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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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
LTV1 | Protein 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
RPS31 | Fusion protein cleaved to yield ribosomal protein S31 and ubiquitin; ubiquitin may facilitate assembly of the ribosomal protein into ribosomes; interacts genetically with translation factor eIF2B; homologous to mammalian ribosomal protein S27A, no bacterial homolog. (152 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) | ||||
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) | ||||
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) | ||||
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) | ||||
GSP1 | GTP-binding nuclear protein GSP1/CNR1; Ran GTPase; GTP binding protein (mammalian Ranp homolog) involved in the maintenance of nuclear organization, RNA processing and transport; regulated by Srm1p, Rna1p, Yrb1p, Yrb2p, Yrp4p, Yrb30p, Cse1p and Kap95p; GSP1 has a paralog, GSP2, that arose from the whole genome duplication; Belongs to the small GTPase superfamily. Ran family. (219 aa) | ||||
AFG2 | ATPase family gene 2 protein; ATPase of the CDC48/PAS1/SEC18 (AAA) family, forms a hexameric complex; is essential for pre-60S maturation and release of several preribosome maturation factors; releases Rlp24p from purified pre-60S particles in vitro; target of the ribosomal biosynthesis inhibitor diazaborine; may be involved in degradation of aberrant mRNAs. (780 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) | ||||
ECM16 | Essential DEAH-box ATP-dependent RNA helicase specific to U3 snoRNP; predominantly nucleolar in distribution; required for 18S rRNA synthesis. (1267 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) | ||||
RPS10B | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S10, no bacterial homolog; RPS10B has a paralog, RPS10A, that arose from the whole genome duplication; mutations in the human homolog associated with Diamond-Blackfan anemia. (105 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) | ||||
NGL2 | RNA exonuclease NGL2; Protein involved in 5.8S rRNA processing; Ccr4p-like RNase required for correct 3'-end formation of 5.8S rRNA at site E; similar to Ngl1p; NGL2 has a paralog, NGL3, that arose from the whole genome duplication. (515 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
RIA1 | Ribosome assembly protein 1; Cytoplasmic GTPase/eEF2-like factor involved in ribosomal biogenesis; with Sdo1p, a guanine nucleotide exchange factor (GEF), promotes release of Tif6p from 60S ribosomal subunits in the cytoplasm so that they can assemble with 40S subunits to generate mature ribosomes; required for quality control check of newly made large ribosomal subunits before they are released into the pool of translating ribosomes; Belongs to the TRAFAC class translation factor GTPase superfamily. Classic translation factor GTPase family. (1110 aa) | ||||
RPS3 | Protein 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) | ||||
RIO2 | Serine/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) | ||||
MPP6 | M-phase phosphoprotein 6 homolog; Nuclear exosome-associated RNA binding protein; involved in surveillance of pre-rRNAs and pre-mRNAs, and the degradation of cryptic non-coding RNAs (ncRNA); copurifies with ribosomes; relocalizes to the cytosol in response to hypoxia. (186 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
RAT1 | 5'-3' exoribonuclease 2; Nuclear 5' to 3' single-stranded RNA exonuclease; involved in RNA metabolism, including rRNA and snoRNA processing, as well as poly (A+) dependent and independent mRNA transcription termination; required for cotranscriptional pre-rRNA cleavage; displaces Cdk1p from elongating transcripts, especially as RNAPII reaches the poly(A) site, negatively regulates phosphorylation of the CTD of RNAPII, and inhibits RNAPII transcriptional elongation; Belongs to the 5'-3' exonuclease family. XRN2/RAT1 subfamily. (1006 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) | ||||
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) | ||||
RPS10A | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S10, no bacterial homolog; RPS10A has a paralog, RPS10B, that arose from the whole genome duplication; mutations in the human homolog associated with Diamond-Blackfan anemia. (105 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
FAP7 | Adenylate 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
LSG1 | Large subunit GTPase 1; Putative GTPase involved in 60S ribosomal subunit biogenesis; required for the release of Nmd3p from 60S subunits in the cytoplasm; Belongs to the TRAFAC class YlqF/YawG GTPase family. LSG1 subfamily. (640 aa) | ||||
PRP43 | Pre-mRNA-splicing factor ATP-dependent RNA helicase PRP43; RNA helicase in the DEAH-box family; functions in both RNA polymerase I and polymerase II transcript metabolism; catalyzes removal of U2, U5, and U6 snRNPs from the postsplicing lariat-intron ribonucleoprotein complex; required for efficient biogenesis of both small- and large-subunit rRNAs; acts with Sqs1p to promote 20S to 18S rRNA processing catalyzed by endonuclease Nob1p; Belongs to the DEAD box helicase family. DEAH subfamily. DDX15/PRP43 sub-subfamily. (767 aa) | ||||
XRN1 | 5'-3' exoribonuclease 1; Evolutionarily-conserved 5'-3' exonuclease; component of cytoplasmic processing (P) bodies involved in mRNA decay; also enters the nucleus and positively regulates transcription initiation and elongation; plays a role in microtubule-mediated processes, filamentous growth, ribosomal RNA maturation, and telomere maintenance; activated by the scavenger decapping enzyme Dcs1p. (1528 aa) | ||||
RPS26A | Protein 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) | ||||
RAI1 | Nuclear protein with decapping endonuclease activity; targets mRNAs with unmethylated 7-methylguanosine cap structures and 5'-triphosphates; binds to and stabilizes the exoribonuclease Rat1p; required for pre-rRNA processing; relocalizes to the cytosol in response to hypoxia; homologous to human DOM3Z; Belongs to the DXO/Dom3Z family. (387 aa) | ||||
CRM1 | Exportin-1; Major karyopherin; involved in export of proteins, RNAs, and ribosomal subunits from the nucleus; exportin. (1084 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) | ||||
NOP10 | Subunit of box H/ACA snoRNP complex; required for pseudouridylation and processing of pre-18S rRNA. (58 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) |