved in production of 18S rRNA and assembly of small ribosomal subunit; Belongs to the UTP14 family. (899 aa) |
| | FPR3 | FK506-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) |
| | PIF1 | DNA helicase, potent G-quadruplex DNA binder/unwinder; possesses strand annealing activity; promotes DNA synthesis during break-induced replication; important for crossover recombination; translation from different start sites produces mitochondrial and nuclear forms; nuclear form is a catalytic inhibitor of telomerase; mitochondrial form involved in DNA repair and recombination; mutations affect Zn, Fe homeostasis; regulated by Rad53p-dependent phosphorylation in rho0 cells. (859 aa) |
| | RRN11 | RNA polymerase I-specific transcription initiation factor RRN11; Component of the core factor (CF) rDNA transcription factor complex; CF is required for transcription of 35S rRNA genes by RNA polymerase I and is composed of Rrn6p, Rrn7p, and Rrn11p. (507 aa) |
| | FPR4 | FK506-binding protein 4; Peptidyl-prolyl cis-trans isomerase (PPIase); nuclear proline isomerase; affects expression of multiple genes via its role in nucleosome assembly; catalyzes isomerization of proline residues in histones H3 and H4, which affects lysine methylation of those histones; PPIase domain acts as a transcriptional repressor when tethered to DNA by lexA, and repressor activity is dependent on PPIase activity; contains a nucleoplasmin-like fold and can form pentamers; Belongs to the FKBP-type PPIase family. FKBP3/4 subfamily. (392 aa) |
| | SIR3 | Regulatory protein SIR3; Silencing protein; interacts with Sir2p, Sir4p, and histone H3/H4 tails to establish transcriptionally silent chromatin; required for spreading of silenced chromatin; recruited to chromatin through interaction with Rap1p; C-terminus assumes variant winged helix-turn-helix (wH) fold that mediates homodimerization, which is critical for holo-SIR complex loading; required for telomere hypercluster formation in quiescent yeast cells; has paralog ORC1 from whole genome duplication. (978 aa) |
| | LSM3 | U6 snRNA-associated Sm-like protein LSm3; Lsm (Like Sm) protein; part of heteroheptameric complexes (Lsm2p-7p and either Lsm1p or 8p): cytoplasmic Lsm1p complex involved in mRNA decay; nuclear Lsm8p complex part of U6 snRNP and possibly involved in processing tRNA, snoRNA, and rRNA; protein increases in abundance and relocalizes from nucleus to cytoplasmic foci upon DNA replication stress. (89 aa) |
| | UTP21 | U3 small nucleolar RNA-associated protein 21; Subunit of U3-containing 90S preribosome and SSU processome complexes; involved in production of 18S rRNA and assembly of small ribosomal subunit; synthetic defect with STI1 Hsp90 cochaperone; human homolog linked to glaucoma; Small Subunit processome is also known as SSU processome. (939 aa) |
| | SGD1 | Suppressor of glycerol defect protein 1; Essential nuclear protein; required for biogenesis of the small ribosomal subunit; has a possible role in the osmoregulatory glycerol response; putative homolog of human NOM1 which is implicated in acute myeloid leukemia. (899 aa) |
| | EST2 | Reverse transcriptase subunit of the telomerase holoenzyme; essential for telomerase core catalytic activity, involved in other aspects of telomerase assembly and function; mutations in human homolog are associated with aplastic anemia. (884 aa) |
| | DBP9 | ATP-dependent RNA helicase DBP9; DEAD-box protein required for 27S rRNA processing; exhibits DNA, RNA and DNA/RNA helicase activities; ATPase activity shows preference for DNA over RNA; DNA helicase activity abolished by mutation in RNA-binding domain; Belongs to the DEAD box helicase family. DDX56/DBP9 subfamily. (594 aa) |
| | YCS4 | Subunit 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) |
| | EST1 | Telomere elongation protein EST1; TLC1 RNA-associated factor involved in telomere length regulation; recruitment subunit of telomerase; has G-quadruplex promoting activity required for telomere elongation; role in activating telomere-bound Est2p-TLC1-RNA; EST1 has a paralog, EBS1, that arose from the whole genome duplication. (699 aa) |
| | IFH1 | Protein IFH1; Coactivator, regulates transcription of ribosomal protein (RP) genes; recruited to RP gene promoters during optimal growth conditions via Fhl1p; subunit of CURI, a complex that coordinates RP production and pre-rRNA processing; regulated by acetylation and phosphorylation at different growth states via TORC1 signaling; IFH1 has a paralog, CRF1, that arose from the whole genome duplication. (1085 aa) |
| | UTP13 | Nucleolar protein; component of the small subunit (SSU) processome containing the U3 snoRNA that is involved in processing of pre-18S rRNA. (817 aa) |
| | RSA3 | Ribosome assembly protein 3; Protein with a likely role in ribosomal maturation; required for accumulation of wild-type levels of large (60S) ribosomal subunits; binds to the helicase Dbp6p in pre-60S ribosomal particles in the nucleolus. (220 aa) |
| | NOP56 | Nucleolar protein 56; Essential evolutionarily-conserved nucleolar protein; component of the box C/D snoRNP complexes that direct 2'-O-methylation of pre-rRNA during its maturation; overexpression causes spindle orientation defects. (504 aa) |
| | PWP1 | Periodic tryptophan protein 1; Protein with WD-40 repeats involved in rRNA processing; associates with trans-acting ribosome biogenesis factors; similar to beta-transducin superfamily; Belongs to the WD repeat PWP1 family. (576 aa) |
| | EMG1 | Ribosomal RNA small subunit methyltransferase NEP1; Methyltransferase for rRNA; methylates pseudouridine 18S rRNA residue 1191; member of the SPOUT methyltransferase family; required for maturation of 18S rRNA and for 40S ribosomal subunit production independent of methyltransferase activity; forms homodimers; human ortholog is mutated in Bowen-Conradi syndrome, and equivalent yeast mutation affects Emg1p dimerization and localization but not methyltransferase activity; human EMG1 complements lethality of null and ts mutant; Belongs to the class IV-like SAM-binding methyltransferase su [...] (252 aa) |
| | CBF5 | H/ACA ribonucleoprotein complex subunit CBF5; Pseudouridine synthase catalytic subunit of box H/ACA snoRNPs; acts on large and small rRNAs, on snRNA U2, and on some mRNAs; mutations in human ortholog dyskerin cause the disorder dyskeratosis congenita; small nucleolar ribonucleoprotein particles are also known as snoRNPs. (483 aa) |
| | ACS2 | Acetyl-coA synthetase isoform; along with Acs1p, acetyl-coA synthetase isoform is the nuclear source of acetyl-coA for histone acetylation; mutants affect global transcription; required for growth on glucose; expressed under anaerobic conditions; Belongs to the ATP-dependent AMP-binding enzyme family. (683 aa) |
| | RMP1 | Ribonuclease MRP protein subunit RMP1; Subunit of RNase MRP; RNase MRP processes pre-rRNA and has a role in cell cycle-regulated degradation of daughter cell-specific mRNAs; unlike most subunits, not shared between RNase MRP and nuclear RNase P. (201 aa) |
