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RRN3 RRN3 FUN12 FUN12 UTP20 UTP20 SLX1 SLX1 ENP1 ENP1 KRR1 KRR1 TSR1 TSR1 NOP14 NOP14 SAS10 SAS10 FAP7 FAP7 MAK21 MAK21 RLI1 RLI1 BFR2 BFR2 FCF1 FCF1 EFT2 EFT2 UTP7 UTP7 LCP5 LCP5 RPS26B RPS26B ROK1 ROK1 RPS26A RPS26A SLX9 SLX9 ENP2 ENP2 CRM1 CRM1 LRP1 LRP1 DBP8 DBP8 SCH9 SCH9 FAF1 FAF1 CYR1 CYR1 NOP9 NOP9 HCA4 HCA4 MTR4 MTR4 UTP18 UTP18 RPA34 RPA34 DHR2 DHR2 UTP11 UTP11 LTV1 LTV1 UTP30 UTP30 SOF1 SOF1 KNS1 KNS1 FCF2 FCF2 EMG1 EMG1 IFH1 IFH1 UTP21 UTP21 TSR2 TSR2 UTP14 UTP14 ECM16 ECM16 RRP5 RRP5 HAS1 HAS1 DOM34 DOM34 IMP4 IMP4 KRE33 KRE33 RPS3 RPS3 RIO2 RIO2 RPA49 RPA49 KRI1 KRI1 ESF2 ESF2 RCL1 RCL1 UTP23 UTP23 NOB1 NOB1 RIO1 RIO1 PNO1 PNO1 NOC2 NOC2 RPA43 RPA43 RRP12 RRP12 NOP53 NOP53 HRR25 HRR25 BMS1 BMS1 YAR1 YAR1 RRP9 RRP9 NOC4 NOC4
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RRN3Protein 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)
FUN12Translation initiation factor eIF5B; GTPase that promotes Met-tRNAiMet binding to ribosomes and ribosomal subunit joining; promotes GTP-dependent maturation of 18S rRNA by Nob1p; protein abundance increases in response to DNA replication stress; homolog of bacterial IF2. (1002 aa)
UTP20U3 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)
SLX1Endonuclease involved in DNA recombination and repair; subunit of a complex, with Slx4p, that hydrolyzes 5' branches from duplex DNA in response to stalled or converging replication forks; function overlaps with that of Sgs1p-Top3p. (304 aa)
ENP1Protein 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)
KRR1KRR1 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)
TSR1Ribosome 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)
NOP14Nucleolar 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)
SAS10Something 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)
FAP7Adenylate kinase isoenzyme 6 homolog FAP7; Essential NTPase required for small ribosome subunit synthesis; mediates processing of the 20S pre-rRNA at site D in the cytoplasm but associates only transiently with 43S preribosomes via Rps14p; complex with Rps14 is conserved between humans, yeast, and arches; may be the endonuclease for site D; depletion leads to accumulation of pre-40S ribosomes in 80S-like ribosomes; human TAF9 functionally complements the lethality of the null mutation. (197 aa)
MAK21Ribosome 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)
RLI1Translation initiation factor RLI1; Essential Fe-S protein; required for ribosome biogenesis, translation initiation/termination; facilitates binding of multifactor complex (MFC) of initiation factors to small ribosomal subunit; Dom34-Hbs1 complex and Rli1p work in dissociating inactive ribosomes, thereby facilitating translation restart; forms complex with Lto1p and Yae1p; dependency on ROS-labile FeS clusters, activity in nuclear ribosomal-subunit export impaired by mild oxidative stress. (608 aa)
BFR2Protein 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)
FCF1rRNA-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)
EFT2Elongation factor 2 (EF-2), also encoded by EFT1; catalyzes ribosomal translocation during protein synthesis; contains diphthamide, the unique posttranslationally modified histidine residue specifically ADP-ribosylated by diphtheria toxin; EFT2 has a paralog, EFT1, that arose from the whole genome duplication. (842 aa)
UTP7Nucleolar protein; component of the small subunit (SSU) processome containing the U3 snoRNA that is involved in processing of pre-18S rRNA. (554 aa)
LCP5U3 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)
RPS26BProtein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S26, no bacterial homolog; RPS26B has a paralog, RPS26A, that arose from the whole genome duplication; human homolog can partially complement an RPS26A, RPS26B double null mutant; mutations in the human gene are associated with Diamond-Blackfan anemia. (119 aa)
ROK1ATP-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)
RPS26AProtein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S26, no bacterial homolog; RPS26A has a paralog, RPS26B, that arose from the whole genome duplication; human homolog can partially complement an RPS26A, RPS26B double null mutant; mutations in the human gene are associated with Diamond-Blackfan anemia. (119 aa)
SLX9Ribosome 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)
ENP2Ribosome 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)
CRM1Exportin-1; Major karyopherin; involved in export of proteins, RNAs, and ribosomal subunits from the nucleus; exportin. (1084 aa)
LRP1Nuclear 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)
DBP8ATPase, 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)
SCH9Serine/threonine-protein kinase SCH9; AGC family protein kinase; functional ortholog of mammalian S6 kinase; phosphorylated by Tor1p and required for TORC1-mediated regulation of ribosome biogenesis, translation initiation, and entry into G0 phase; involved in transactivation of osmostress-responsive genes; regulates G1 progression, cAPK activity and nitrogen activation of the FGM pathway; integrates nutrient signals and stress signals from sphingolipids to regulate lifespan. (824 aa)
FAF1Protein required for pre-rRNA processing; also required for 40S ribosomal subunit assembly. (346 aa)
CYR1Adenylate cyclase; required for cAMP production and cAMP-dependent protein kinase signaling; the cAMP pathway controls a variety of cellular processes, including metabolism, cell cycle, stress response, stationary phase, and sporulation. (2026 aa)
