Major pathway of rRNA processing in the nucleolus and cytosol (Homo sapiens)

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1, 13, 15, 19, 21...15, 18, 2010, 12, 28, 29, 3720, 34, 386, 7, 9, 14, 20...8, 17, 27, 30, 39...2, 16, 31, 33-35, 38...40, 46, 48, 50nucleoplasmcytosolRPL35 WDR75 RRP36 UTP14A KRR1 GNL3:EBNA1BP2:DDX21:PES1UTP18 RPS18 NOP14 KRR1 WDR43 PES1 WDR18 RPLP0 RPL14 RRP36 HEATR1 RPS19 NOL11WDR46 RRP9 WDR43 RPS6 DIS3RPS2CIRH1A RPS14 NOL6 SKIV2L2NOP56 CIRH1A CSNK1E 32S pre-rRNAHEATR1 RPS18 FBL DDX47 RPSA TBL3 RPL27A 21S pre-rRNA:SSUprocessomeRPL13A PWP2 RCL1 NOP14 UTP3 RIOK3EXOSC1 NOL6 RPS25 RPL40 PELP1 UTP20 28S rRNA RPL23 RPS4Y2 PES1 RPS23 WDR46 NHP2L1 CIRH1A U3 snoRNA RRP1RPS9 IMP4 RPS7 HEATR1 RPS27 RPS14 RPS9 IMP3 RPS4Y1 UTP15 WDR3 UTP18 PDCD11 UTP20 IMP4 RPS20 PeBoW complexNOP56 RPS19 TEX10 45S pre-rRNA RPP38 FAU WDR36 RPS13UTP18RPL12 RPS23WDR36 WDR75 EXOSC10FCF1 RPL11 TBL3 BOP1PDCD11 RPL23A DIEXF FCF1 NOP56 NHP2L1 UTP11L RPS9 NOP58 NHP2L1 PDCD11 RPS10 NOP56 NOL11WDR3 RPS7 SKIV2L2EXOSC5 U3 snoRNA DDX47 RPL39 RPS24 RPL41 LAS1LRPS8RPS14 DCAF13 UTP14C MPHOSPH10 NOP14 RPS11 NOL9RPL8 RPLP1 UTP20 RPL4 RPS12 RRP7A NOP58 RPL10L 40S ribosomalcomplexRPS11NOP14 RPS2 RRP36 XRN2MPHOSPH10 RPS13 RPS21 WDR43 UTP3 NOP58 RPL36AL RPS28 DHX37 NIP7UTP6 NOB1NOC4L RCL1 SENP3 DDX49 UTP11L RPS27L RPS15ARPL39L RPS9 SKIV2L2UTP15 KRR1 28SrRNA:12Spre-rRNA:Rix1:LAS1L:NOL9:SENP3:PeBoWRPS5 RPL31 RPL3L RPS3 RPL30 DDX49 RPS2 UTP3 DDX52 RPL19 RPL36 WDR12 WBSCR22NOL6 WDR75 BOP1 ERI1RPL35A IMP3 UTP15 nascent pre-rRNA transcript PWP2 XRN2NOC4L C1D NOC4L UTPA complexRPL17 RPL22L1 RPS3 RPS12 EXOSC10EMG1 DHX37 UTP11L RPS7 UTP20 RPL18 MPHOSPH6RPS4X DCAF13 RPS17 CIRH1A UTPC complexFAU RRP9 FTSJ3MPHOSPH10 UTP15 RPS16RPS6 GNL3 DDX49 DIEXF RPL10 NOL12TSR1RPS17 RPS23 RPP30 HEATR1 WDR75 WDR43 TBL3 MPHOSPH10 TBL3 EXOSC9 18SE pre-rRNA:SSUprocessomeRCL1 PNO1 WDR12 C1DRPL26 NCLTBL3 FBL RPS28EMG1 RPS15A RRP9 UTP6 DDX21 RBM28WDR36 UTP14A PWP2 DIEXF EXOSC2 RPL7A WDR75 RRP7A WDR18 RPS16 NOL9 NOC4L NOP14 RPS11 KRR1 RPS27 12S pre-rRNA RPS26 RPP40 EXOSC8 RPP14 RPL26L1 NOL6 UTP3 EXOSC4 UTP14A UTP14A DDX52 DCAF13 UTP6 RPS16 DHX37 UTP14C EXOSC10:C1D:MPHOSPH6:SKIV2L2:ExosomeWDR3 LAS1L RPS2 RRP36 RPL29 RPS29 RRP7A UTP11L RPL38 PNO1EMG1 LTV147S pre-rRNA:SSUprocessomeWDR43 WDR3 BMS1 DDX49 UTP14A RPS2 SKIV2L2 FCF1 EXOSC10 UTP18 DDX52 RPS14 RPS21 WDR36 RPS7 WDR46 UTP15 DDX52 RCL1 RPP25 RPS27LNOL11 RPL10A RPL37A RPS27L DIEXF EXOSC7 RPS14UTP3 WDR46 RRP36 BMS1 RPS9 RIOK2 RPS918SE pre-rRNA RPS28 45S pre-rRNA:SSUprocessomeRPL37 EXOSC3 RPL21 CSNK1DHEATR1 5.8S rRNA 30S pre-rRNA WDR75 CSNK1ERIOK1 DDX52 RPS20 U3 snoRNA RPL18A RRP9 BYSL UTP15 IMP3 RPS3A MPHOSPH6 RPL3 NOL11 DDX47 BMS1 RPS7 RPL24 RPS27A(77-156) UTP6 RPL34 NOL6 DDX47 FBL BOP1 RPS5 U3 snoRNA TSR1 RPS7 ISG20L2IMP4 RIOK2RPL7 CIRH1A UTP11L NOP58 FBL RPL6 RRP7A IMP3 NOB1 RPL36A KRR1 RPS9 5S rRNA RPS6RPS4Y2 NOL6 UTP14C EXOSC6 EMG1 RPL28 CIRH1A NOL9 RPS24 HEATR1 RPS2418SE pre-40SparticleRPL22 RPS14 RPS7 RPL5 UTP18 RPS5RPS10 NOL11 RPS9 RPS15 RRP7A UTP15 UTPA complex32S pre-rRNA WDR75 DDX49 RPS6 EXOSC5RPS8 PES1 RPS2 BMS1 RRP7A RPS14 RPS4Y1 RPS2 RPS26 HEATR1 RPS6 RPS8 FCF1RIOK1RPS25 DDX47 DIEXF CIRH1A WDR46 RPS13 U3 snoRNA RPS3AIMP4 32Spre-rRNA:Rix1:LAS1L:NOL9:SENP3NHP2L1 IMP4 MPHOSPH10 NOP56 TEX10 RPS4X FCF1 PDCD11 WDR36 PELP1 RPS2 DHX37 RRP36 RPL13 DCAF13 FCF1 60S ribosomalcomplexRPS14 RPP21 18S rRNA IMP3 RPS6 RPSA UTP14C DCAF13 BYSLRPL9 EBNA1BP2 UTP14C NOC4L RPS15 EMG1 RPL32 NOP58 RPL15 SENP3 PWP2 RPS27A(77-156) CSNK1D UTP18 NOL11 RPS6 RPLP2 BMS1 RPL27 RPS6 FBL PWP2 RRP9 DHX37 LTV1 RPS15A RPS27WDR3 RPS3A UTP6 18SE pre-rRNA UTP20 21S pre-rRNA NHP2L1 30S pre-rRNA:SSUprocessomeWDR43 28S rRNA RPS7RPS29 PDCD11 RCL1 WDR43 NOL11 LAS1L 2831, 38, 41, 4328283, 118, 17, 27, 42, 452830452828238, 17, 27, 42, 45283, 1152920, 4654


