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

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5, 10, 12, 17, 19...21, 30, 47, 4920, 37, 458, 37, 404, 7, 8, 22, 33...1, 9, 18, 25, 28...3, 5, 6, 15, 31...2, 16, 24, 29, 35cytosolnucleoplasm5.8S rRNA HEATR1 GNL3:EBNA1BP2:DDX21:PES1RPL8 FBL RPS3A UTPC complexRPL27A RPS20 RPL7 EBNA1BP2 CSNK1D RPS17 RPS28 UTPA complexRPS27 RPS16TEX10 RPL18 IMP4 TSR1RRP9 RPS10 RPL32 SENP3 NOL9 RPL9 DDX21 WDR18 RPS20 RPS26 IMP4 RPS6RPS7RPP14 RPS15 RPL36A RPS27RCL1 LTV1RPS3 RPS28RPL3L NHP2L1 RPL12 RPS27LRPS13 RPS23 RPSA XRN2RPL27 MPHOSPH6 LAS1L NOP56 RPL35A NHP2L1 NOL9 RPS3ARRP9 EXOSC4 RPS12 EXOSC3 LAS1L RRP9 RPL37 DIS3RPS25 RPL19 PES1 EXOSC7 NOL11RPL39 RPLP1 BMS1 BYSL WDR75 RPL28 RCL1 45S pre-rRNA:SSUprocessomeRPS14 18SE pre-40SparticleNCLSKIV2L2RPS4Y2 SKIV2L2 RPL36AL RPL17 RPL39L CSNK1DSENP3 RPL18A BYSL5S rRNA RPL24 NOP58 RPS13 PeBoW complexRPS28 RPS9 BOP1WDR43 IMP4 EXOSC10RPS8 IMP3 RPL14 WDR12 RPS6 RPS2NOP58 RPS21 RPL41 RPS15 HEATR1 WDR18 NOP56 RPS2 21S pre-rRNA RBM28RPS24 EXOSC2 RPL13 RPL37A RPL22L1 RPS23 RPS16 NOP58 NHP2L1 RPS4Y1 ERI1UTPA complexRPLP2 PNO1 SKIV2L2RPS11 EXOSC1 RPS4X SKIV2L2RPL10L 28S rRNA FBL IMP4 RPL38 RPS17 RPS23LAS1LRPS10 RPS27A(77-156) RPL29 FBL CSNK1E60S ribosomalcomplexEXOSC10:C1D:MPHOSPH6:SKIV2L2:ExosomeRPS24RPS27 RPS6 WBSCR22RPS2 EXOSC6 32Spre-rRNA:Rix1:LAS1L:NOL9:SENP3RRP7A RCL1 32S pre-rRNA RIOK2RPS26 RRP1RPL23A RPS12 LTV1 28SrRNA:12Spre-rRNA:Rix1:LAS1L:NOL9:SENP3:PeBoWRPL35 UTP18FBL RPS4Y1 RPS11RPS8NOP56 RPS9UTP15 RPS24 32S pre-rRNANOL11FAU RPL4 NOP56 PNO130S pre-rRNA:SSUprocessome18SE pre-rRNA NOL9RPS19 RPL34 RRP9 RPS18 RPS4X RPS27A(77-156) RPS29 FAU NOP58 RPS5RPS7 RPL31 NHP2L1 RPS7 MPHOSPH6RPS9 RPL22 RPS16 RIOK2 RPS5 RPL10A IMP3 RPS15A RPP38 RPL13A WDR12 RIOK3CIRH1A RPP40 RPP21 RPL30 18SE pre-rRNA:SSUprocessomeRPS5 18S rRNA EXOSC9 NOL6 RPS15A RPL40 TEX10 RPL21 RPL26L1 47S pre-rRNA:SSUprocessome21S pre-rRNA:SSUprocessomeFCF1BMS1 ISG20L2NIP7RPS21 MPHOSPH10 RPS3 PELP1 CIRH1A RPS13PES1 XRN2RPP30 RPL11 PES1 EXOSC8 RPS14 WDR43 WDR75 RPP25 PELP1 RPSA RPS27L RIOK1 UTP15 C1D RPS29 RPS18 RPS25 EXOSC10 RPL6 MPHOSPH10 RPS11 MPHOSPH10 40S ribosomalcomplexRPL3 FTSJ3RPL10 RPS3A RPS14IMP3 GNL3 BMS1 EXOSC5BOP1 TSR1 RPL7A MPHOSPH10 C1DRPL36 RPS19 EXOSC10RPL15 EXOSC5 RPS8 NOB1NOB1 RPL23 RPS4Y2 RPL26 RPS27L 18SE pre-rRNA RRP36 RCL1 RPS15A28S rRNA 12S pre-rRNA RPL5 IMP3 BOP1 RIOK1RPLP0 NOL12CSNK1E BMS1 1122827251, 9, 18, 28, 438, 22, 33, 393613, 143613, 141, 9, 18, 28, 43353530, 37


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.


