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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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
RLP7 | Ribosome biogenesis protein RLP7; Nucleolar protein similar to large ribosomal subunit L7 proteins; constituent of 66S pre-ribosomal particles; plays an essential role in processing of precursors to the large ribosomal subunit RNAs; binds junction of ITS2 and ITS2-proximal stem between the 3' end of 5.8S rRNA and the 5' end of 25S rRNA. (322 aa) | ||||
HAS1 | ATP-dependent RNA helicase; involved in the biogenesis of 40S and 60S ribosome subunits; localizes to both the nuclear periphery and nucleolus; highly enriched in nuclear pore complex fractions; constituent of 66S pre-ribosomal particles. (505 aa) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
RLP24 | Essential protein required for ribosomal large subunit biogenesis; associated with pre-60S ribosomal subunits; stimulates the ATPase activity of Afg2p, which is required for release of Rlp24p from the pre-60S particle; has similarity to Rpl24Ap and Rpl24Bp. (199 aa) | ||||
NOC3 | Nucleolar complex-associated protein 3; Subunit of a nuclear complex with Noc2p and pre-replicative complexes; the Noc2p-Noc3p complex binds to 66S ribosomal precursors to mediate their maturation and intranuclear transport; binds to chromatin at active replication origins, and is required for pre-RC formation and maintenance during DNA replication licensing. (663 aa) | ||||
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) | ||||
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) | ||||
RPF2 | Ribosome biogenesis protein RPF2; Essential protein involved in rRNA maturation and ribosomal assembly; involved in the processing of pre-rRNA and the assembly of the 60S ribosomal subunit; interacts with ribosomal protein L11; localizes predominantly to the nucleolus; constituent of 66S pre-ribosomal particles. (344 aa) | ||||
LAS1 | Protein LAS1; Endonuclease involved in pre-rRNA processing at both ends of ITS2; functions with Grc3p in a conserved mechanism to modulate rRNA processing and ribosome biogenesis; may coordinate the action of the Rat1p-Rai1p exoRNAse; required for the G1/S cell cycle transition; human ortholog is Las1L; mutants require the SSD1-v allele for viability. (502 aa) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
NUG1 | Nuclear GTP-binding protein NUG1; GTPase that associates with nuclear 60S pre-ribosomes; required for export of 60S ribosomal subunits from the nucleus. (520 aa) | ||||
NOP16 | Nucleolar protein 16; Constituent of 66S pre-ribosomal particles; involved in 60S ribosomal subunit biogenesis; Belongs to the NOP16 family. (231 aa) | ||||
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) | ||||
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) | ||||
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) | ||||
ARX1 | Probable metalloprotease ARX1; Nuclear export factor for the ribosomal pre-60S subunit; shuttling factor which directly binds FG rich nucleoporins and facilities translocation through the nuclear pore complex; interacts directly with Alb1p; responsible for Tif6p recycling defects in the absence of Rei1; associated with the ribosomal export complex. (593 aa) | ||||
RRP1 | Ribosomal RNA-processing protein 1; Essential evolutionarily conserved nucleolar protein; necessary for biogenesis of 60S ribosomal subunits and for processing of pre-rRNAs to mature rRNA; associated with several distinct 66S pre-ribosomal particles; Belongs to the RRP1 family. (278 aa) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) |