NOP9Nucleolar 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)
HCA4DEAD 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)
MTR4ATP-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)
UTP18Small-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)
RPA34RNA 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)
DHR2Predominantly nucleolar DEAH-box ATP-dependent RNA helicase; required for 18S rRNA synthesis; Belongs to the DEAD box helicase family. DEAH subfamily. (735 aa)
UTP11U3 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)
LTV1Protein LTV1; Component of the GSE complex; GSE is required for proper sorting of amino acid permease Gap1p; required for ribosomal small subunit export from nucleus; required for growth at low temperature; Belongs to the LTV1 family. (463 aa)
UTP30Ribosome 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)
SOF1Protein 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)
KNS1Dual specificity protein kinase KNS1; Protein kinase involved in negative regulation of PolIII transcription; effector kinase of the TOR signaling pathway and phosphorylates Rpc53p to regulate ribosome and tRNA biosynthesis; member of the LAMMER family of protein kinases, which are serine/threonine kinases also capable of phosphorylating tyrosine residues; capable of autophosphorylation. (737 aa)
FCF2rRNA-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)
EMG1Ribosomal 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)
IFH1Protein 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)
UTP21U3 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)
TSR2Pre-rRNA-processing protein TSR2; Protein with a potential role in pre-rRNA processing; Belongs to the TSR2 family. (205 aa)
UTP14U3 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)
ECM16Essential DEAH-box ATP-dependent RNA helicase specific to U3 snoRNP; predominantly nucleolar in distribution; required for 18S rRNA synthesis. (1267 aa)
RRP5rRNA 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)
HAS1ATP-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)
DOM34Protein that facilitates ribosomal subunit dissociation; Dom34-Hbs1 complex and Rli1p have roles in dissociating inactive ribosomes to facilitate translation restart, particularly ribosomes stalled in 3' UTRs; required for RNA cleavage in no-go decay, but reports conflict on endonuclease activity; Pelota ortholog; protein abundance increases in response to DNA replication stress; DOM34 has a paralog, YCL001W-B, that arose from the whole genome duplication. (386 aa)
IMP4U3 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)
KRE33RNA 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)
RPS3Protein component of the small (40S) ribosomal subunit; has apurinic/apyrimidinic (AP) endonuclease activity; essential for viability; nascent Rps3p is bound by specific chaperone Yar1p during translation; homologous to mammalian ribosomal protein S3 and bacterial S3. (240 aa)
RIO2Serine/threonine-protein kinase RIO2; Essential serine kinase involved in the processing of 20S pre-rRNA; involved in the processing of the 20S pre-rRNA into mature 18S rRNA; has similarity to Rio1p. (425 aa)
RPA49RNA polymerase I subunit A49; essential for nucleolar assembly and for high polymerase loading rate; required for nucleolar localization of Rpa34p. (415 aa)
KRI1Protein 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)
ESF2Pre-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)
RCL1RNA 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)
UTP23rRNA-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)
NOB120S-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)
RIO1Serine/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)
PNO1Pre-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)
NOC2Nucleolar 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)
RPA43RNA polymerase I subunit A43; Belongs to the eukaryotic RPA43 RNA polymerase subunit family. (326 aa)
RRP12Protein 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)
NOP53Nucleolar 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)
HRR25Conserved casein kinase; regulates diverse events including: vesicular traffic, DNA repair, the CVT pathway, monopolar attachment of sister kinetochores at meiosis I, and ribosomal subunit biogenesis; monopolin subunit; binds the RNAPII CTD; phosphorylates COPII coat subunits; interacts with Sit4p phosphatase; antagonizes calcineurin signaling, reducing nuclear accumulation of Crz1p; phosphorylates Dsn1p, the kinetochore receptor for monopolin; homolog of mammalian CK1delta. (494 aa)
BMS1Ribosome 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)
YAR1Ankyrin repeat-containing protein YAR1; Ankyrin-repeat containing, nucleocytoplasmic shuttling chaperone; prevents aggregation of Rps3p in the cytoplasm, associates with nascent Rps3p during its translation in the cytoplasm and delivers it to the 90S in the nucleus; required for 40S ribosomal subunit export, biogenesis and adaptation to osmotic and oxidative stress; expression repressed by heat shock. (200 aa)
RRP9Ribosomal 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)
NOC4Nucleolar 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)
Your Current Organism:
Saccharomyces cerevisiae
NCBI taxonomy Id: 4932
Other names: ATCC 18824, Candida robusta, Mycoderma cerevisiae, NRRL Y-12632, S. cerevisiae, Saccharomyces capensis, Saccharomyces italicus, Saccharomyces oviformis, Saccharomyces uvarum var. melibiosus, yeast
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