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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) | ||||
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) | ||||
RPL19B | Ribosomal 60S subunit protein L19B; rpl19a and rpl19b single null mutations result in slow growth, while the double null mutation is lethal; homologous to mammalian ribosomal protein L19, no bacterial homolog; RPL19B has a paralog, RPL19A, that arose from the whole genome duplication. (189 aa) | ||||
RPL4A | Ribosomal 60S subunit protein L4A; N-terminally acetylated; homologous to mammalian ribosomal protein L4 and bacterial L4; RPL4A has a paralog, RPL4B, that arose from the whole genome duplication. (362 aa) | ||||
RPL19A | Ribosomal 60S subunit protein L19A; rpl19a and rpl19b single null mutations result in slow growth, while the double null mutation is lethal; homologous to mammalian ribosomal protein L19, no bacterial homolog; RPL19A has a paralog, RPL19B, that arose from the whole genome duplication. (189 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) | ||||
RPL31A | Ribosomal 60S subunit protein L31A; associates with karyopherin Sxm1p; loss of both Rpl31p and Rpl39p confers lethality; homologous to mammalian ribosomal protein L31, no bacterial homolog; RPL31A has a paralog, RPL31B, that arose from the whole genome duplication. (113 aa) | ||||
SSB1 | Cytoplasmic ATPase that is a ribosome-associated molecular chaperone; functions with J-protein partner Zuo1p; may be involved in folding of newly-made polypeptide chains; member of the HSP70 family; interacts with phosphatase subunit Reg1p; SSB1 has a paralog, SSB2, that arose from the whole genome duplication; Belongs to the heat shock protein 70 family. Ssb-type Hsp70 subfamily. (613 aa) | ||||
RPL4B | Ribosomal 60S subunit protein L4B; homologous to mammalian ribosomal protein L4 and bacterial L4; RPL4B has a paralog, RPL4A, that arose from the whole genome duplication. (362 aa) | ||||
ARX1 | Probable metalloprotease ARX1; Nuclear export factor for the ribosomal pre-60S subunit; shuttling factor which directly binds FG rich nucleoporins and facilities translocation through the nuclear pore complex; interacts directly with Alb1p; responsible for Tif6p recycling defects in the absence of Rei1; associated with the ribosomal export complex. (593 aa) | ||||
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) | ||||
URA3 | Orotidine-5'-phosphate (OMP) decarboxylase; catalyzes the sixth enzymatic step in the de novo biosynthesis of pyrimidines, converting OMP into uridine monophosphate (UMP); converts 5-FOA into 5-fluorouracil, a toxic compound. (267 aa) | ||||
NOP16 | Nucleolar protein 16; Constituent of 66S pre-ribosomal particles; involved in 60S ribosomal subunit biogenesis; Belongs to the NOP16 family. (231 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) | ||||
RPL2A | Ribosomal 60S subunit protein L2A; homologous to mammalian ribosomal protein L2 and bacterial L2; RPL2A has a paralog, RPL2B, that arose from the whole genome duplication. (254 aa) | ||||
NSA1 | Ribosome biogenesis protein NSA1; Constituent of 66S pre-ribosomal particles; involved in 60S ribosomal subunit biogenesis. (463 aa) | ||||
SDA1 | Protein required for actin organization and passage through Start; highly conserved nuclear protein; required for actin cytoskeleton organization; plays a critical role in G1 events; binds Nap1p; involved in 60S ribosome biogenesis; Belongs to the SDA1 family. (767 aa) | ||||
ZUO1 | Zuotin; Ribosome-associated chaperone; zuotin functions in ribosome biogenesis and as a chaperone for nascent polypeptide chains in partnership with Ssz1p and SSb1/2; contains a DnaJ domain and functions as a J-protein partner for Ssb1p and Ssb2p; human gene DNAJC2 can partially complement yeast zuo1 null mutant. (433 aa) | ||||
NMD3 | Protein involved in nuclear export of the large ribosomal subunit; acts as a Crm1p-dependent adapter protein for export of nascent ribosomal subunits through the nuclear pore complex; Belongs to the NMD3 family. (518 aa) | ||||
