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| | 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) |
| | RPS1A | Ribosomal protein 10 (rp10) of the small (40S) subunit; homologous to mammalian ribosomal protein S3A, no bacterial homolog; RPS1A has a paralog, RPS1B, that arose from the whole genome duplication. (255 aa) |
| | RPS1B | Ribosomal protein 10 (rp10) of the small (40S) subunit; homologous to mammalian ribosomal protein S3A, no bacterial homolog; RPS1B has a paralog, RPS1A, that arose from the whole genome duplication. (255 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) |
| | 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) |
| | 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) |
| | 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) |
| | ECM16 | Essential DEAH-box ATP-dependent RNA helicase specific to U3 snoRNP; predominantly nucleolar in distribution; required for 18S rRNA synthesis. (1267 aa) |
| | RPS16A | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S16 and bacterial S9; RPS16A has a paralog, RPS16B, that arose from the whole genome duplication. (143 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | RPS5 | Protein component of the small (40S) ribosomal subunit; least basic of non-acidic ribosomal proteins; phosphorylated in vivo; essential for viability; homologous to mammalian ribosomal protein S5 and bacterial S7. (225 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) |
| | RPL17B | Ribosomal 60S subunit protein L17B; required for processing of 27SB pre-rRNA and formation of stable 66S assembly intermediates; homologous to mammalian ribosomal protein L17 and bacterial L22; RPL17B has a paralog, RPL17A, that arose from the whole genome duplication. (184 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) |
| | 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) |
| | 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) |
| | 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) |
| | UTP25 | U3 small nucleolar RNA-associated protein 25; Nucleolar protein; required for 35S pre-RNA processing and 40S ribosomal subunit biogenesis. (721 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) |
| | RPS4B | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S4, no bacterial homolog; RPS4B has a paralog, RPS4A, that arose from the whole genome duplication. (261 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | RPS0A | Ribosomal 40S subunit protein S0A; required for maturation of 18S rRNA along with Rps0Bp; deletion of either RPS0 gene reduces growth rate, deletion of both genes is lethal; homologous to human ribosomal protein SA and bacterial S2; RPS0A has a paralog, RPS0B, that arose from the whole genome duplication. (252 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | RPS11A | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S11 and bacterial S17; N-terminally propionylated in vivo; RPS11A has a paralog, RPS11B, that arose from the whole genome duplication. (156 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) |
| | RPL35A | Ribosomal 60S subunit protein L35A; homologous to mammalian ribosomal protein L35 and bacterial L29; RPL35A has a paralog, RPL35B, that arose from the whole genome duplication. (120 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) |
| | RPL35B | Ribosomal 60S subunit protein L35B; homologous to mammalian ribosomal protein L35 and bacterial L29; RPL35B has a paralog, RPL35A, that arose from the whole genome duplication. (120 aa) |
| | RPS16B | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S16 and bacterial S9; RPS16B has a paralog, RPS16A, that arose from the whole genome duplication. (143 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | REI1 | Cytoplasmic pre-60S factor; required for the correct recycling of shuttling factors Alb1, Arx1 and Tif6 at the end of the ribosomal large subunit biogenesis; involved in bud growth in the mitotic signaling network. (393 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) |
| | 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) |
| | 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) |
| | RPS11B | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S11 and bacterial S17; RPS11B has a paralog, RPS11A, that arose from the whole genome duplication. (156 aa) |
| | RPS8A | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S8, no bacterial homolog; RPS8A has a paralog, RPS8B, that arose from the whole genome duplication. (200 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) |
| | 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) |
| | 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) |
| | RPS26B | Protein 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) |
| | 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) |
| | RPS8B | Protein component of the small (40S) ribosomal subunit; homologous to mammalian ribosomal protein S8, no bacterial homolog; RPS8B has a paralog, RPS8A, that arose from the whole genome duplication. (200 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) |
| | RPL34A | Ribosomal 60S subunit protein L34A; homologous to mammalian ribosomal protein L34, no bacterial homolog; RPL34A has a paralog, RPL34B, that arose from the whole genome duplication. (121 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | RPL25 | Ribosomal 60S subunit protein L25; primary rRNA-binding ribosomal protein component of large ribosomal subunit; binds to 25S rRNA via a conserved C-terminal motif; homologous to mammalian ribosomal protein L23A and bacterial L23; Belongs to the universal ribosomal protein uL23 family. (142 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | RPL27B | Ribosomal 60S subunit protein L27B; homologous to mammalian ribosomal protein L27, no bacterial homolog; RPL27B has a paralog, RPL27A, that arose from the whole genome duplication. (136 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) |
| | RPL37B | Ribosomal 60S subunit protein L37B; required for processing of 27SB pre-rRNA and formation of stable 66S assembly intermediates; protein abundance increases in response to DNA replication stress; homologous to mammalian ribosomal protein L37, no bacterial homolog; RPL37B has a paralog, RPL37A, that arose from the whole genome duplication. (88 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) |
| | 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) |
| | NOP16 | Nucleolar protein 16; Constituent of 66S pre-ribosomal particles; involved in 60S ribosomal subunit biogenesis; Belongs to the NOP16 family. (231 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) |
| | RPS4A | Protein component of the small (40S) ribosomal subunit; mutation affects 20S pre-rRNA processing; homologous to mammalian ribosomal protein S4, no bacterial homolog; RPS4A has a paralog, RPS4B, that arose from the whole genome duplication. (261 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) |
| | 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) |
| | 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) |
| | RPL17A | Ribosomal 60S subunit protein L17A; required for processing of 27SB pre-rRNA and formation of stable 66S assembly intermediates; copurifies with the Dam1 complex (aka DASH complex); homologous to mammalian ribosomal protein L17 and bacterial L22; RPL17A has a paralog, RPL17B, that arose from the whole genome duplication; protein abundance increases in response to DNA replication stress. (184 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | CMS1 | Protein CMS1; Putative subunit of the 90S preribosome processome complex; overexpression rescues supressor mutant of mcm10; null mutant is viable; relocalizes from nucleus to cytoplasm upon DNA replication stress. (291 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | 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) |
| | BUD20 | Bud site selection protein 20; C2H2-type zinc finger protein required for ribosome assembly; shuttling factor which associates with pre-60S particles in the nucleus, accompanying them to the cytoplasm; cytoplasmic dissociation of Bud20p requires Drg1p; N-terminus harbors a nuclear localization signal (NLS) and a nuclear export signal (NES); cytoplasmic Bud20p is reimported by Kap123-dependent pathway; involved in bud-site selection; diploid mutants display a random budding pattern; similar to human ZNF593. (166 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) |
| | 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) |
| | RPL37A | Ribosomal 60S subunit protein L37A; required for processing of 27SB pre-rRNA and formation of stable 66S assembly intermediates; homologous to mammalian ribosomal protein L37, no bacterial homolog; RPL37A has a paralog, RPL37B, that arose from the whole genome duplication. (88 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) |
| | 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) |
| | 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) |
| | RPP0 | Conserved ribosomal protein P0 of the ribosomal stalk; involved in interaction between translational elongation factors and the ribosome; phosphorylated on serine 302; homologous to mammalian ribosomal protein LP0 and bacterial L10. (312 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) |
| | 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) |
| | 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) |
| | 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) |