| | RRN5 | Protein involved in transcription of rDNA by RNA polymerase I; transcription factor, member of UAF (upstream activation factor) family along with Rrn9p and Rrn10p. (363 aa) |
| | DIP2 | Nucleolar protein; specifically associated with the U3 snoRNA, part of the large ribonucleoprotein complex known as the small subunit (SSU) processome, required for 18S rRNA biogenesis, part of the active pre-rRNA processing complex. (943 aa) |
| | MDN1 | Midasin; Huge dynein-related AAA-type ATPase (midasin); forms extended pre-60S particle with the Rix1 complex (Rix1p-Ipi1p-Ipi3p); acts in removal of ribosomal biogenesis factors at successive steps of pre-60S assembly and export from nucleus. (4910 aa) |
| | FYV7 | rRNA-processing protein FYV7; Essential protein required for maturation of 18S rRNA; required for survival upon exposure to K1 killer toxin. (151 aa) |
| | FCF2 | rRNA-processing protein FCF2; Nucleolar protein involved in the early steps of 35S rRNA processing; interacts with Faf1p; member of a transcriptionally co-regulated set of genes called the RRB regulon; essential gene. (217 aa) |
| | RLP24 | Essential protein required for ribosomal large subunit biogenesis; associated with pre-60S ribosomal subunits; stimulates the ATPase activity of Afg2p, which is required for release of Rlp24p from the pre-60S particle; has similarity to Rpl24Ap and Rpl24Bp. (199 aa) |
| | NOC3 | Nucleolar complex-associated protein 3; Subunit of a nuclear complex with Noc2p and pre-replicative complexes; the Noc2p-Noc3p complex binds to 66S ribosomal precursors to mediate their maturation and intranuclear transport; binds to chromatin at active replication origins, and is required for pre-RC formation and maintenance during DNA replication licensing. (663 aa) |
| | GRC3 | Polynucleotide 5'-hydroxyl-kinase GRC3; Polynucleotide kinase present on rDNA; required for efficient transcription termination by RNA polymerase I; functions with Las1p in a conserved mechanism to modulate rRNA processing and ribosome biogenesis; required for cell growth; mRNA is cell-cycle regulated; Belongs to the Clp1 family. NOL9/GRC3 subfamily. (632 aa) |
| | RIX7 | Ribosome biogenesis ATPase RIX7; Putative ATPase of the AAA family; required for export of pre-ribosomal large subunits from the nucleus; distributed between the nucleolus, nucleoplasm, and nuclear periphery depending on growth conditions. (837 aa) |
| | SOF1 | Protein required for biogenesis of 40S (small) ribosomal subunit; has similarity to the beta subunit of trimeric G-proteins and the splicing factor Prp4p; essential gene; Belongs to the WD repeat DCAF13/WDSOF1 family. (489 aa) |
| | DRS1 | ATP-dependent RNA helicase DRS1; Nucleolar DEAD-box protein required for ribosome assembly and function; including synthesis of 60S ribosomal subunits; constituent of 66S pre-ribosomal particles; Belongs to the DEAD box helicase family. DDX27/DRS1 subfamily. (752 aa) |
| | SRP40 | Suppressor protein SRP40; Nucleolar serine-rich protein; role in preribosome assembly or transport; may function as a chaperone of small nucleolar ribonucleoprotein particles (snoRNPs); immunologically and structurally to rat Nopp140. (406 aa) |
| | RPF2 | Ribosome biogenesis protein RPF2; Essential protein involved in rRNA maturation and ribosomal assembly; involved in the processing of pre-rRNA and the assembly of the 60S ribosomal subunit; interacts with ribosomal protein L11; localizes predominantly to the nucleolus; constituent of 66S pre-ribosomal particles. (344 aa) |
| | LAS1 | Protein LAS1; Endonuclease involved in pre-rRNA processing at both ends of ITS2; functions with Grc3p in a conserved mechanism to modulate rRNA processing and ribosome biogenesis; may coordinate the action of the Rat1p-Rai1p exoRNAse; required for the G1/S cell cycle transition; human ortholog is Las1L; mutants require the SSD1-v allele for viability. (502 aa) |
| | UTP30 | Ribosome biogenesis protein UTP30; Putative subunit of U3-containing 90S preribosome complex; complex is involved in production of 18S rRNA and assembly of small ribosomal subunit; Belongs to the universal ribosomal protein uL1 family. Highly divergent. (274 aa) |
| | DBP7 | Putative ATP-dependent RNA helicase of the DEAD-box family; involved in ribosomal biogenesis; required at post-transcriptional step for efficient retrotransposition; essential for growth under anaerobic conditions. (742 aa) |
| | TOF2 | Topoisomerase 1-associated factor 2; Protein required for rDNA silencing and mitotic rDNA condensation; stimulates Cdc14p phosphatase activity and biphasic release to promote rDNA repeat segregation; required for condensin recruitment to the replication fork barrier site; TOF2 has a paralog, NET1, that arose from the whole genome duplication; To yeast YJL076w. (771 aa) |
| | EBP2 | rRNA-processing protein EBP2; Required for 25S rRNA maturation and 60S ribosomal subunit assembly; localizes to the nucleolus and in foci along nuclear periphery; constituent of 66S pre-ribosomal particles; cooperates with Rrs1p and Mps3p to mediate telomere clustering by binding Sir4p, but is not involved in telomere tethering. (427 aa) |
| | CTK1 | Catalytic (alpha) subunit of C-terminal domain kinase I (CTDK-I); phosphorylates both RNA pol II subunit Rpo21p to affect transcription and pre-mRNA 3' end processing, and ribosomal protein Rps2p to increase translational fidelity; required for H3K36 trimethylation but not dimethylation by Set2p; suggested stimulatory role in 80S formation during translation initiation; similar to the Drosophila dCDK12 and human CDK12 and probably CDK13; Belongs to the protein kinase superfamily. CMGC Ser/Thr protein kinase family. CDC2/CDKX subfamily. (528 aa) |
| | RRN3 | Protein required for transcription of rDNA by RNA polymerase I; transcription factor independent of DNA template; involved in recruitment of RNA polymerase I to rDNA; structure reveals unique HEAT repeat fold and a surface serine patch; phosphorylation of serine patch impairs cell growth and reduces RNA polymerase I binding in vitro and RNA polymerase I recruitment to the rDNA gene in vivo; Belongs to the RRN3 family. (627 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) |
| | UTP11 | U3 small nucleolar RNA-associated protein 11; Subunit of U3-containing Small Subunit (SSU) processome complex; involved in production of 18S rRNA and assembly of small ribosomal subunit. (250 aa) |
| | RRP14 | Essential protein, constituent of 66S pre-ribosomal particles; interacts with proteins involved in ribosomal biogenesis and cell polarity; member of the SURF-6 family; Belongs to the SURF6 family. (434 aa) |
| | DHR2 | Predominantly nucleolar DEAH-box ATP-dependent RNA helicase; required for 18S rRNA synthesis; Belongs to the DEAD box helicase family. DEAH subfamily. (735 aa) |