RIX1 | Pre-rRNA-processing protein RIX1; Component of the Rix1 complex and possibly pre-replicative complexes; required for processing of ITS2 sequences from 35S pre-rRNA; component of the pre-60S ribosomal particle with the dynein-related AAA-type ATPase Mdn1p; required for pre-replicative complex (pre-RC) formation and maintenance during DNA replication licensing; relocalizes to the cytosol in response to hypoxia; essential gene; Belongs to the RIX1/PELP1 family. (763 aa) | ||||
RPL2B | Ribosomal 60S subunit protein L2B; homologous to mammalian ribosomal protein L2 and bacterial L2; RPL2B has a paralog, RPL2A, that arose from the whole genome duplication; expression is upregulated at low temperatures. (254 aa) | ||||
ALB1 | Ribosome biogenesis protein ALB1; Shuttling pre-60S factor; involved in the biogenesis of ribosomal large subunit; interacts directly with Arx1p; responsible for Tif6p recycling defects in absence of Rei1p. (175 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) | ||||
RPL39 | Ribosomal 60S subunit protein L39; required for ribosome biogenesis; loss of both Rpl31p and Rpl39p confers lethality; also exhibits genetic interactions with SIS1 and PAB1; homologous to mammalian ribosomal protein L39, no bacterial homolog. (51 aa) | ||||
RPL43B | Ribosomal 60S subunit protein L43B; homologous to mammalian ribosomal protein L37A, no bacterial homolog; RPL43B has a paralog, RPL43A, that arose from the whole genome duplication; protein abundance increases in response to DNA replication stress. (92 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
PWP1 | Periodic tryptophan protein 1; Protein with WD-40 repeats involved in rRNA processing; associates with trans-acting ribosome biogenesis factors; similar to beta-transducin superfamily; Belongs to the WD repeat PWP1 family. (576 aa) | ||||
RPL31B | Ribosomal 60S subunit protein L31B; associates with karyopherin Sxm1p; loss of both Rpl31p and Rpl39p confers lethality; homologous to mammalian ribosomal protein L31, no bacterial homolog; RPL31B has a paralog, RPL31A, that arose from the whole genome duplication. (113 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) | ||||
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) | ||||
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) | ||||
IPI3 | Pre-rRNA-processing protein IPI3; Component of the Rix1 complex and pre-replicative complexes (pre-RCs); required for processing of ITS2 sequences from 35S pre-rRNA; component of the pre-60S ribosomal particle with the dynein-related AAA-type ATPase Mdn1p; required for pre-RC formation and maintenance during DNA replication licensing; highly conserved protein which contains several WD40 motifs; IPI3 is an essential gene; other members include Rix1p, Ipi1p, and Ipi3p. (555 aa) | ||||
SSB2 | Cytoplasmic ATPase that is a ribosome-associated molecular chaperone; functions with J-protein partner Zuo1p; may be involved in the folding of newly-synthesized polypeptide chains; member of the HSP70 family; SSB2 has a paralog, SSB1, that arose from the whole genome duplication; Belongs to the heat shock protein 70 family. Ssb-type Hsp70 subfamily. (613 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) | ||||
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) | ||||
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) | ||||
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) | ||||
RPL3 | Ribosomal 60S subunit protein L3; homologous to mammalian ribosomal protein L3 and bacterial L3; plays an important role in function of eIF5B in stimulating 3' end processing of 18S rRNA in context of 80S ribosomes that have not yet engaged in translation; involved in replication and maintenance of killer double stranded RNA virus; Belongs to the universal ribosomal protein uL3 family. (387 aa) | ||||
RPL43A | Ribosomal 60S subunit protein L43A; null mutation confers a dominant lethal phenotype; homologous to mammalian ribosomal protein L37A, no bacterial homolog; RPL43A has a paralog, RPL43B, that arose from the whole genome duplication. (92 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) | ||||
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) | ||||
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) |