Description

In humans, a 47S precursor rRNA (pre-rRNA) is transcribed by RNA polymerase I from rRNA-encoding genes (rDNA) at the boundary of the fibrillar center and the dense fibrillar components of the nucleolus (Stanek et al. 2001). The 47S precursor is processed over the course of about 5-8 minutes (Popov et al. 2013) by endoribonucleases and exoribonucleases to yield the 28S rRNA and 5.8S rRNA of the 60S subunit and the 18S rRNA of the 40S subunit (reviewed in Mullineus and Lafontaine 2012, Henras et al. 2015). As the pre-rRNA is being transcribed, a large protein complex, the small subunit (SSU) processome, assembles in the region of the 18S rRNA sequence, forming terminal knobs on the pre-rRNA (reviewed in Phipps et al. 2011, inferred from yeast in Dragon et al. 2002). The SSU processome contains both ribosomal proteins of the small subunit and processing factors which process the pre-rRNA and modify nucleotides. Through addition of subunits the SSU processome appears to be converted into the larger 90S pre-ribosome (inferred from yeast in Grandi et al. 2002). An analogous large subunit processome (LSU) assembles in the region of the 28S rRNA, however the LSU is less well characterized (inferred from yeast in McCann et al. 2015).
Following cleavage of the pre-rRNA within internal transcribed spacer 1 (ITS1), the pre-ribosomal particle separates into a pre-60S subunit and a pre-40S subunit in the nucleolus (reviewed in Hernandez-Verdun et al. 2010, Phipps et al. 2011). The pre-60S and pre-40S ribosomal particles are then exported from the nucleus to the cytoplasm where the processing factors dissociate and recycle back to the nucleus
Nuclease digestions of the 47S pre-rRNA can follow several paths. In the major pathway, the ends of the 47S pre-rRNA are trimmed to yield the 45S pre-rRNA. Digestion at site 2 (also called site 2b in mouse, see Henras et al. 2015 for nomenclature) cleaves the 45S pre-rRNA to yield the 30S pre-rRNA containing the 18S rRNA of the small subunit and the 32S pre-rRNA containing the 5.8S rRNA and the 28S rRNA of the large subunit. The 32S pre-rRNA is digested in the nucleus to yield the 5.8S rRNA and the 28S rRNA while the 30S pre-rRNA is digested in the nucleus to yield the 18SE pre-rRNA which is then processed in the nucleus and cytosol to yield the 18S rRNA. At least 286 human proteins, 74 of which have no yeast homolog, are required for efficient processing of pre-rRNA in the nucleus (Tafforeau et al. 2013) View original pathway at Reactome.