| | MAK11 | Protein involved in an early step of 60S ribosomal subunit biogenesis; essential for cell growth and replication of killer M1 dsRNA virus; contains four beta-transducin repeats. (468 aa) |
| | URB1 | Protein required for the normal accumulation of 25S and 5.8S rRNAs; nucleolar protein; associated with the 27SA2 pre-ribosomal particle; proposed to be involved in the biogenesis of the 60S ribosomal subunit. (1764 aa) |
| | MRT4 | Protein involved in mRNA turnover and ribosome assembly; required at post-transcriptional step for efficient retrotransposition; localizes to the nucleolus; Belongs to the universal ribosomal protein uL10 family. (236 aa) |
| | RPL14A | Ribosomal 60S subunit protein L14A; N-terminally acetylated; homologous to mammalian ribosomal protein L14, no bacterial homolog; RPL14A has a paralog, RPL14B, that arose from the whole genome duplication. (138 aa) |
| | RPA12 | RNA polymerase I subunit A12.2; contains two zinc binding domains, and the N terminal domain is responsible for anchoring to the RNA pol I complex; physically interacts with transcriptional activator Msn4p, to regulate transcription of AYR1, a gene involved in lipid metabolism; Belongs to the archaeal RpoM/eukaryotic RPA12/RPB9/RPC11 RNA polymerase family. (125 aa) |
| | URB2 | Protein required for normal metabolism of the rRNA primary transcript; nucleolar protein; proposed to be involved in ribosome biogenesis. (1174 aa) |
| | LSM8 | U6 snRNA-associated Sm-like protein LSm8; Lsm (Like Sm) protein; forms heteroheptameric complex (with Lsm2p, Lsm3p, Lsm4p, Lsm5p, Lsm6p, and Lsm7p) that is part of spliceosomal U6 snRNP and is also implicated in processing of pre-tRNA, pre-snoRNA, and pre-rRNA. (109 aa) |
| | MPP10 | U3 small nucleolar RNA-associated protein MPP10; Component of the SSU processome and 90S preribosome; required for pre-18S rRNA processing, interacts with and controls the stability of Imp3p and Imp4p, essential for viability; similar to human Mpp10p. (593 aa) |
| | RPS14B | Protein component of the small (40S) ribosomal subunit; required for ribosome assembly and 20S pre-rRNA processing; mutations confer cryptopleurine resistance; homologous to mammalian ribosomal protein S14 and bacterial S11; RPS14B has a paralog, RPS14A, that arose from the whole genome duplication. (138 aa) |
| | RPA34 | RNA polymerase I subunit A34.5; essential for nucleolar assembly and for high polymerase loading rate; nucleolar localization depends on Rpa49p; Belongs to the eukaryotic RPA34 RNA polymerase subunit family. (233 aa) |
| | UTP10 | Nucleolar protein; component of the small subunit (SSU) processome containing the U3 snoRNA that is involved in processing of pre-18S rRNA; mutant has increased aneuploidy tolerance. (1769 aa) |
| | NET1 | Nucleolar protein NET1; Core subunit of the RENT complex; involved in nucleolar silencing and telophase exit; stimulates transcription by RNA polymerase I and regulates nucleolar structure; NET1 has a paralog, TOF2, that arose from the whole genome duplication; To yeast YKR010c. (1189 aa) |
| | UTP18 | Small-subunit processome protein involved in pre-18S rRNA maturation; part of a subunit of the 90S preribosomal particle capable of interacting directly with the 5' ETS of the 35S pre-rRNA; contains WD40 repeats. (594 aa) |
| | MTR4 | ATP-dependent 3'-5' RNA helicase of the DExD/H family; involved in nuclear RNA processing and degradation both as a component of TRAMP complex and in TRAMP-independent processes; TRAMP unwinds RNA duplexes, with Mtr4p unwinding activity stimulated by Pap2p/Air2p but not dependent on ongoing polyadenylation; contains an arch domain, with two coiled-coil arms/stalks and a globular fist/KOW domain, which has RNA binding activity and is required for 5.8S rRNA processing; Belongs to the helicase family. SKI2 subfamily. (1073 aa) |
| | HCA4 | DEAD box RNA helicase; component of the SSU; interacts with Bfr2p and Enp2p; high-copy number suppression of a U14 snoRNA processing mutant suggests an involvement in 18S rRNA synthesis; Belongs to the DEAD box helicase family. DDX10/DBP4 subfamily. (770 aa) |
| | RRN7 | RNA polymerase I-specific transcription initiation factor RRN7; Component of the core factor (CF) rDNA transcription factor complex; CF is required for transcription of 35S rRNA genes by RNA polymerase I and is composed of Rrn6p, Rrn7p, and Rrn11p; Belongs to the RRN7/TAF1B family. (514 aa) |
| | NOP9 | Nucleolar protein 9; Essential subunit of U3-containing 90S preribosome; involved in production of 18S rRNA and assembly of small ribosomal subunit; also part of pre-40S ribosome and required for its export into cytoplasm; binds RNA and contains pumilio domain; Belongs to the NOP9 family. (666 aa) |
| | CTK2 | Beta subunit of C-terminal domain kinase I (CTDK-I); which phosphorylates both RNA pol II subunit Rpo21p to affect transcription and pre-mRNA 3' end processing, and ribosomal protein Rps2p to increase translational fidelity; relocalizes to the cytosol in response to hypoxia. (323 aa) |
| | RPR2 | Ribonuclease P protein subunit RPR2; Subunit of nuclear RNase P; nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs; not shared between RNase MRP and RNase P, in contrast to all other RNase P protein subunits; protein abundance increases in response to DNA replication stress. (144 aa) |
| | RPL16A | Ribosomal 60S subunit protein L16A; N-terminally acetylated, binds 5.8 S rRNA; transcriptionally regulated by Rap1p; homologous to mammalian ribosomal protein L13A and bacterial L13; RPL16A has a paralog, RPL16B, that arose from the whole genome duplication; protein abundance increases in response to DNA replication stress. (199 aa) |
| | RRT14 | Regulator of rDNA transcription protein 14; Putative protein of unknown function; identified in a screen for mutants with decreased levels of rDNA transcription; green fluorescent protein (GFP)-fusion protein localizes to the nucleolus; predicted to be involved in ribosome biogenesis. (206 aa) |
| | BMT5 | Methyltransferase required for m3U2634 methylation of the 25S rRNA; S-adenosylmethionine-dependent; associates with precursors of the 60S ribosomal subunit; predicted to be involved in ribosome biogenesis; Belongs to the class I-like SAM-binding methyltransferase superfamily. BMT5 family. (336 aa) |
| | UTP25 | U3 small nucleolar RNA-associated protein 25; Nucleolar protein; required for 35S pre-RNA processing and 40S ribosomal subunit biogenesis. (721 aa) |
| | AIR1 | Zinc knuckle protein; involved in nuclear RNA processing and degradation as a component of the TRAMP complex; stimulates the poly(A) polymerase activity of Pap2p in vitro; AIR1 has a paralog, AIR2, that arose from the whole genome duplication; although Air1p and Air2p are homologous TRAMP subunits, they have nonredundant roles in regulation of substrate specificity of the exosome. (360 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) |