Pathway is converted from Reactome ID: 6791226
Reactome version: 66
Reactome Author 
Reactome Author: May, Bruce

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  16. Morello LG, Hesling C, Coltri PP, Castilho BA, Rimokh R, Zanchin NI.; ''The NIP7 protein is required for accurate pre-rRNA processing in human cells.''; PubMed Europe PMC
  17. Popov A, Smirnov E, Kováčik L, Raška O, Hagen G, Stixová L, Raška I.; ''Duration of the first steps of the human rRNA processing.''; PubMed Europe PMC
  18. Hölzel M, Rohrmoser M, Schlee M, Grimm T, Harasim T, Malamoussi A, Gruber-Eber A, Kremmer E, Hiddemann W, Bornkamm GW, Eick D.; ''Mammalian WDR12 is a novel member of the Pes1-Bop1 complex and is required for ribosome biogenesis and cell proliferation.''; PubMed Europe PMC
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  20. Grandi P, Rybin V, Bassler J, Petfalski E, Strauss D, Marzioch M, Schäfer T, Kuster B, Tschochner H, Tollervey D, Gavin AC, Hurt E.; ''90S pre-ribosomes include the 35S pre-rRNA, the U3 snoRNP, and 40S subunit processing factors but predominantly lack 60S synthesis factors.''; PubMed Europe PMC
  21. Couté Y, Kindbeiter K, Belin S, Dieckmann R, Duret L, Bezin L, Sanchez JC, Diaz JJ.; ''ISG20L2, a novel vertebrate nucleolar exoribonuclease involved in ribosome biogenesis.''; PubMed Europe PMC
  22. Widmann B, Wandrey F, Badertscher L, Wyler E, Pfannstiel J, Zemp I, Kutay U.; ''The kinase activity of human Rio1 is required for final steps of cytoplasmic maturation of 40S subunits.''; PubMed Europe PMC
  23. Stanek D, Koberna K, Pliss A, Malínský J, Masata M, Vecerová J, Risueño MC, Raska I.; ''Non-isotopic mapping of ribosomal RNA synthesis and processing in the nucleolus.''; PubMed Europe PMC
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  26. Hernandez-Verdun D, Roussel P, Thiry M, Sirri V, Lafontaine DL.; ''The nucleolus: structure/function relationship in RNA metabolism.''; PubMed Europe PMC
  27. Zhou GJ, Zhang Y, Wang J, Guo JH, Ni J, Zhong ZM, Wang LQ, Dang YJ, Dai JF, Yu L.; ''Cloning and characterization of a novel human RNA binding protein gene PNO1.''; PubMed Europe PMC
  28. Rohrmoser M, Hölzel M, Grimm T, Malamoussi A, Harasim T, Orban M, Pfisterer I, Gruber-Eber A, Kremmer E, Eick D.; ''Interdependence of Pes1, Bop1, and WDR12 controls nucleolar localization and assembly of the PeBoW complex required for maturation of the 60S ribosomal subunit.''; PubMed Europe PMC
  29. O'Donohue MF, Choesmel V, Faubladier M, Fichant G, Gleizes PE.; ''Functional dichotomy of ribosomal proteins during the synthesis of mammalian 40S ribosomal subunits.''; PubMed Europe PMC
  30. Schilders G, van Dijk E, Pruijn GJ.; ''C1D and hMtr4p associate with the human exosome subunit PM/Scl-100 and are involved in pre-rRNA processing.''; PubMed Europe PMC
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  32. Ginisty H, Amalric F, Bouvet P.; ''Nucleolin functions in the first step of ribosomal RNA processing.''; PubMed Europe PMC
  33. Wyler E, Zimmermann M, Widmann B, Gstaiger M, Pfannstiel J, Kutay U, Zemp I.; ''Tandem affinity purification combined with inducible shRNA expression as a tool to study the maturation of macromolecular assemblies.''; PubMed Europe PMC
  34. Wang M, Pestov DG.; ''5'-end surveillance by Xrn2 acts as a shared mechanism for mammalian pre-rRNA maturation and decay.''; PubMed Europe PMC
  35. Freed EF, Prieto JL, McCann KL, McStay B, Baserga SJ.; ''NOL11, implicated in the pathogenesis of North American Indian childhood cirrhosis, is required for pre-rRNA transcription and processing.''; PubMed Europe PMC
  36. Haindl M, Harasim T, Eick D, Muller S.; ''The nucleolar SUMO-specific protease SENP3 reverses SUMO modification of nucleophosmin and is required for rRNA processing.''; PubMed Europe PMC
  37. Sloan KE, Mattijssen S, Lebaron S, Tollervey D, Pruijn GJ, Watkins NJ.; ''Both endonucleolytic and exonucleolytic cleavage mediate ITS1 removal during human ribosomal RNA processing.''; PubMed Europe PMC
  38. Haag S, Kretschmer J, Bohnsack MT.; ''WBSCR22/Merm1 is required for late nuclear pre-ribosomal RNA processing and mediates N7-methylation of G1639 in human 18S rRNA.''; PubMed Europe PMC
  39. Zemp I, Wild T, O'Donohue MF, Wandrey F, Widmann B, Gleizes PE, Kutay U.; ''Distinct cytoplasmic maturation steps of 40S ribosomal subunit precursors require hRio2.''; PubMed Europe PMC
  40. Preti M, O'Donohue MF, Montel-Lehry N, Bortolin-Cavaillé ML, Choesmel V, Gleizes PE.; ''Gradual processing of the ITS1 from the nucleolus to the cytoplasm during synthesis of the human 18S rRNA.''; PubMed Europe PMC
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  42. Tafforeau L, Zorbas C, Langhendries JL, Mullineux ST, Stamatopoulou V, Mullier R, Wacheul L, Lafontaine DL.; ''The complexity of human ribosome biogenesis revealed by systematic nucleolar screening of Pre-rRNA processing factors.''; PubMed Europe PMC
  43. Castle CD, Cassimere EK, Denicourt C.; ''LAS1L interacts with the mammalian Rix1 complex to regulate ribosome biogenesis.''; PubMed Europe PMC
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  46. Hölzel M, Orban M, Hochstatter J, Rohrmoser M, Harasim T, Malamoussi A, Kremmer E, Längst G, Eick D.; ''Defects in 18 S or 28 S rRNA processing activate the p53 pathway.''; PubMed Europe PMC
  47. Tomecki R, Tomecki R, Kristiansen MS, Lykke-Andersen S, Chlebowski A, Larsen KM, Szczesny RJ, Drazkowska K, Pastula A, Andersen JS, Stepien PP, Dziembowski A, Jensen TH.; ''The human core exosome interacts with differentially localized processive RNases: hDIS3 and hDIS3L.''; PubMed Europe PMC
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  49. Ansel KM, Pastor WA, Rath N, Lapan AD, Glasmacher E, Wolf C, Smith LC, Papadopoulou N, Lamperti ED, Tahiliani M, Ellwart JW, Shi Y, Kremmer E, Rao A, Heissmeyer V.; ''Mouse Eri1 interacts with the ribosome and catalyzes 5.8S rRNA processing.''; PubMed Europe PMC
  50. Heindl K, Martinez J.; ''Nol9 is a novel polynucleotide 5'-kinase involved in ribosomal RNA processing.''; PubMed Europe PMC


