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

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4, 5, 10, 14, 27...8, 13, 24, 30, 38...6, 9, 11, 14, 16...17, 18, 2017, 26, 271, 7, 11, 23, 3315, 34, 402, 3, 12, 18, 20...nucleoplasmcytosolNOB1RPL37 ISG20L245S pre-rRNA:SSUprocessomeMPHOSPH6 RPS10 RPL13 IMP4 RPS9 RPS3 NOP56 RPS9 RPL37A RPL41 FBL IMP3 RPP14 RPS23 18SE pre-rRNA RRP9 RPS25 RPS26 RIOK1EXOSC5 RPS27 RPL34 RPS27A(77-156) RPL29 RPL8 TEX10 RPL22 RPS15A NOP56 21S pre-rRNA NOP58 FBL RPL30 CSNK1ERPL7 RPS24 NOB1NOP58 IMP3 21S pre-rRNA:SSUprocessomeNHP2L1 RPL10A IMP3 SENP3 RPL35 RPS13 RPS4X XRN2MPHOSPH10 EXOSC7 18SE pre-rRNA:SSUprocessomeRPS12 IMP3 RPS2 RCL1 RPS27L RPS28 RPL6 RCL1 RPL14 RPL5 RPS17 IMP4 12S pre-rRNA LTV1 RPL4 RPS7 RPS15 RPS16 EXOSC10 MPHOSPH10 RPS17 PNO1RPS11 RPL27 RIOK2RPSA NHP2L1 LAS1L RPL17 FAU RPL18 RRP9 NOP56 RPL31 EXOSC4 PES1 28S rRNA FBL 28S rRNA RPL15 RPLP0 RPS20 RCL1 RPP38 RPS8 30S pre-rRNA:SSUprocessomeRPP30 EXOSC3 RPL10 TEX10 RPL32 RPP40 RPLP2 RPS28 RPLP1 NHP2L1 IMP4 RPL26L1 RPP21 TSR1 RPS14 EXOSC9 RPL23A RPS3A RPS12 RPL27A PELP1 RRP9 RPS6 18S rRNA RPS19 RPS4Y1 IMP4 RPS5 FAU RPS3A RPS3 RPL26 RPL18A RPS21 BMS1 RPL3 NHP2L1 RPL12 RPL36A RPS15 RPS27A(77-156) RPS18 NOP58 CSNK1D32Spre-rRNA:Rix1:LAS1L:NOL9:SENP35.8S rRNA RPS14 RPS11 RPS29 WDR18 PELP1 LAS1L BOP1 EXOSC2 RPL40 RPS8 TSR132S pre-rRNA 60S ribosomalcomplexMPHOSPH10 RPS24 C1D SENP3 RPS15A 18SE pre-rRNA RPS26 RPL24 RPS23 WDR12 LTV1RPS2 NOP58 NOP56 RPS6 RPS25 CSNK1E BYSLRPL19 RPS20 RIOK2 EXOSC6 RPS13 CSNK1D BMS1 SKIV2L2 RPS18 NOL9 BYSL RPL21 RPS16 RCL1 RPL35A EXOSC1 RPL38 RPL3L PNO1 NOB1 RPL11 EXOSC10:C1D:MPHOSPH6:SKIV2L2:ExosomeRIOK1 BMS1 RIOK3RPL28 RPS5 EXOSC8 RPL7A RPS4Y1 32S pre-rRNARPL9 RPS7 RPS10 18SE pre-40SparticleRPP25 RPL39 WDR18 RPS29 RPS21 RPS27 RPSA RPL23 5S rRNA RPS19 NOL9 FBL MPHOSPH10 28SrRNA:12Spre-rRNA:Rix1:LAS1L:NOL9:SENP3:PeBoWBMS1 RPS4X RPL13A RPL36 47S pre-rRNA:SSUprocessome40S ribosomalcomplexRRP9 8, 24, 30, 38, 3917, 40218, 24, 30, 38, 392, 3, 18, 452521


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.