Comments

Reactome-Converter 
Pathway is converted from Reactome ID: 6791226
Reactome-version 
Reactome version: 75
Reactome Author 
Reactome Author: May, Bruce

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Bibliography

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History

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CompareRevisionActionTimeUserComment
114630view16:09, 25 January 2021ReactomeTeamReactome version 75
113078view11:13, 2 November 2020ReactomeTeamReactome version 74
112312view15:23, 9 October 2020ReactomeTeamReactome version 73
101211view11:10, 1 November 2018ReactomeTeamreactome version 66
100749view20:35, 31 October 2018ReactomeTeamreactome version 65
100293view19:12, 31 October 2018ReactomeTeamreactome version 64
99839view15:56, 31 October 2018ReactomeTeamreactome version 63
99396view14:34, 31 October 2018ReactomeTeamreactome version 62 (2nd attempt)
99090view12:39, 31 October 2018ReactomeTeamreactome version 62
94494view08:59, 14 September 2017Mkutmonreactome version 61
86967view13:41, 15 July 2016MkutmonOntology Term : 'RNA processing pathway' added !
86396view09:17, 11 July 2016ReactomeTeamNew pathway

External references

DataNodes

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NameTypeDatabase referenceComment
12S pre-rRNA R-HSA-6791201 (Reactome)
18S rRNA ProteinX03205 (EMBL)
18SE pre-40S particleComplexR-HSA-6791531 (Reactome)
18SE pre-rRNA R-HSA-6791204 (Reactome)
18SE pre-rRNA:SSU processomeComplexR-HSA-6791187 (Reactome)
21S pre-rRNA R-HSA-6791194 (Reactome)
21S pre-rRNA:SSU processomeComplexR-HSA-6791199 (Reactome)
28S