| | FAF1 | Protein required for pre-rRNA processing; also required for 40S ribosomal subunit assembly. (346 aa) |
| | UTP9 | Nucleolar protein; component of the small subunit (SSU) processome containing the U3 snoRNA that is involved in processing of pre-18S rRNA. (575 aa) |
| | DBP8 | ATPase, putative RNA helicase of the DEAD-box family; component of 90S preribosome complex involved in production of 18S rRNA and assembly of 40S small ribosomal subunit; ATPase activity stimulated by association with Esf2p; Belongs to the DEAD box helicase family. DDX49/DBP8 subfamily. (431 aa) |
| | SPO12 | Sporulation-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) |
| | IMP3 | U3 small nucleolar ribonucleoprotein protein IMP3; Component of the SSU processome; SSU processome is required for pre-18S rRNA processing, essential protein that interacts with Mpp10p and mediates interactions of Imp4p and Mpp10p with U3 snoRNA; Belongs to the universal ribosomal protein uS4 family. (183 aa) |
| | RPC10 | RNA polymerase subunit ABC10-alpha, found in RNA pol I, II, and III; relocalizes from nucleolus to cytoplasm upon DNA replication stress. (70 aa) |
| | GAR1 | H/ACA ribonucleoprotein complex subunit GAR1; Protein component of the H/ACA snoRNP pseudouridylase complex; involved in the modification and cleavage of the 18S pre-rRNA; Belongs to the GAR1 family. (205 aa) |
| | RPF1 | Ribosome production factor 1; Protein involved in assembly and export of the large ribosomal subunit; nucleolar protein; constituent of 66S pre-ribosomal particles; contains a sigma(70)-like motif, which is thought to bind RNA. (295 aa) |
| | LRP1 | Nuclear exosome-associated nucleic acid binding protein; involved in RNA processing, surveillance, degradation, tethering, and export; forms a stable heterodimer with Rrp6p and regulates its exonucleolytic activity; rapidly degraded by the proteasome in the absence of Rrp6p; homolog of mammalian nuclear matrix protein C1D involved in regulation of DNA repair and recombination. (184 aa) |
| | NOP10 | Subunit of box H/ACA snoRNP complex; required for pseudouridylation and processing of pre-18S rRNA. (58 aa) |
| | RRP4 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; predicted to contain RNA binding domains; has similarity to human hRrp4p (EXOSC2). (359 aa) |
| | SSF1 | Ribosome biogenesis protein SSF1; Constituent of 66S pre-ribosomal particles; required for ribosomal large subunit maturation; functionally redundant with Ssf2p; member of the Brix family; SSF1 has a paralog, SSF2, that arose from the whole genome duplication. (453 aa) |
| | RRP3 | ATP-dependent rRNA helicase RRP3; Protein involved in rRNA processing; required for maturation of the 35S primary transcript of pre-rRNA and for cleavage leading to mature 18S rRNA; homologous to eIF-4a, which is a DEAD box RNA-dependent ATPase with helicase activity. (501 aa) |
| | RPP1 | Ribonuclease P/MRP protein subunit RPP1; Subunit of both RNase MRP and nuclear RNase P; RNase MRP cleaves pre-rRNA, while nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs; relocalizes to the cytosol in response to hypoxia; Belongs to the eukaryotic/archaeal RNase P protein component 3 family. (293 aa) |
| | CIC1 | Proteasome-interacting protein CIC1; Essential protein that interacts with proteasome components; has a potential role in proteasome substrate specificity; also copurifies with 66S pre-ribosomal particles. (376 aa) |
| | SBP1 | Single-stranded nucleic acid-binding protein; Protein that binds eIF4G and has a role in repression of translation; has an RGG motif; found in cytoplasmic P bodies; binds to mRNAs under glucose starvation stress, most often in the 5' UTR; found associated with small nucleolar RNAs snR10 and snR11; SBP1 has a paralog, RNP1, that arose from the whole genome duplication; Belongs to the RRM GAR family. (294 aa) |
| | SHU1 | Suppressor of HU sensitivity involved in recombination protein 1; Component of Shu complex (aka PCSS complex); Shu complex also includes Psy3, Csm2, Shu2, and promotes error-free DNA repair, mediates inhibition of Srs2p function; essential for promoting the establishment of homolog bias during meiotic homologous recombination; promotes both crossover (CO) and non-crossover (NCO) pathways of meiotic recombination and formation of Rad51p filaments. (150 aa) |
| | RPL14B | Ribosomal 60S subunit protein L14B; homologous to mammalian ribosomal protein L14, no bacterial homolog; RPL14B has a paralog, RPL14A, that arose from the whole genome duplication; protein abundance increases in response to DNA replication stress. (138 aa) |
| | ZUO1 | Zuotin; Ribosome-associated chaperone; zuotin functions in ribosome biogenesis and as a chaperone for nascent polypeptide chains in partnership with Ssz1p and SSb1/2; contains a DnaJ domain and functions as a J-protein partner for Ssb1p and Ssb2p; human gene DNAJC2 can partially complement yeast zuo1 null mutant. (433 aa) |
| | YGR283C | Putative methyltransferase; may interact with ribosomes, based on co-purification experiments; predicted to be involved in ribosome biogenesis; null mutant is resistant to fluconazole; GFP-fusion protein localizes to the nucleolus; YGR283C has a paralog, YMR310C, that arose from the whole genome duplication; Belongs to the class IV-like SAM-binding methyltransferase superfamily. (341 aa) |
| | PXR1 | Essential protein involved in rRNA and snoRNA maturation; competes with TLC1 RNA for binding to Est2p, suggesting a role in negative regulation of telomerase; human homolog inhibits telomerase; contains a G-patch RNA interacting domain. (271 aa) |
| | EFG1 | rRNA-processing protein EFG1; Essential protein required for maturation of 18S rRNA; null mutant is sensitive to hydroxyurea and is delayed in recovering from alpha-factor arrest; green fluorescent protein (GFP)-fusion protein localizes to the nucleolus. (233 aa) |
| | NOP19 | Nucleolar protein 19; Ribosome biogenesis factor; nucleolar protein associated with pre-rRNA components of the 90S preribosome, required for cleavage of pre-rRNA at A0, A1 and A2 sites; interacts with RNA helicase Dhr2p and RNA helicase-like protein Utp25p; required for incorporation of Utp25p into preribosomes. (196 aa) |
| | SDA1 | Protein required for actin organization and passage through Start; highly conserved nuclear protein; required for actin cytoskeleton organization; plays a critical role in G1 events; binds Nap1p; involved in 60S ribosome biogenesis; Belongs to the SDA1 family. (767 aa) |
| | SKI6 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; has similarity to E. coli RNase PH and to human hRrp41p (EXOSC4). (246 aa) |
| | NSR1 | Nucleolar protein that binds nuclear localization sequences; required for pre-rRNA processing and ribosome biogenesis; binds to single stranded telomeric DNA and mRNA; methylated by Hmt1p; interaction with Top1p and nucleolar localization are negatively regulated by polyphosphorylation. (414 aa) |
