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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) | ||||
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) | ||||
FUN12 | Translation initiation factor eIF5B; GTPase that promotes Met-tRNAiMet binding to ribosomes and ribosomal subunit joining; promotes GTP-dependent maturation of 18S rRNA by Nob1p; protein abundance increases in response to DNA replication stress; homolog of bacterial IF2. (1002 aa) | ||||
MAK5 | Essential nucleolar protein; putative DEAD-box RNA helicase required for maintenance of M1 dsRNA virus; involved in biogenesis of large (60S) ribosomal subunits; Belongs to the DEAD box helicase family. DDX24/MAK5 subfamily. (773 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) | ||||
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) | ||||
BUD23 | Methyltransferase that methylates residue G1575 of 18S rRNA; required for rRNA processing and nuclear export of 40S ribosomal subunits independently of methylation activity; diploid mutant displays random budding pattern; functional homolog of human WBSCR22; Belongs to the class I-like SAM-binding methyltransferase superfamily. BUD23/WBSCR22 family. (275 aa) | ||||
RSA4 | WD-repeat protein involved in ribosome biogenesis; may interact with ribosomes; required for maturation and efficient intra-nuclear transport or pre-60S ribosomal subunits, localizes to the nucleolus; Belongs to the NLE1/RSA4 family. (515 aa) | ||||
SYO1 | Synchronized import protein 1; Transport adaptor or symportin; facilitates synchronized nuclear coimport of the two 5S-rRNA binding proteins Rpl5p and Rpl11p; binds to nascent Rpl5p during translation; required for biogenesis of the large ribosomal subunit; green fluorescent protein (GFP)-fusion protein localizes to the cytoplasm and nucleus. (620 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) | ||||
ACL4 | Specific assembly chaperone for ribosomal protein Rpl4p; binds to an evolutionarily conserved surface extension of nascent Rpl4p and chaperones Rpl4p until its assembly into the pre-ribosome; transcriptionally co-regulated with rRNA and ribosome biosynthesis genes; Belongs to the ACL4 family. (387 aa) | ||||
BCP1 | Essential protein involved in nuclear export of Mss4p; Mss4p is a lipid kinase that generates phosphatidylinositol 4,5-biphosphate and plays a role in actin cytoskeleton organization and vesicular transport; Belongs to the BCP1 family. (283 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) | ||||
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) | ||||
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) | ||||
NSA1 | Ribosome biogenesis protein NSA1; Constituent of 66S pre-ribosomal particles; involved in 60S ribosomal subunit biogenesis. (463 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) | ||||
RPL11B | Ribosomal 60S subunit protein L11B; expressed at half the level of Rpl11Ap; involved in ribosomal assembly; depletion causes degradation of 60S proteins and RNA; homologous to mammalian ribosomal protein L11 and bacterial L5; RPL11B has a paralog, RPL11A, that arose from the whole genome duplication. (174 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) | ||||
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) | ||||
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) | ||||
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) | ||||
URB1 | Protein required for the normal accumulation of 25S and 5.8S rRNAs; nucleolar protein; associated with the 27SA2 pre-ribosomal particle; proposed to be involved in the biogenesis of the 60S ribosomal subunit. (1764 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) | ||||
DBP7 | Putative ATP-dependent RNA helicase of the DEAD-box family; involved in ribosomal biogenesis; required at post-transcriptional step for efficient retrotransposition; essential for growth under anaerobic conditions. (742 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) | ||||
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) | ||||
REX2 | Oligoribonuclease, mitochondrial; 3'-5' RNA exonuclease; involved in 3'-end processing of U4 and U5 snRNAs, 5S and 5.8S rRNAs, and RNase P and RNase MRP RNA; localized to mitochondria and null suppresses escape of mtDNA to nucleus in yme1 yme2 mutants; RNase D exonuclease. (269 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) | ||||
REX3 | RNA exonuclease; required for maturation of the RNA component of RNase MRP; functions redundantly with Rnh70p and Rex2p in processing of U5 snRNA and RNase P RNA; member of RNase D family of exonucleases; Belongs to the REXO1/REXO3 family. (404 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) | ||||
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) | ||||
DBP9 | ATP-dependent RNA helicase DBP9; DEAD-box protein required for 27S rRNA processing; exhibits DNA, RNA and DNA/RNA helicase activities; ATPase activity shows preference for DNA over RNA; DNA helicase activity abolished by mutation in RNA-binding domain; Belongs to the DEAD box helicase family. DDX56/DBP9 subfamily. (594 aa) | ||||
REH1 | Cytoplasmic 60S subunit biogenesis factor; associates with pre-60S particles; similar to Rei1p and shares partially redundant function in cytoplasmic 60S subunit maturation; contains dispersed C2H2 zinc finger domains; Belongs to the REI1 family. (432 aa) | ||||
TSR2 | Pre-rRNA-processing protein TSR2; Protein with a potential role in pre-rRNA processing; Belongs to the TSR2 family. (205 aa) | ||||
ECM16 | Essential DEAH-box ATP-dependent RNA helicase specific to U3 snoRNP; predominantly nucleolar in distribution; required for 18S rRNA synthesis. (1267 aa) | ||||
RRB1 | Ribosome assembly protein RRB1; Specific chaperone for ribosomal protein Rpl3p; binds to nascent Rpl3p during translation; essential gene. (511 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
TRM112 | Multifunctional methyltransferase subunit TRM112; Protein involved in methylation of tRNA, rRNA, and translation factors; also involved in ribosome biogenesis; subunit of tRNA methyltransferase (MTase) complexes in combination with Trm9p and Trm11p; N7-methylates G1575 of 18S rRNA as complex with Bud23p; subunit of complex with Mtq2p that methylates Sup45p (eRF1) in the ternary complex eRF1-eRF3-GTP; relative distribution to the nucleus increases upon DNA replication stress; functional homolog of human TRMT112. (135 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) | ||||
NOP8 | Nucleolar protein required for 60S ribosomal subunit biogenesis. (484 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) | ||||
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) | ||||
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) | ||||
NOP4 | Nucleolar protein; essential for processing and maturation of 27S pre-rRNA and large ribosomal subunit biogenesis; constituent of 66S pre-ribosomal particles; contains four RNA recognition motifs (RRMs). (685 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) | ||||
RPL5 | Ribosomal 60S subunit protein L5; nascent Rpl5p is bound by specific chaperone Syo1p during translation; homologous to mammalian ribosomal protein L5 and bacterial L18; binds 5S rRNA and is required for 60S subunit assembly; Belongs to the universal ribosomal protein uL18 family. (297 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) | ||||
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) | ||||
YAR1 | Ankyrin repeat-containing protein YAR1; Ankyrin-repeat containing, nucleocytoplasmic shuttling chaperone; prevents aggregation of Rps3p in the cytoplasm, associates with nascent Rps3p during its translation in the cytoplasm and delivers it to the 90S in the nucleus; required for 40S ribosomal subunit export, biogenesis and adaptation to osmotic and oxidative stress; expression repressed by heat shock. (200 aa) | ||||
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) | ||||
RPL11A | Ribosomal 60S subunit protein L11A; expressed at twice the level of Rpl11Bp; involved in ribosomal assembly; depletion causes degradation of 60S proteins and RNA; homologous to mammalian ribosomal protein L11 and bacterial L5; RPL11A has a paralog, RPL11B, that arose from the whole genome duplication. (174 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) |