rRNA:12S

pre-rRNA:Rix1:LAS1L:NOL9:SENP3:PeBoW		ComplexR-HSA-6791568 (Reactome)
28S rRNA ProteinM11167 (EMBL)
30S pre-rRNA R-HSA-6791193 (Reactome)
30S pre-rRNA:SSU processomeComplexR-HSA-6791185 (Reactome)
32S pre-rRNA:Rix1:LAS1L:NOL9:SENP3ComplexR-HSA-6791562 (Reactome) The PeBoW complex and the Rix1:LAS1L:NOL9:SENP3 complex associate with pre-60S ribosome subunits during maturation and are required for efficient rRNA processing. Note that this complex contains many ribsomal subunit proteins and processing factors but the precise composition is not known.
32S pre-rRNA R-HSA-6791200 (Reactome)
32S pre-rRNAR-HSA-6791200 (Reactome)
40S ribosomal complexComplexR-HSA-72392 (Reactome)
45S pre-rRNA R-HSA-6791208 (Reactome)
45S pre-rRNA:SSU processomeComplexR-HSA-6791211 (Reactome)
47S pre-rRNA:SSU processomeComplexR-HSA-6791184 (Reactome)
5.8S rRNA ProteinJ01866 (EMBL)
5S rRNA ProteinV00589 (EMBL)
60S ribosomal complexComplexR-HSA-72499 (Reactome)
BMS1 ProteinQ14692 (Uniprot-TrEMBL)
BOP1 ProteinQ14137 (Uniprot-TrEMBL)
BOP1ProteinQ14137 (Uniprot-TrEMBL)
BYSL ProteinQ13895 (Uniprot-TrEMBL)
BYSLProteinQ13895 (Uniprot-TrEMBL)
C1D ProteinQ13901 (Uniprot-TrEMBL)
C1DProteinQ13901 (Uniprot-TrEMBL)
CIRH1A ProteinQ969X6 (Uniprot-TrEMBL)
CSNK1D ProteinP48730 (Uniprot-TrEMBL)
CSNK1DProteinP48730 (Uniprot-TrEMBL)
CSNK1E ProteinP49674 (Uniprot-TrEMBL)
CSNK1EProteinP49674 (Uniprot-TrEMBL)
DCAF13 ProteinQ9NV06 (Uniprot-TrEMBL)
DDX21 ProteinQ9NR30 (Uniprot-TrEMBL)
DDX47 ProteinQ9H0S4 (Uniprot-TrEMBL)
DDX49 ProteinQ9Y6V7 (Uniprot-TrEMBL)
DDX52 ProteinQ9Y2R4 (Uniprot-TrEMBL)
DHX37 ProteinQ8IY37 (Uniprot-TrEMBL)
DIEXF ProteinQ68CQ4 (Uniprot-TrEMBL)
DIS3ProteinQ9Y2L1 (Uniprot-TrEMBL)
EBNA1BP2 ProteinQ99848 (Uniprot-TrEMBL)
EMG1 ProteinQ92979 (Uniprot-TrEMBL)
ERI1ProteinQ8IV48 (Uniprot-TrEMBL)
EXOSC1 ProteinQ9Y3B2 (Uniprot-TrEMBL)
EXOSC10 ProteinQ01780 (Uniprot-TrEMBL)
EXOSC10:C1D:MPHOSPH6:SKIV2L2:ExosomeComplexR-HSA-6791581 (Reactome)
EXOSC10ProteinQ01780 (Uniprot-TrEMBL)
EXOSC2 ProteinQ13868 (Uniprot-TrEMBL)
EXOSC3 ProteinQ9NQT5 (Uniprot-TrEMBL)
EXOSC4 ProteinQ9NPD3 (Uniprot-TrEMBL)
EXOSC5 ProteinQ9NQT4 (Uniprot-TrEMBL)
EXOSC5ProteinQ9NQT4 (Uniprot-TrEMBL)
EXOSC6 ProteinQ5RKV6 (Uniprot-TrEMBL)
EXOSC7 ProteinQ15024 (Uniprot-TrEMBL)
EXOSC8 ProteinQ96B26 (Uniprot-TrEMBL)
EXOSC9 ProteinQ06265 (Uniprot-TrEMBL)
FAU ProteinP62861 (Uniprot-TrEMBL)
FBL ProteinP22087 (Uniprot-TrEMBL)
FCF1 ProteinQ9Y324 (Uniprot-TrEMBL)
FCF1ProteinQ9Y324 (Uniprot-TrEMBL)
FTSJ3ProteinQ8IY81 (Uniprot-TrEMBL)
GNL3 ProteinQ9BVP2 (Uniprot-TrEMBL)
GNL3:EBNA1BP2:DDX21:PES1ComplexR-HSA-8869320 (Reactome)
HEATR1 ProteinQ9H583 (Uniprot-TrEMBL)
IMP3 ProteinQ9NV31 (Uniprot-TrEMBL)
IMP4 ProteinQ96G21 (Uniprot-TrEMBL)
ISG20L2ProteinQ9H9L3 (Uniprot-TrEMBL)
KRR1 ProteinQ13601 (Uniprot-TrEMBL)
LAS1L ProteinQ9Y4W2 (Uniprot-TrEMBL)
LAS1LProteinQ9Y4W2 (Uniprot-TrEMBL)
LTV1 ProteinQ96GA3 (Uniprot-TrEMBL)
LTV1ProteinQ96GA3 (Uniprot-TrEMBL)
MPHOSPH10 ProteinO00566 (Uniprot-TrEMBL)
MPHOSPH6 ProteinQ99547 (Uniprot-TrEMBL)
MPHOSPH6ProteinQ99547 (Uniprot-TrEMBL)
NCLProteinP19338 (Uniprot-TrEMBL)
NHP2L1 ProteinP55769 (Uniprot-TrEMBL)
NIP7ProteinQ9Y221 (Uniprot-TrEMBL)
