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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) | ||||
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) | ||||
TSR2 | Pre-rRNA-processing protein TSR2; Protein with a potential role in pre-rRNA processing; Belongs to the TSR2 family. (205 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
RPL7B | Ribosomal 60S subunit protein L7B; required for processing of 27SA3 pre-rRNA to 27SB pre-rRNA during assembly of large ribosomal subunit; depletion leads to a turnover of pre-rRNA; contains a conserved C-terminal Nucleic acid Binding Domain (NDB2); binds to Domain II of 25S and 5.8S rRNAs; homologous to mammalian ribosomal protein L7 and bacterial L30; RPL7B has a paralog, RPL7A, that arose from the whole genome duplication. (244 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
GRC3 | Polynucleotide 5'-hydroxyl-kinase GRC3; Polynucleotide kinase present on rDNA; required for efficient transcription termination by RNA polymerase I; functions with Las1p in a conserved mechanism to modulate rRNA processing and ribosome biogenesis; required for cell growth; mRNA is cell-cycle regulated; Belongs to the Clp1 family. NOL9/GRC3 subfamily. (632 aa) | ||||
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) | ||||
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) | ||||
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) | ||||
MAK11 | Protein involved in an early step of 60S ribosomal subunit biogenesis; essential for cell growth and replication of killer M1 dsRNA virus; contains four beta-transducin repeats. (468 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) | ||||
HIT1 | Protein involved in C/D snoRNP assembly; regulates abundance of Rsa1p; required for growth at high temperature; similar to human ZNHIT3. (164 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) | ||||
IPI1 | Pre-rRNA-processing protein IPI1; 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 IPI1/TEX10 family. (334 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) | ||||
BCD1 | Box C/D snoRNA protein 1; Essential protein required for the accumulation of box C/D snoRNA. (366 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) | ||||
RAI1 | Nuclear protein with decapping endonuclease activity; targets mRNAs with unmethylated 7-methylguanosine cap structures and 5'-triphosphates; binds to and stabilizes the exoribonuclease Rat1p; required for pre-rRNA processing; relocalizes to the cytosol in response to hypoxia; homologous to human DOM3Z; Belongs to the DXO/Dom3Z family. (387 aa) | ||||
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) | ||||
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) | ||||
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) | ||||
RPL7A | Ribosomal 60S subunit protein L7A; required for processing of 27SA3 pre-rRNA to 27SB pre-rRNA during assembly of large ribosomal subunit; depletion leads to a turnover of pre-rRNA; contains a conserved C-terminal Nucleic acid Binding Domain (NDB2); binds to Domain II of 25S and 5.8S rRNAs; homologous to mammalian ribosomal protein L7 and bacterial L30; RPL7A has a paralog, RPL7B, that arose from the whole genome duplication. (244 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) | ||||
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) | ||||
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) | ||||
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) | ||||
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) | ||||
TOM1 | E3 ubiquitin ligase of the hect-domain class; has a role in mRNA export from the nucleus and may regulate transcriptional coactivators; involved in degradation of excess histones; interacts with Dia2p and is required for Dia2p degradation; required to target Cdc6p for ubiquitin-mediated destruction during G1 phase; Belongs to the UPL family. TOM1/PTR1 subfamily. (3268 aa) | ||||
RRP17 | Ribosomal RNA-processing protein 17; Component of the pre-60S pre-ribosomal particle; required for cell viability under standard (aerobic) conditions but not under anaerobic conditions; exonuclease required for 5' end processing of pre-60S ribosomal RNA. (235 aa) |