| | MTR3 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; has similarity to E. coli RNase PH and to human hMtr3p (EXOSC6). (250 aa) |
| | ENP2 | Ribosome biogenesis protein ENP2; Component of the SSU; required for pre-18S rRNA processing, biogenesis of the small ribosomal subunit; interacts with U3 snoRNA, Mpp10p and Bfr2p; contains WD repeats, and has homology to Spb1p. (707 aa) |
| | UTP8 | U3 small nucleolar RNA-associated protein 8; Nucleolar protein required for export of tRNAs from the nucleus; also copurifies with the small subunit (SSU) processome containing the U3 snoRNA that is involved in processing of pre-18S rRNA. (713 aa) |
| | NOP7 | Pescadillo homolog; Component of several different pre-ribosomal particles; forms a complex with Ytm1p and Erb1p that is required for maturation of the large ribosomal subunit; required for exit from G<sub>0</sub> and the initiation of cell proliferation. (605 aa) |
| | RRP46 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; has similarity to E. coli RNase PH and to human hRrp46p (EXOSC5). (223 aa) |
| | UTP22 | U3 small nucleolar RNA-associated protein 22; Component of the small-subunit processome; required for nuclear export of tRNAs; may facilitate binding of Utp8p to aminoacylated tRNAs and their delivery to Los1p for export; conserved from yeast to mammals. (1237 aa) |
| | SLX9 | Ribosome biogenesis protein SLX9; Protein required for pre-rRNA processing; associated with the 90S pre-ribosome and 43S small ribosomal subunit precursor; interacts with U3 snoRNA; deletion mutant has synthetic fitness defect with an sgs1 deletion mutant; Belongs to the SLX9 family. (210 aa) |
| | UPF3 | Component of the nonsense-mediated mRNA decay (NMD) pathway; along with Nam7p and Nmd2p; involved in decay of mRNA containing nonsense codons; involved in telomere maintenance; Belongs to the RENT3 family. (387 aa) |
| | POP6 | Ribonucleases P/MRP protein subunit POP6; Subunit of RNase MRP, nuclear RNase P and telomerase; forms a soluble heterodimer with Pop7p that binds P3 domain of RNase MRP and RNase P RNAs; RNase MRP cleaves pre-rRNA, nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs, while telomerase replenishes telomeric DNA; relocalizes to the cytosol in response to hypoxia. (158 aa) |
| | IME4 | N6-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) |
| | ROK1 | ATP-dependent RNA helicase ROK1; RNA-dependent ATPase; involved in pre-rRNA processing at sites A0, A1, and A2, and in control of cell cycle progression; contains two upstream open reading frames (uORFs) in 5' untranslated region which regulate translation; Belongs to the DEAD box helicase family. DDX52/ROK1 subfamily. (564 aa) |
| | RPS2 | Protein component of the small (40S) subunit; essential for control of translational accuracy; phosphorylation by C-terminal domain kinase I (CTDK-I) enhances translational accuracy; methylated on one or more arginine residues by Hmt1p; homologous to mammalian ribosomal protein S2 and bacterial S5. (254 aa) |
| | NSA1 | Ribosome biogenesis protein NSA1; Constituent of 66S pre-ribosomal particles; involved in 60S ribosomal subunit biogenesis. (463 aa) |
| | DBP3 | ATP-dependent RNA helicase DBP3; RNA-Dependent ATPase, member of DExD/H-box family; involved in cleavage of site A3 within the ITS1 spacer during rRNA processing; not essential for growth, but deletion causes severe slow-growth phenotype. (523 aa) |
| | RPB9 | RNA polymerase II subunit B12.6; contacts DNA; mutations affect transcription start site selection and fidelity of transcription. (122 aa) |
| | CGR1 | rRNA-processing protein CGR1; Protein involved in nucleolar integrity and processing of pre-rRNA; has a role in processing rRNA for the 60S ribosome subunit; transcript is induced in response to cytotoxic stress but not genotoxic stress; relocalizes from nucleus to nucleolus upon DNA replication stress. (120 aa) |
| | CKB1 | Beta regulatory 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. (278 aa) |
| | CDC14 | Tyrosine-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) |
| | LOC1 | 60S ribosomal subunit assembly/export protein LOC1; Nuclear protein involved in asymmetric localization of ASH1 mRNA; binds double-stranded RNA in vitro; constituent of 66S pre-ribosomal particles; required at post-transcriptional step for efficient retrotransposition; absence results in decreased Ty1 Gag:GFP protein levels; relocalizes from nucleus to cytoplasm upon DNA replication stress. (204 aa) |
| | CDC4 | Cell division control protein 4; F-box protein required for both the G1/S and G2/M phase transitions; modular substrate specificity factor which associates with core SCF (Cdc53p, Skp1p and Hrt1p/Rbx1p) to form the SCFCdc4 complex; SCFCdc4 acts as a ubiquitin-protein ligase directing ubiquitination of cyclin-dependent kinase (CDK) phosphorylated substrates, such as: Sic1p, Far1p, Cdc6p, Clb6p, and Cln3p. (779 aa) |
| | SPB4 | Putative ATP-dependent RNA helicase; nucleolar protein required for synthesis of 60S ribosomal subunits at a late step in the pathway; sediments with 66S pre-ribosomes in sucrose gradients; Belongs to the DEAD box helicase family. DDX55/SPB4 subfamily. (606 aa) |
| | CHD1 | Chromo domain-containing protein 1; Chromatin remodeler that regulates various aspects of transcription; acts in in conjunction with Isw1b to regulate chromatin structure and maintain chromatin integrity during transcription elongation by RNAP II by preventing trans-histone exchange over coding regions; contains a chromo domain, a helicase domain and a DNA-binding domain; component of both the SAGA and SLIK complexes. (1468 aa) |
| | LSM5 | U6 snRNA-associated Sm-like protein LSm5; Lsm (Like Sm) protein; part of heteroheptameric complexes (Lsm2p-7p and either Lsm1p or 8p): cytoplasmic Lsm1p complex involved in mRNA decay; nuclear Lsm8p complex part of U6 snRNP and possibly involved in processing tRNA, snoRNA, and rRNA. (93 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) |
| | LCP5 | U3 small nucleolar ribonucleoprotein protein LCP5; Essential protein involved in maturation of 18S rRNA; depletion leads to inhibited pre-rRNA processing and reduced polysome levels; localizes primarily to the nucleolus. (357 aa) |
| | NSA2 | Ribosome biogenesis protein NSA2; Protein constituent of 66S pre-ribosomal particles; contributes to processing of the 27S pre-rRNA; recruited by ribosomal proteins L17, L35, and L37 to assembling ribosomes after 27SB pre-rRNA is generated, immediately preceding removal of ITS2. (261 aa) |
| | LSM4 | U6 snRNA-associated Sm-like protein LSm4; Lsm (Like Sm) protein; part of heteroheptameric complexes (Lsm2p-7p and either Lsm1p or 8p): cytoplasmic Lsm1p complex involved in mRNA decay; nuclear Lsm8p complex part of U6 snRNP and possibly involved in processing tRNA, snoRNA, and rRNA; forms cytoplasmic foci upon DNA replication stress. (187 aa) |