NOB1 ProteinQ9ULX3 (Uniprot-TrEMBL)
NOB1ProteinQ9ULX3 (Uniprot-TrEMBL)
NOC4L ProteinQ9BVI4 (Uniprot-TrEMBL)
NOL11 ProteinQ9H8H0 (Uniprot-TrEMBL)
NOL11ProteinQ9H8H0 (Uniprot-TrEMBL)
NOL12ProteinQ9UGY1 (Uniprot-TrEMBL)
NOL6 ProteinQ9H6R4 (Uniprot-TrEMBL)
NOL9 ProteinQ5SY16 (Uniprot-TrEMBL)
NOL9ProteinQ5SY16 (Uniprot-TrEMBL)
NOP14 ProteinP78316 (Uniprot-TrEMBL)
NOP56 ProteinO00567 (Uniprot-TrEMBL)
NOP58 ProteinQ9Y2X3 (Uniprot-TrEMBL)
PDCD11 ProteinQ14690 (Uniprot-TrEMBL)
PELP1 ProteinQ8IZL8 (Uniprot-TrEMBL)
PES1 ProteinO00541 (Uniprot-TrEMBL)
PNO1 ProteinQ9NRX1 (Uniprot-TrEMBL)
PNO1ProteinQ9NRX1 (Uniprot-TrEMBL)
PWP2 ProteinQ15269 (Uniprot-TrEMBL)
PeBoW complexComplexR-HSA-6791214 (Reactome)
RBM28ProteinQ9NW13 (Uniprot-TrEMBL)
RCL1 ProteinQ9Y2P8 (Uniprot-TrEMBL)
RIOK1 ProteinQ9BRS2 (Uniprot-TrEMBL)
RIOK1ProteinQ9BRS2 (Uniprot-TrEMBL)
RIOK2 ProteinQ9BVS4 (Uniprot-TrEMBL)
RIOK2ProteinQ9BVS4 (Uniprot-TrEMBL)
RIOK3ProteinO14730 (Uniprot-TrEMBL)
RPL10 ProteinP27635 (Uniprot-TrEMBL)
RPL10A ProteinP62906 (Uniprot-TrEMBL)
RPL10L ProteinQ96L21 (Uniprot-TrEMBL)
RPL11 ProteinP62913 (Uniprot-TrEMBL)
RPL12 ProteinP30050 (Uniprot-TrEMBL)
RPL13 ProteinP26373 (Uniprot-TrEMBL)
RPL13A ProteinP40429 (Uniprot-TrEMBL)
RPL14 ProteinP50914 (Uniprot-TrEMBL)
RPL15 ProteinP61313 (Uniprot-TrEMBL)
RPL17 ProteinP18621 (Uniprot-TrEMBL)
RPL18 ProteinQ07020 (Uniprot-TrEMBL)
RPL18A ProteinQ02543 (Uniprot-TrEMBL)
RPL19 ProteinP84098 (Uniprot-TrEMBL)
RPL21 ProteinP46778 (Uniprot-TrEMBL)
RPL22 ProteinP35268 (Uniprot-TrEMBL)
RPL22L1 ProteinQ6P5R6 (Uniprot-TrEMBL)
RPL23 ProteinP62829 (Uniprot-TrEMBL)
RPL23A ProteinP62750 (Uniprot-TrEMBL)
RPL24 ProteinP83731 (Uniprot-TrEMBL)
RPL26 ProteinP61254 (Uniprot-TrEMBL)
RPL26L1 ProteinQ9UNX3 (Uniprot-TrEMBL)
RPL27 ProteinP61353 (Uniprot-TrEMBL)
RPL27A ProteinP46776 (Uniprot-TrEMBL)
RPL28 ProteinP46779 (Uniprot-TrEMBL)
RPL29 ProteinP47914 (Uniprot-TrEMBL)
RPL3 ProteinP39023 (Uniprot-TrEMBL)
RPL30 ProteinP62888 (Uniprot-TrEMBL)
RPL31 ProteinP62899 (Uniprot-TrEMBL)
RPL32 ProteinP62910 (Uniprot-TrEMBL)
RPL34 ProteinP49207 (Uniprot-TrEMBL)
RPL35 ProteinP42766 (Uniprot-TrEMBL)
RPL35A ProteinP18077 (Uniprot-TrEMBL)
RPL36 ProteinQ9Y3U8 (Uniprot-TrEMBL)
RPL36A ProteinP83881 (Uniprot-TrEMBL)
RPL36AL ProteinQ969Q0 (Uniprot-TrEMBL)
RPL37 ProteinP61927 (Uniprot-TrEMBL)
RPL37A ProteinP61513 (Uniprot-TrEMBL)
RPL38 ProteinP63173 (Uniprot-TrEMBL)
RPL39 ProteinP62891 (Uniprot-TrEMBL)
RPL39L ProteinQ96EH5 (Uniprot-TrEMBL)
RPL3L ProteinQ92901 (Uniprot-TrEMBL)
RPL4 ProteinP36578 (Uniprot-TrEMBL)
RPL40 ProteinP62987 (Uniprot-TrEMBL)
RPL41 ProteinP62945 (Uniprot-TrEMBL)
RPL5 ProteinP46777 (Uniprot-TrEMBL)
RPL6 ProteinQ02878 (Uniprot-TrEMBL)
RPL7 ProteinP18124 (Uniprot-TrEMBL)
RPL7A ProteinP62424 (Uniprot-TrEMBL)
RPL8 ProteinP62917 (Uniprot-TrEMBL)
RPL9 ProteinP32969 (Uniprot-TrEMBL)
RPLP0 ProteinP05388 (Uniprot-TrEMBL)
RPLP1 ProteinP05386 (Uniprot-TrEMBL)
RPLP2 ProteinP05387 (Uniprot-TrEMBL)
RPP14 ProteinO95059 (Uniprot-TrEMBL)
RPP21 ProteinQ9H633 (Uniprot-TrEMBL)
RPP25 ProteinQ9BUL9 (Uniprot-TrEMBL)
RPP30 ProteinP78346 (Uniprot-TrEMBL)
RPP38 ProteinP78345 (Uniprot-TrEMBL)
RPP40 ProteinO75818 (Uniprot-TrEMBL)