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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


View all...
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)


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)
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)
DDX21 ProteinQ9NR30 (Uniprot-TrEMBL)
DIS3ProteinQ9Y2L1 (Uniprot-TrEMBL)
EBNA1BP2 ProteinQ99848 (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)
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)
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)
NOL11ProteinQ9H8H0 (Uniprot-TrEMBL)
NOL12ProteinQ9UGY1 (Uniprot-TrEMBL)
NOL6 ProteinQ9H6R4 (Uniprot-TrEMBL)
NOL9 ProteinQ5SY16 (Uniprot-TrEMBL)
NOL9ProteinQ5SY16 (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)
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)
TEX10 ProteinQ9NXF1 (Uniprot-TrEMBL)
TSR1 ProteinQ2NL82 (Uniprot-TrEMBL)
TSR1ProteinQ2NL82 (Uniprot-TrEMBL)
U3 snoRNA Protein
UTP15 ProteinQ8TED0 (Uniprot-TrEMBL)
UTP18ProteinQ9Y5J1 (Uniprot-TrEMBL)
UTPA complexComplexR-HSA-6790880 (Reactome)
UTPC complexComplexR-HSA-6790889 (Reactome)
WBSCR22ProteinO43709 (Uniprot-TrEMBL)
WDR12 ProteinQ9GZL7 (Uniprot-TrEMBL)
WDR18 ProteinQ9BV38 (Uniprot-TrEMBL)
WDR43 ProteinQ15061 (Uniprot-TrEMBL)
WDR75 ProteinQ8IWA0 (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)


ArrowR-HSA-6791219 (Reactome)


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)
ERI1mim-catalysisR-HSA-6791218 (Reactome)
EXOSC10:C1D:MPHOSPH6:SKIV2L2:Exosomemim-catalysisR-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)
ISG20L2mim-catalysisR-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)
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