| | UTP7 | Nucleolar protein; component of the small subunit (SSU) processome containing the U3 snoRNA that is involved in processing of pre-18S rRNA. (554 aa) |
| | EDC2 | Enhancer of mRNA-decapping protein 2; RNA-binding protein that directly activates mRNA decapping; binds mRNA substrate and enhances activity of decapping proteins Dcp1p and Dcp2p; has a role in translation during heat stress; protein increases in abundance and relocalizes to nucleolus and to nuclear foci upon DNA replication stress; EDC2 has a paralog, EDC1, that arose from the whole genome duplication. (145 aa) |
| | NUG1 | Nuclear GTP-binding protein NUG1; GTPase that associates with nuclear 60S pre-ribosomes; required for export of 60S ribosomal subunits from the nucleus. (520 aa) |
| | NOP16 | Nucleolar protein 16; Constituent of 66S pre-ribosomal particles; involved in 60S ribosomal subunit biogenesis; Belongs to the NOP16 family. (231 aa) |
| | POL5 | rDNA transcriptional regulator POL5; DNA Polymerase phi; has sequence similarity to the human MybBP1A and weak sequence similarity to B-type DNA polymerases, not required for chromosomal DNA replication; required for the synthesis of rRNA. (1022 aa) |
| | SNU13 | 13 kDa ribonucleoprotein-associated protein; RNA binding protein; part of U3 snoRNP involved in rRNA processing, part of U4/U6-U5 tri-snRNP involved in mRNA splicing, similar to human 15.5K protein; Belongs to the eukaryotic ribosomal protein eL8 family. (126 aa) |
| | PUF6 | Pumilio-homology domain protein; binds the 3' UTR of ASH1 mRNA and represses its translation, resulting in proper asymmetric localization of ASH1 mRNA; required at post-transcriptional step for efficient retrotransposition; absence results in decreased Ty1 Gag:GFP protein levels; co-sediments with the 60S ribosomal subunit and is required for its biogenesis; Belongs to the PUF6 family. (656 aa) |
| | SNM1 | Ribonuclease MRP complex subunit; ribonuclease (RNase) MRP cleaves pre-rRNA and has a role in cell cycle-regulated degradation of daughter cell-specific mRNAs; binds to the NME1 RNA subunit of RNase MRP. (198 aa) |
| | TOM1 | E3 ubiquitin ligase of the hect-domain class; has a role in mRNA export from the nucleus and may regulate transcriptional coactivators; involved in degradation of excess histones; interacts with Dia2p and is required for Dia2p degradation; required to target Cdc6p for ubiquitin-mediated destruction during G1 phase; Belongs to the UPL family. TOM1/PTR1 subfamily. (3268 aa) |
| | UTP6 | Nucleolar protein; component of the small subunit (SSU) processome containing the U3 snoRNA that is involved in processing of pre-18S rRNA. (440 aa) |
| | LRS4 | Monopolin 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) |
| | RRP17 | Ribosomal RNA-processing protein 17; Component of the pre-60S pre-ribosomal particle; required for cell viability under standard (aerobic) conditions but not under anaerobic conditions; exonuclease required for 5' end processing of pre-60S ribosomal RNA. (235 aa) |
| | UTP5 | U3 small nucleolar RNA-associated protein 5; Subunit of U3-containing Small Subunit (SSU) processome complex; involved in production of 18S rRNA and assembly of small ribosomal subunit. (643 aa) |
| | LSM6 | U6 snRNA-associated Sm-like protein LSm6; Lsm (Like Sm) protein; part of heteroheptameric complexes (Lsm2p-7p and either Lsm1p or 8p): cytoplasmic Lsm1p complex involved in mRNA decay; nuclear Lsm8p complex part of U6 snRNP and possibly involved in processing tRNA, snoRNA, and rRNA; Belongs to the snRNP Sm proteins family. SmF/LSm6 subfamily. (86 aa) |
| | ESF1 | Pre-rRNA-processing protein ESF1; Nucleolar protein involved in pre-rRNA processing; depletion causes severely decreased 18S rRNA levels. (628 aa) |
| | FCF1 | rRNA-processing protein FCF1; Putative PINc domain nuclease; required for early cleavages of 35S pre-rRNA and maturation of 18S rRNA; component of the SSU (small subunit) processome involved in 40S ribosomal subunit biogenesis; copurifies with Faf1p. (189 aa) |
| | UTP4 | U3 small nucleolar RNA-associated protein 4; Subunit of U3-containing 90S preribosome and SSU processome complexes; involved in production of 18S rRNA and assembly of small ribosomal subunit; member of t-Utp subcomplex involved with transcription of 35S rRNA transcript; Small Subunit processome is also known as SSU processome. (776 aa) |
| | SSF2 | Ribosome biogenesis protein SSF2; Protein required for ribosomal large subunit maturation; functionally redundant with Ssf1p; member of the Brix family; SSF2 has a paralog, SSF1, that arose from the whole genome duplication. (453 aa) |
| | BFR2 | Protein BFR2; Component of the SSU and 90S preribosomes; involved in pre-18S rRNA processing; binds to U3 snoRNA and Mpp10p; multicopy suppressor of sensitivity to Brefeldin A; expression is induced during lag phase and also by cold shock. (534 aa) |
| | RRP45 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; has similarity to E. coli RNase PH and to human hRrp45p (PM/SCL-75, EXOSC9); protein abundance increases in response to DNA replication stress. (305 aa) |
| | HMO1 | Chromatin associated high mobility group (HMG) family member; involved in compacting, bending, bridging and looping DNA; rDNA-binding component that regulates transcription from RNA polymerase I promoters; regulates start site selection of ribosomal protein genes via RNA polymerase II promoters; role in genome maintenance; associates with a 5'-3' DNA helicase and Fpr1p, a prolyl isomerase; relocalizes to the cytosol in response to hypoxia. (246 aa) |
| | RPA14 | RNA polymerase I subunit A14. (137 aa) |
| | FOB1 | Nucleolar protein that binds the rDNA replication fork barrier site; required for replication fork blocking, recombinational hotspot activity, condensin recruitment to replication fork barrier (RFB), and rDNA repeat segregation; related to retroviral integrases. (566 aa) |
| | ARX1 | Probable metalloprotease ARX1; Nuclear export factor for the ribosomal pre-60S subunit; shuttling factor which directly binds FG rich nucleoporins and facilities translocation through the nuclear pore complex; interacts directly with Alb1p; responsible for Tif6p recycling defects in the absence of Rei1; associated with the ribosomal export complex. (593 aa) |
| | RRP1 | Ribosomal RNA-processing protein 1; Essential evolutionarily conserved nucleolar protein; necessary for biogenesis of 60S ribosomal subunits and for processing of pre-rRNAs to mature rRNA; associated with several distinct 66S pre-ribosomal particles; Belongs to the RRP1 family. (278 aa) |
| | RRP8 | 25S rRNA (adenine(645)-N(1))-methyltransferase; Nucleolar S-adenosylmethionine-dependent rRNA methyltransferase; methylates adenine (m1A) of the large subunit (LSU) rRNA at position 645; involved in pre-rRNA cleavage at site A2; mutation is synthetically lethal with a gar1 mutation; deletion disrupts telomere maintenance by influencing the expression of neighboring gene STN1; Belongs to the methyltransferase superfamily. RRP8 family. (392 aa) |