RPS10 ProteinP46783 (Uniprot-TrEMBL)
RPS11 ProteinP62280 (Uniprot-TrEMBL)
RPS11ProteinP62280 (Uniprot-TrEMBL)
RPS12 ProteinP25398 (Uniprot-TrEMBL)
RPS13 ProteinP62277 (Uniprot-TrEMBL)
RPS13ProteinP62277 (Uniprot-TrEMBL)
RPS14 ProteinP62263 (Uniprot-TrEMBL)
RPS14ProteinP62263 (Uniprot-TrEMBL)
RPS15 ProteinP62841 (Uniprot-TrEMBL)
RPS15A ProteinP62244 (Uniprot-TrEMBL)
RPS15AProteinP62244 (Uniprot-TrEMBL)
RPS16 ProteinP62249 (Uniprot-TrEMBL)
RPS16ProteinP62249 (Uniprot-TrEMBL)
RPS17 ProteinP08708 (Uniprot-TrEMBL)
RPS18 ProteinP62269 (Uniprot-TrEMBL)
RPS19 ProteinP39019 (Uniprot-TrEMBL)
RPS2 ProteinP15880 (Uniprot-TrEMBL)
RPS20 ProteinP60866 (Uniprot-TrEMBL)
RPS21 ProteinP63220 (Uniprot-TrEMBL)
RPS23 ProteinP62266 (Uniprot-TrEMBL)
RPS23ProteinP62266 (Uniprot-TrEMBL)
RPS24 ProteinP62847 (Uniprot-TrEMBL)
RPS24ProteinP62847 (Uniprot-TrEMBL)
RPS25 ProteinP62851 (Uniprot-TrEMBL)
RPS26 ProteinP62854 (Uniprot-TrEMBL)
RPS27 ProteinP42677 (Uniprot-TrEMBL)
RPS27A(77-156) ProteinP62979 (Uniprot-TrEMBL)
RPS27L ProteinQ71UM5 (Uniprot-TrEMBL)
RPS27LProteinQ71UM5 (Uniprot-TrEMBL)
RPS27ProteinP42677 (Uniprot-TrEMBL)
RPS28 ProteinP62857 (Uniprot-TrEMBL)
RPS28ProteinP62857 (Uniprot-TrEMBL)
RPS29 ProteinP62273 (Uniprot-TrEMBL)
RPS2ProteinP15880 (Uniprot-TrEMBL)
RPS3 ProteinP23396 (Uniprot-TrEMBL)
RPS3A ProteinP61247 (Uniprot-TrEMBL)
RPS3AProteinP61247 (Uniprot-TrEMBL)
RPS4X ProteinP62701 (Uniprot-TrEMBL)
RPS4Y1 ProteinP22090 (Uniprot-TrEMBL)
RPS4Y2 ProteinQ8TD47 (Uniprot-TrEMBL)
RPS5 ProteinP46782 (Uniprot-TrEMBL)
RPS5ProteinP46782 (Uniprot-TrEMBL)
RPS6 ProteinP62753 (Uniprot-TrEMBL)
RPS6ProteinP62753 (Uniprot-TrEMBL)
RPS7 ProteinP62081 (Uniprot-TrEMBL)
RPS7ProteinP62081 (Uniprot-TrEMBL)
RPS8 ProteinP62241 (Uniprot-TrEMBL)
RPS8ProteinP62241 (Uniprot-TrEMBL)
RPS9 ProteinP46781 (Uniprot-TrEMBL)
RPS9ProteinP46781 (Uniprot-TrEMBL)
RPSA ProteinP08865 (Uniprot-TrEMBL)
RRP1ProteinP56182 (Uniprot-TrEMBL)
RRP36 ProteinQ96EU6 (Uniprot-TrEMBL)
RRP7A ProteinQ9Y3A4 (Uniprot-TrEMBL)
RRP9 ProteinO43818 (Uniprot-TrEMBL)
SENP3 ProteinQ9H4L4 (Uniprot-TrEMBL)
SKIV2L2 ProteinP42285 (Uniprot-TrEMBL)
SKIV2L2ProteinP42285 (Uniprot-TrEMBL)
TBL3 ProteinQ12788 (Uniprot-TrEMBL)
TEX10 ProteinQ9NXF1 (Uniprot-TrEMBL)
TSR1 ProteinQ2NL82 (Uniprot-TrEMBL)
TSR1ProteinQ2NL82 (Uniprot-TrEMBL)
U3 snoRNA ProteinNR_006880 (NCBI Protein)
UTP11L ProteinQ9Y3A2 (Uniprot-TrEMBL)
UTP14A ProteinQ9BVJ6 (Uniprot-TrEMBL)
UTP14C ProteinQ5TAP6 (Uniprot-TrEMBL)
UTP15 ProteinQ8TED0 (Uniprot-TrEMBL)
UTP18 ProteinQ9Y5J1 (Uniprot-TrEMBL)
UTP18ProteinQ9Y5J1 (Uniprot-TrEMBL)
UTP20 ProteinO75691 (Uniprot-TrEMBL)
UTP3 ProteinQ9NQZ2 (Uniprot-TrEMBL)
UTP6 ProteinQ9NYH9 (Uniprot-TrEMBL)
UTPA complexComplexR-HSA-6790880 (Reactome)
UTPC complexComplexR-HSA-6790889 (Reactome)
WBSCR22ProteinO43709 (Uniprot-TrEMBL)
WDR12 ProteinQ9GZL7 (Uniprot-TrEMBL)
WDR18 ProteinQ9BV38 (Uniprot-TrEMBL)
WDR3 ProteinQ9UNX4 (Uniprot-TrEMBL)
WDR36 ProteinQ8NI36 (Uniprot-TrEMBL)
WDR43 ProteinQ15061 (Uniprot-TrEMBL)
WDR46 ProteinO15213 (Uniprot-TrEMBL)
WDR75 ProteinQ8IWA0 (Uniprot-TrEMBL)
XRN2ProteinQ9H0D6 (Uniprot-TrEMBL)
nascent pre-rRNA transcript ProteinNR_046235 (NCBI Protein)