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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: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: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)
BYSL ProteinQ13895 (Uniprot-TrEMBL)
BYSLProteinQ13895 (Uniprot-TrEMBL)
C1D ProteinQ13901 (Uniprot-TrEMBL)
CSNK1D ProteinP48730 (Uniprot-TrEMBL)
CSNK1DProteinP48730 (Uniprot-TrEMBL)
CSNK1E ProteinP49674 (Uniprot-TrEMBL)
CSNK1EProteinP49674 (Uniprot-TrEMBL)
EXOSC1 ProteinQ9Y3B2 (Uniprot-TrEMBL)
EXOSC10 ProteinQ01780 (Uniprot-TrEMBL)
EXOSC10:C1D:MPHOSPH6:SKIV2L2:ExosomeComplexR-HSA-6791581 (Reactome)
EXOSC2 ProteinQ13868 (Uniprot-TrEMBL)
EXOSC3 ProteinQ9NQT5 (Uniprot-TrEMBL)
EXOSC4 ProteinQ9NPD3 (Uniprot-TrEMBL)
EXOSC5 ProteinQ9NQT4 (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)
IMP3 ProteinQ9NV31 (Uniprot-TrEMBL)
IMP4 ProteinQ96G21 (Uniprot-TrEMBL)
ISG20L2ProteinQ9H9L3 (Uniprot-TrEMBL)
LAS1L ProteinQ9Y4W2 (Uniprot-TrEMBL)
LTV1 ProteinQ96GA3 (Uniprot-TrEMBL)
LTV1ProteinQ96GA3 (Uniprot-TrEMBL)
MPHOSPH10 ProteinO00566 (Uniprot-TrEMBL)
MPHOSPH6 ProteinQ99547 (Uniprot-TrEMBL)
NHP2L1 ProteinP55769 (Uniprot-TrEMBL)
NOB1 ProteinQ9ULX3 (Uniprot-TrEMBL)
NOB1ProteinQ9ULX3 (Uniprot-TrEMBL)
NOL9 ProteinQ5SY16 (Uniprot-TrEMBL)
NOP56 ProteinO00567 (Uniprot-TrEMBL)
NOP58 ProteinQ9Y2X3 (Uniprot-TrEMBL)
PELP1 ProteinQ8IZL8 (Uniprot-TrEMBL)
PES1 ProteinO00541 (Uniprot-TrEMBL)
PNO1 ProteinQ9NRX1 (Uniprot-TrEMBL)
PNO1ProteinQ9NRX1 (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)
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)
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)
RPL37 ProteinP61927 (Uniprot-TrEMBL)
RPL37A ProteinP61513 (Uniprot-TrEMBL)
RPL38 ProteinP63173 (Uniprot-TrEMBL)
RPL39 ProteinP62891 (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)
RPS12 ProteinP25398 (Uniprot-TrEMBL)
RPS13 ProteinP62277 (Uniprot-TrEMBL)
RPS14 ProteinP62263 (Uniprot-TrEMBL)
RPS15 ProteinP62841 (Uniprot-TrEMBL)
RPS15A ProteinP62244 (Uniprot-TrEMBL)
RPS16 ProteinP62249 (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)
RPS24 ProteinP62847 (Uniprot-TrEMBL)
RPS25 ProteinP62851 (Uniprot-TrEMBL)
RPS26 ProteinP62854 (Uniprot-TrEMBL)
RPS27 ProteinP42677 (Uniprot-TrEMBL)
RPS27A(77-156) ProteinP62979 (Uniprot-TrEMBL)
RPS27L ProteinQ71UM5 (Uniprot-TrEMBL)
RPS28 ProteinP62857 (Uniprot-TrEMBL)
RPS29 ProteinP62273 (Uniprot-TrEMBL)
RPS3 ProteinP23396 (Uniprot-TrEMBL)
RPS3A ProteinP61247 (Uniprot-TrEMBL)
RPS4X ProteinP62701 (Uniprot-TrEMBL)
RPS4Y1 ProteinP22090 (Uniprot-TrEMBL)
RPS5 ProteinP46782 (Uniprot-TrEMBL)
RPS6 ProteinP62753 (Uniprot-TrEMBL)
RPS7 ProteinP62081 (Uniprot-TrEMBL)
RPS8 ProteinP62241 (Uniprot-TrEMBL)
RPS9 ProteinP46781 (Uniprot-TrEMBL)
RPSA ProteinP08865 (Uniprot-TrEMBL)
RRP9 ProteinO43818 (Uniprot-TrEMBL)
SENP3 ProteinQ9H4L4 (Uniprot-TrEMBL)
SKIV2L2 ProteinP42285 (Uniprot-TrEMBL)
TEX10 ProteinQ9NXF1 (Uniprot-TrEMBL)
TSR1 ProteinQ2NL82 (Uniprot-TrEMBL)
TSR1ProteinQ2NL82 (Uniprot-TrEMBL)
U3 snoRNA Protein
WDR12 ProteinQ9GZL7 (Uniprot-TrEMBL)
WDR18 ProteinQ9BV38 (Uniprot-TrEMBL)
XRN2ProteinQ9H0D6 (Uniprot-TrEMBL)
nascent pre-rRNA transcript Protein

Annotated Interactions

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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)
BYSLArrowR-HSA-6791223 (Reactome)
CSNK1DArrowR-HSA-6791223 (Reactome)
CSNK1Dmim-catalysisR-HSA-6791223 (Reactome)
CSNK1EArrowR-HSA-6791223 (Reactome)
CSNK1Emim-catalysisR-HSA-6791223 (Reactome)
EXOSC10:C1D:MPHOSPH6:SKIV2L2:Exosomemim-catalysisR-HSA-6791218 (Reactome)
EXOSC10:C1D:MPHOSPH6:SKIV2L2:Exosomemim-catalysisR-HSA-6791222 (Reactome)
ISG20L2mim-catalysisR-HSA-6791218 (Reactome)
LTV1ArrowR-HSA-6791223 (Reactome)
NOB1ArrowR-HSA-6791223 (Reactome)
NOB1mim-catalysisR-HSA-6791223 (Reactome)
PNO1ArrowR-HSA-6791223 (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.
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).
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).
RIOK1ArrowR-HSA-6791223 (Reactome)
RIOK2ArrowR-HSA-6791223 (Reactome)
RIOK3ArrowR-HSA-6791223 (Reactome)
RIOK3R-HSA-6791223 (Reactome)
TSR1ArrowR-HSA-6791223 (Reactome)
XRN2mim-catalysisR-HSA-6791223 (Reactome)
XRN2mim-catalysisR-HSA-6791227 (Reactome)
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