| | RPS13 | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S13 and bacterial S15. (151 aa) |
| | MAK21 | Ribosome biogenesis protein MAK21; Constituent of 66S pre-ribosomal particles; required for large (60S) ribosomal subunit biogenesis; acts as part of a Mak21p-Noc2p-Rrp5p module that associates with nascent pre-rRNA during transcription and has a role in bigenesis of the large ribosomal subunit; involved in nuclear export of pre-ribosomes; required for maintenance of dsRNA virus; homolog of human CAATT-binding protein. (1025 aa) |
| | RPC11 | RNA polymerase III subunit C11; mediates pol III RNA cleavage activity and is important for termination of transcription; homologous to TFIIS. (110 aa) |
| | NSI1 | RNA polymerase I termination factor; binds to rDNA terminator element, required for efficient Pol I termination; required for rDNA silencing at NTS1; facilities association of Sir2p with NTS1, contributes to rDNA stability and cell longevity; interacts physically with Fob1p and RENT subunits, Sir2p and Net1p; may interact with ribosomes, based on co-purification experiments; Myb-like DNA-binding protein; NSI1 has a paralog, REB1, that arose from the whole genome duplication. (570 aa) |
| | FAL1 | ATP-dependent RNA helicase FAL1; Nucleolar protein required for maturation of 18S rRNA; member of the eIF4A subfamily of DEAD-box ATP-dependent RNA helicases; 18S rRNA biogenesis defect of the null mutant is functionally complemented by human EIF4A3; Belongs to the DEAD box helicase family. DDX48/FAL1 subfamily. (399 aa) |
| | NOP6 | Nucleolar protein 6; rRNA-binding protein required for 40S ribosomal subunit biogenesis; contains an RNA recognition motif (RRM); hydrophilin essential to overcome the stress of the desiccation-rehydration process; NOP6 may be a fungal-specific gene as no homologs have been yet identified in higher eukaryotes. (225 aa) |
| | NHP2 | H/ACA ribonucleoprotein complex subunit NHP2; Protein related to mammalian high mobility group (HMG) proteins; nuclear protein; essential for function of H/ACA-type snoRNPs, which are involved in 18S rRNA processing; Belongs to the eukaryotic ribosomal protein eL8 family. (156 aa) |
| | AIR2 | Protein AIR2; RNA-binding subunit of the TRAMP nuclear RNA surveillance complex; involved in nuclear RNA processing and degradation; involved in TRAMP complex assembly as a bridge between Mtr4p and Trf4p; stimulates the poly(A) polymerase activity of Pap2p in vitro; has 5 zinc knuckle motifs; AIR2 has a paralog, AIR1, that arose from the whole genome duplication; Air2p and Air1p have nonredundant roles in regulation of substrate specificity of the exosome. (344 aa) |
| | SAS10 | Something about silencing protein 10; Subunit of U3-containing Small Subunit (SSU) processome complex; involved in production of 18S rRNA and assembly of small ribosomal subunit; disrupts silencing when overproduced; mutant has increased aneuploidy tolerance; essential gene. (610 aa) |
| | NOP14 | Nucleolar protein; forms a complex with Noc4p that mediates maturation and nuclear export of 40S ribosomal subunits; also present in the small subunit processome complex, which is required for processing of pre-18S rRNA; Belongs to the NOP14 family. (810 aa) |
| | RRP42 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; has similarity to E. coli RNase PH and to human hRrp42p (EXOSC7). (265 aa) |
| | TSR1 | Ribosome biogenesis protein TSR1; Protein required for processing of 20S pre-rRNA in the cytoplasm; associates with pre-40S ribosomal particles; inhibits the premature association of 60S subunits with assembling 40S subunits in the cytoplasm; similar to Bms1p; relocalizes from nucleus to cytoplasm upon DNA replication stress; Belongs to the TRAFAC class translation factor GTPase superfamily. Bms1-like GTPase family. TSR1 subfamily. (788 aa) |
| | LHP1 | La protein homolog; RNA binding protein required for maturation of tRNA and U6 snRNA; acts as a molecular chaperone for RNAs transcribed by polymerase III; homologous to human La (SS-B) autoantigen. (275 aa) |
| | SIR2 | Conserved NAD+ dependent histone deacetylase of the Sirtuin family; deacetylation targets are primarily nuclear proteins; required for telomere hypercluster formation in quiescent yeast cells; involved in regulation of lifespan; plays roles in silencing at HML, HMR, telomeres, and rDNA; negatively regulates initiation of DNA replication; functions as regulator of autophagy like mammalian homolog SIRT1, and also of mitophagy. (562 aa) |
| | DBP10 | Putative ATP-dependent RNA helicase of the DEAD-box protein family; constituent of 66S pre-ribosomal particles; essential protein involved in ribosome biogenesis; Belongs to the DEAD box helicase family. DDX54/DBP10 subfamily. (995 aa) |
| | NOP1 | rRNA 2'-O-methyltransferase fibrillarin; Histone glutamine methyltransferase, modifies H2A at Q105 in nucleolus; component of the small subunit processome complex, which is required for processing of pre-18S rRNA; ortholog of mammalian fibrillarin; inviability of the null mutant is functionally complemented by human FBL. (327 aa) |
| | YCR087C-A | UPF0743 protein YCR087C-A; Putative protein of unknown function; green fluorescent protein (GFP)-fusion protein localizes to the nucleolus; YCR087C-A is not an essential gene; Belongs to the UPF0743 family. (153 aa) |
| | CSM1 | Monopolin 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) |
| | RSA4 | WD-repeat protein involved in ribosome biogenesis; may interact with ribosomes; required for maturation and efficient intra-nuclear transport or pre-60S ribosomal subunits, localizes to the nucleolus; Belongs to the NLE1/RSA4 family. (515 aa) |
| | PWP2 | Periodic tryptophan protein 2; Conserved 90S pre-ribosomal component; essential for proper endonucleolytic cleavage of the 35 S rRNA precursor at A0, A1, and A2 sites; contains eight WD-repeats; PWP2 deletion leads to defects in cell cycle and bud morphogenesis. (923 aa) |
| | BUD23 | Methyltransferase that methylates residue G1575 of 18S rRNA; required for rRNA processing and nuclear export of 40S ribosomal subunits independently of methylation activity; diploid mutant displays random budding pattern; functional homolog of human WBSCR22; Belongs to the class I-like SAM-binding methyltransferase superfamily. BUD23/WBSCR22 family. (275 aa) |
| | RRP43 | Exosome non-catalytic core component; involved in 3'-5' RNA processing and degradation in both the nucleus and the cytoplasm; has similarity to E. coli RNase PH and to human hRrp43p (OIP2, EXOSC8); protein abundance increases in response to DNA replication stress. (394 aa) |
| | RPS14A | Protein component of the small (40S) ribosomal subunit; required for ribosome assembly and 20S pre-rRNA processing; mutations confer cryptopleurine resistance; homologous to mammalian ribosomal protein S14 and bacterial S11; RPS14A has a paralog, RPS14B, that arose from the whole genome duplication. (137 aa) |