Annotated Interactions

View all...
SourceTargetTypeDatabase referenceComment
18SE pre-40S particleR-HSA-6791223 (Reactome)
18SE pre-rRNA:SSU processomeArrowR-HSA-6791222 (Reactome)
21S pre-rRNA:SSU processomeArrowR-HSA-6791221 (Reactome)
21S pre-rRNA:SSU processomeR-HSA-6791222 (Reactome)
28S

rRNA:12S

pre-rRNA:Rix1:LAS1L:NOL9:SENP3:PeBoW		ArrowR-HSA-6791219 (Reactome)
28S

rRNA:12S

pre-rRNA:Rix1:LAS1L:NOL9:SENP3:PeBoW		R-HSA-6791218 (Reactome)
30S pre-rRNA:SSU processomeArrowR-HSA-6791228 (Reactome)
30S pre-rRNA:SSU processomeR-HSA-6791221 (Reactome)
32S pre-rRNA:Rix1:LAS1L:NOL9:SENP3R-HSA-6791219 (Reactome)
32S pre-rRNAArrowR-HSA-6791228 (Reactome)
40S ribosomal complexArrowR-HSA-6791223 (Reactome)
45S pre-rRNA:SSU processomeArrowR-HSA-6791227 (Reactome)
45S pre-rRNA:SSU processomeR-HSA-6791228 (Reactome)
47S pre-rRNA:SSU processomeR-HSA-6791227 (Reactome)
60S ribosomal complexArrowR-HSA-6791218 (Reactome)
BOP1ArrowR-HSA-6791228 (Reactome)
BYSLArrowR-HSA-6791223 (Reactome)
C1DArrowR-HSA-6791218 (Reactome)
CSNK1DArrowR-HSA-6791223 (Reactome)
CSNK1EArrowR-HSA-6791223 (Reactome)
DIS3ArrowR-HSA-6791218 (Reactome)
ERI1ArrowR-HSA-6791218 (Reactome)
EXOSC10:C1D:MPHOSPH6:SKIV2L2:Exosomemim-catalysisR-HSA-6791222 (Reactome)
EXOSC10ArrowR-HSA-6791218 (Reactome)
EXOSC10ArrowR-HSA-6791222 (Reactome)
EXOSC10ArrowR-HSA-6791228 (Reactome)
EXOSC5ArrowR-HSA-6791222 (Reactome)
FCF1ArrowR-HSA-6791221 (Reactome)
FTSJ3ArrowR-HSA-6791221 (Reactome)
GNL3:EBNA1BP2:DDX21:PES1ArrowR-HSA-6791219 (Reactome)
ISG20L2ArrowR-HSA-6791218 (Reactome)
LAS1LArrowR-HSA-6791219 (Reactome)
LTV1ArrowR-HSA-6791223 (Reactome)
MPHOSPH6ArrowR-HSA-6791218 (Reactome)
NCLArrowR-HSA-6791227 (Reactome)
NIP7ArrowR-HSA-6791221 (Reactome)
NOB1ArrowR-HSA-6791223 (Reactome)
NOB1mim-catalysisR-HSA-6791223 (Reactome)
NOL11ArrowR-HSA-6791221 (Reactome)
NOL11ArrowR-HSA-6791227 (Reactome)
NOL12ArrowR-HSA-6791228 (Reactome)
NOL9ArrowR-HSA-6791219 (Reactome)
PNO1ArrowR-HSA-6791223 (Reactome)
PeBoW complexArrowR-HSA-6791219 (Reactome)
R-HSA-6791218 (Reactome) The 12S pre-rRNA is nucleolytically cleaved to yield 5.8S rRNA. C1D, MPHOSPH6 (MPP6), and EXOSC10 (Pm/Scl-100) of the exosome associate and, together with SKIV2L2 (MTR4), are required for 3' processing of mature 5.8S rRNA therefore the exonuclease activity of the exosome seems to be involved (Schilders et al. 2007). Similarly, the DIS3 subunit of the nuclear exosome is required for production of 5.8S rRNA (Tomecki et al. 2010) and the 3'-5' exoribonuclease ISG20L2 (Coute et al. 2008) are also required for production of 5.8S rRNA from 12S pre-rRNA. As inferred from the mouse homolog, the 3'-5' exonuclease ERI1 also plays a role in trimming the 3' end of pre-5.8S rRNA (Ansel et al. 2008).
R-HSA-6791219 (Reactome) Unknown nucleases process the 32S precursor rRNA (pre-rRNA) at site 4, yielding the 28S rRNA and the 12S pre-RNA that will be further processed to the mature 5.8S rRNA. Processing occurs in precursor 60S ribosomal (pre-60S) subunits that contain ribosomal proteins and processing factors. The PES1:BOP1:WDR12 complex (PeBoW complex) associates with pre-60S subunits in both the nucleus and cytosol where it is involved in processing 32S rRNA and recycling pre-60S subunit processing factors. Perturbation of the PeBoW complex prevents processing of the 32S pre-rRNA (Holzel et al. 2005, Grimm et al. 2006, Holzel et al 2007, Rohrmoser et al. 2007). The polynucleotide kinase activity of NOL9, which is associated with pre-60s subunits, is also required for processing of pre-32S rRNA (Heindl and Martinez 2010). LAS1L interacts with PELP1:TEX10:WDR18, NOL9, and SENP3 in pre-60S subunits where it is required for processing of the internal transcribed spacer 2 (ITS2) in pre-32S rRNA (Castle et al. 2012). The PELP1:TEX10:WDR18 complex is the mammalian homolog of the yeast Rix1 complex (Castle et al. 2012). A complex containing GNL3 (Nucleostemin), EBNA1BP2, DDX21, and PES1 is also required for processing 32S rRNA to 28S rRNA (Romanova et al. 2009). Both the Nucleostemin complex and the PeBoW complex both contain PES1 and therefore may be part of a single larger complex.
R-HSA-6791221 (Reactome) An unknown endonuclease cleaves site 1 (also called site A1), removing the remainder of the 5' external transcribed spacer (5' ETS) from 30S pre-rRNA containing the 18S rRNA (Freed et al. 2012, Tomecki et al. 2015). Sixteen proteins of the small ribosomal subunit are required for processing sequences flanking the 18S rRNA in the 30S pre-rRNA (O'Donohue et al. 2010) presumably due to their assembly onto pre-rRNA as processing proceeds. Additionally, FCF1 (hUTP24, part of the SSU processome) is required for cleavage at site 1 (Tomecki et al. 2015) and NOL11 and CIRH1A (Cirhin, part of the UTPA complex) interact and are required for cleavage at site 1 and other sites (A', A0, and 2) (Freed et al. 2012). FTSJ3 and NIP7 interact and are required for processing at sites 1, 2, and A0 (Morello et al. 2011).
R-HSA-6791222 (Reactome) An unknown endonuclease in the nucleolus cleaves at site E (site 2a in mouse) of 21S pre-rRNA, yielding 18SE pre-rRNA (Preti et al. 2013, Sloan et al. 2013). Evidence also indicates that 18SE may also be produced by an exonucleolytic pathway (Carron et al. 2011, Sloan et al. 2013). BYSL (Bystin, ENG1), SKIV2L2 (MTR4), and the exonuclease activity of EXOSC10 (RRP6), all associated with the exosome, are required for formation of 18SE by the exonucleolytic pathway (Sloan et al. 2013). Sequencing indicates that 18SE molecules can have variable ends, presumably due to exonuclease activity (Preti et al. 2013).
R-HSA-6791223 (Reactome) Exonuclease activity of the exosome (Preti et al. 2013) and endonuclease activity of NOB1 (inferred from yeast, Pertschy et al. 2009) process the 3' end of precursor rRNA (pre-rRNA) 18SE to yield mature 18S rRNA. During the processing, pre-rRNA 18SE is bound in the pre-40S ribosome subunit, which contains ribosomal proteins and processing factors such as NOB1 and BYSL. The pre-40S subunit is exported from the nucleus to the cytosol where processing factors are released and recycled back to the nucleus. The kinases RIOK1, RIOK2, CSNK1D and CSNK1E are associated with the pre-40S rRNA subunit in both the nucleus and cytosol and their kinase activity is required for recycling of processing factors back to the nucleus (Zemp et al. 2009, Zemp et al. 2014). RIOK1 and RIOK2 are also required for 18SE processing (Widmann et al. 2012, Zemp et al. 2009). RIOK3 (RIO3) is a cytosolic kinase that associates with the pre-40S ribosomal particle after export from the nucleus and is required for release of processing factors (Baumas et al. 2012).
R-HSA-6791227 (Reactome) Unknown nucleases concomitantly cleave the 47S precursor rRNA (pre-rRNA) at the A' site (also known as the 01 site or the A1 site), the A0 site in the 5' external transcribed spacer (5' ETS), and site 02 (also known as site 6 in mouse) in the 3' ETS (Sloan et al. 2014). Cleavage occurs when the pre-rRNA is complexed with the small subunit processome (SSU processome) complex, a large protein complex that binds the 5' region of the pre-rRNA after transcription commences (Kass and Sollner-Webb 1990, Sloan et al. 2014, inferred from yeast in Dragon et al. 2002). The UTP-A subcomplex of the SSU processome and SKIV2L2 (MTR4) are required for cleavage at the A' site while the UTP-B subcomplex and U3 snoRNP (Sloan et al. 2014) and RRP36 of the UTPC subcomplex of the SSU processome (Gerus et al. 2010) improve efficiency of cleavage. UTP18 is required for cleavage of the 5' ETS (Holzel et al. 2010). Nucleolin (NCL) interacts with the 47S pre-rRNA (Yanagida et al. 2001, inferred from mouse in Ginisty et al. 1998) and is involved in cleavage at the A' site (inferred from mouse in Ginisty et al. 1998) but its association with the SSU processome is transitory (Turner et al. 2009). NOL11, a component of the SSU processome which interacts with UTP4, increases the efficiency of processing at A', but is not strictly required (Freed et al. 2012). XRN2 exonucleolytically degrades the 5' 01 fragment (Sloan et al. 2013, inferred from mouse homologs in Wang and Pestov 2011).
R-HSA-6791228 (Reactome) An unknown endonuclease cleaves at site 2 (also called site 2b in mouse) in the internal transcribed spacer 1 (ITS1) between the 18S rRNA and the 5.8S rRNA in the 45S precursor rRNA (pre-rRNA) while the pre-rRNA is contained in a 90S particle containing ribosomal proteins and assembly factors (Sloan et al. 2013). (The 90S particle is believed to be produced by addition of further subunits to the complex containing the pre-RNA and the small subunit (SSU) processome (inferred from yeast in Grandi et al. 2002).) The products are a 30S pre-rRNA which contains the 18S rRNA and a 32S pre-rRNA containing the 5.8S rRNA and the 28S rRNA. The cleavage splits the 90S particle into a pre-40S particle and a pre-60S particle and is believed to occur while the 5' region of the 45S rRNA is bound by the SSU processome. BOP1 (a subunit of the PeBoW complex), RBM28, NOL12 and RRP1 (NOP52) also participate in the cleavage (Sloan et al. 2013, Yoshikawa et al. 2015). Exonucleases including XRN2 and EXOSC10 (RRP6) of the exosome complex then remove further nucleotides from the end of the ITS (Sloan et al. 2013).
RBM28ArrowR-HSA-6791228 (Reactome)
RIOK1ArrowR-HSA-6791223 (Reactome)
RIOK2ArrowR-HSA-6791223 (Reactome)
RIOK3ArrowR-HSA-6791223 (Reactome)
RIOK3R-HSA-6791223 (Reactome)
RPS11ArrowR-HSA-6791221 (Reactome)
RPS13ArrowR-HSA-6791221 (Reactome)
RPS14ArrowR-HSA-6791221 (Reactome)
RPS15AArrowR-HSA-6791221 (Reactome)
RPS16ArrowR-HSA-6791221 (Reactome)
RPS23ArrowR-HSA-6791221 (Reactome)
RPS24ArrowR-HSA-6791221 (Reactome)
RPS27ArrowR-HSA-6791221 (Reactome)
RPS27LArrowR-HSA-6791221 (Reactome)
RPS28ArrowR-HSA-6791221 (Reactome)
RPS2ArrowR-HSA-6791221 (Reactome)
RPS3AArrowR-HSA-6791221 (Reactome)
RPS5ArrowR-HSA-6791221 (Reactome)
RPS6ArrowR-HSA-6791221 (Reactome)
RPS7ArrowR-HSA-6791221 (Reactome)
RPS8ArrowR-HSA-6791221 (Reactome)
RPS9ArrowR-HSA-6791221 (Reactome)
RRP1ArrowR-HSA-6791228 (Reactome)
SKIV2L2ArrowR-HSA-6791218 (Reactome)
SKIV2L2ArrowR-HSA-6791222 (Reactome)
SKIV2L2ArrowR-HSA-6791227 (Reactome)
TSR1ArrowR-HSA-6791223 (Reactome)
UTP18ArrowR-HSA-6791227 (Reactome)
UTPA complexArrowR-HSA-6791221 (Reactome)
UTPA complexArrowR-HSA-6791227 (Reactome)
UTPC complexArrowR-HSA-6791227 (Reactome)
WBSCR22ArrowR-HSA-6791223 (Reactome)
XRN2ArrowR-HSA-6791228 (Reactome)
XRN2mim-catalysisR-HSA-6791227 (Reactome)
Retrieved from "https://classic.wikipathways.org/index.php/Pathway:WP3797"
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