| | YCR016W | Uncharacterized protein YCR016W; Putative protein of unknown function; green fluorescent protein (GFP)-fusion protein localizes to the nucleolus and nucleus; predicted to be involved in ribosome biogenesis. (290 aa) |
| | KRR1 | KRR1 small subunit processome component; Nucleolar protein required for rRNA synthesis and ribosomal assembly; required for the synthesis of 18S rRNA and for the assembly of 40S ribosomal subunit; essential gene; Belongs to the KRR1 family. (316 aa) |
| | SPB1 | 27S pre-rRNA (guanosine(2922)-2'-O)-methyltransferase; AdoMet-dependent methyltransferase; involved in rRNA processing and 60S ribosomal subunit maturation; methylates G2922 in the tRNA docking site of the large subunit rRNA and in the absence of snR52, U2921; suppressor of PAB1 mutants. (841 aa) |
| | RRP7 | Ribosomal RNA-processing protein 7; Essential protein involved in rRNA processing and ribosome biogenesis; protein abundance increases in response to DNA replication stress. (297 aa) |
| | POP4 | RNases MRP/P 32.9 kDa subunit; Subunit of both RNase MRP and nuclear RNase P; RNase MRP cleaves pre-rRNA, while nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs; binds to the RPR1 RNA subunit in RNase P. (279 aa) |
| | ENP1 | Protein associated with U3 and U14 snoRNAs; required for pre-rRNA processing and 40S ribosomal subunit synthesis; localized in the nucleus and concentrated in the nucleolus; human BYSL functionally complements the heat sensitivity of an enp1 ts mutant; Belongs to the bystin family. (483 aa) |
| | ISW1 | ISWI chromatin-remodeling complex ATPase ISW1; ATPase subunit of imitation-switch (ISWI) class chromatin remodelers; with Ioc3p forms Isw1a complex involved in repression of transcription initiation; with Ioc2p and Ioc4p forms Isw1b complex involved in regulation of transcription elongation; Isw1b recruited to ORFs by H3K36 methylation and acts with Chd1p to prevent trans-histone exchange over coding regions; Isw1p import into nucleus depends on C-terminal bipartite nuclear targeting signal KRIR X19 KKAK. (1129 aa) |
| | RPS9B | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S9 and bacterial S4; RPS9B has a paralog, RPS9A, that arose from the whole genome duplication. (195 aa) |
| | PCH2 | Pachytene checkpoint protein 2; Hexameric ring ATPase that remodels chromosome axis protein Hop1p; nucleolar component of the pachytene checkpoint, which prevents chromosome segregation when recombination and chromosome synapsis are defective; also represses meiotic interhomolog recombination in rDNA; required for meiotic double-stranded break formation. (564 aa) |
| | RPS6B | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S6, no bacterial homolog; phosphorylated on S233 by Ypk3p in a TORC1-dependent manner, and on S232 in a TORC1/2-dependent manner by Ypk1/2/3p; RPS6B has a paralog, RPS6A, that arose from the whole genome duplication. (236 aa) |
| | POP7 | Ribonucleases P/MRP protein subunit POP7; Subunit of RNase MRP, nuclear RNase P and telomerase; forms a soluble heterodimer with Pop6p that binds P3 domain of RNase MRP and RNase P RNAs; RNase MRP cleaves pre-rRNA, nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs, while telomerase replenishes telomeric DNA. (140 aa) |
| | RPB5 | RNA polymerase subunit ABC27; common to RNA polymerases I, II, and III; contacts DNA and affects transactivation; Belongs to the archaeal RpoH/eukaryotic RPB5 RNA polymerase subunit family. (215 aa) |
| | MAK5 | Essential nucleolar protein; putative DEAD-box RNA helicase required for maintenance of M1 dsRNA virus; involved in biogenesis of large (60S) ribosomal subunits; Belongs to the DEAD box helicase family. DDX24/MAK5 subfamily. (773 aa) |
| | BMT2 | 25S rRNA (adenine(2142)-N(1))-methyltransferase; Nucleolar S-adenosylmethionine-dependent rRNA methyltransferase; methylates adenine (m1A) of the large subunit (LSU) rRNA at position 2142; belongs to Rossmann fold superfamily; green fluorescent protein (GFP)-fusion protein localizes to the nucleolus; YBR141C is not an essential gene. (337 aa) |
| | MUM2 | Protein 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) |
| | YBL028C | UPF0642 protein YBL028C; Protein of unknown function that may interact with ribosomes; green fluorescent protein (GFP)-fusion protein localizes to the nucleolus; predicted to be involved in ribosome biogenesis. (106 aa) |
| | LSM2 | U6 snRNA-associated Sm-like protein LSm2; Lsm (Like Sm) protein; part of heteroheptameric complexes (Lsm2p-7p and either Lsm1p or 8p): cytoplasmic Lsm1p complex involved in mRNA decay; nuclear Lsm8p complex part of U6 snRNP and possibly involved in processing tRNA, snoRNA, and rRNA; relocalizes from nucleus to cytoplasmic foci upon DNA replication stress. (95 aa) |
| | RRN10 | Protein involved in promoting high level transcription of rDNA; subunit of UAF (upstream activation factor) for RNA polymerase I. (145 aa) |
| | NCL1 | Multisite-specific tRNA:(cytosine-C(5))-methyltransferase; S-adenosyl-L-methionine-dependent tRNA: m5C-methyltransferase; methylates cytosine to m5C at several positions in tRNAs and intron-containing pre-tRNAs; increases proportion of tRNALeu(CAA) with m5C at wobble position in response to hydrogen peroxide, causing selective translation of mRNA from genes enriched in TTG codon; loss of NCL1 confers hypersensitivity to oxidative stress; similar to Nop2p and human proliferation associated nucleolar protein p120. (684 aa) |
| | POP8 | Ribonucleases P/MRP protein subunit POP8; Subunit of both RNase MRP and nuclear RNase P; RNase MRP cleaves pre-rRNA, while nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs; relocalizes to the cytosol in response to hypoxia. (133 aa) |
| | RRN6 | RNA polymerase I-specific transcription initiation factor RRN6; Component of the core factor (CF) rDNA transcription factor complex; CF is required for transcription of 35S rRNA genes by RNA polymerase I and is composed of Rrn6p, Rrn7p, and Rrn11p. (894 aa) |
| | UTP20 | U3 small nucleolar RNA-associated protein 20; Component of the small-subunit (SSU) processome; SSU processome is involved in the biogenesis of the 18S rRNA; Belongs to the UTP20 family. (2493 aa) |
| | ECM1 | Shuttling pre-60S factor ECM1; Pre-ribosomal factor involved in 60S ribosomal protein subunit export; associates with the pre-60S particle; shuttles between the nucleus and cytoplasm; Belongs to the ECM1 family. (212 aa) |
| | POP5 | Ribonuclease P/MRP protein subunit POP5; Subunit of both RNase MRP and nuclear RNase P; RNase MRP cleaves pre-rRNA, while nuclear RNase P cleaves tRNA precursors to generate mature 5' ends and facilitates turnover of nuclear RNAs. (173 aa) |
| | MAK16 | Essential nuclear protein; constituent of 66S pre-ribosomal particles; required for maturation of 25S and 5.8S rRNAs; required for maintenance of M1 satellite double-stranded RNA of the L-A virus. (306 aa) |
