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Eukaryotic ribosome biogenesis II


Ed Hurt,1 Janet Iwasa,2 and Roland Beckmann3
1
Heidelberg University Biochemistry Center (BZH), Heidelberg, Germany; 2Department of Biochemistry, University of Utah, Salt Lake City, UT, USA ; 3Gene Center, Ludwig-
Maximilians-Universität München, Munich, Germany

Cleavage sites A0 A1 D A2 A3 B1 C2 C1 B2

Pol I 35S pre-rRNA 5v 18S 5.8S 25S 3v Pol III 5S


ITS1 ITS2

5S rRNA transcription
NUCLEOLAR PRE-60S BIOGENESIS

Pol I Earliest 5S RNP Pol III


Nucleolar +ATP
pre-60S Foot factors factors
25S 5S Rea1
factors factors
ITS2 RNA
Has1 5v uL18
5S RNP uL5 Ytm1-Erb1
5.8S
ITS1 Brx1- assembly
5v 18S Noc1- Spb4
Noc2 Ebp2 Noc2-Noc3
Nucleolar
Pre-rRNA H84
A2 A2 factors 5S
transcription Spb1
Rrp5 cleavage Dbp10 RNA
snoRNPs
Earliest 90S Rlp24 Nog1
factors Further
90S Tif6
factors 90S
90S-pre-60S Pre-60S (state A) Pre-60S (Dbp10-3) Pre-60S (state E)
particle Pre-40S biogenesis

Pre-60S 7S>5.8S
Rixosome
(nuclear Nmd3) processing
+ATP 5S RNP ITS2 Exosome
Rea1 rotated Mtr4
and Rix1 5S RNA
complex rotation AIM
NES Nop53 Nop53
Nmd3
90˚
Nog2
Arx1
Rsa4 Bud20
Exosome
+ATP

Nuclear export
Export Pre-60S Pre-60S Pre-60S Pre-60S
factors
(Rea1-Rix1) (Nop53 exosome Mtr4) (Nog2/Arx1) (NE2)
NPC NUCLEOPLASM NUCLEOPLASMIC PRE-60S BIOGENESIS

CYTOPLASM C Y T O P L A S M I C P R E - 6 0 S M AT U R AT I O N
Central
protuberance
Crm1- Mrt4
L1 P-stalk
RanGTP 5S
Nog1 uL16 stalk
Sdo1 Efl1
Yvh1 5.8S PTC
Lsg1
Rlp24 Tif6
Mex67 eL24 25S rRNA
-Mtr2 18S
PET rRNA
Drg1
60S 40S
Pre-60S in NPC Pre-60S (Drg1) Pre-60S (Lsg1) Pre-60S (Efl1) subunit subunit

Pre-60S assembly factors


Earliest pre-60S factors snoRNPs Nucleolar factors Foot factors Nuclear factors Cytoplasmic
Noc1 Dbp6 H/ACA Cbf5 Nhp2 Rrp1 Ssf1 Tif6 Rix7 Cic1 Rea1 Nog2 factors
Noc2 Upa1 snR35 Gar1 Nop10 Nsa1 Ssf2 Nop2 Dbp3 Nop15 Rix1 Cgr1 Drg1
Rrp5 Upa2 C/D box Nop1 Nop56 Rpf1 Rrp15 Nip7 Dbp7 Rlp7 Ipi3 Rsa4 Yvh1
Urb1 Mak5 snR190 Snu13 Nop58 Mak16 Rrp14 Nog1 Dpb9 Nop7 Ipi1 Nop53 Rei1
Urb2 Rbp95 Brx1 Nop16 Mak11 Dbp10 Export factors Sda1 Arx1 Reh1
Noc2 Nop4 5S RNP Ebp2 Spb4 Has1 Drs1 Crm1 Las1 Alb1 Lsg1
Noc3 Pwp1 5S RNA Syo1 Ytm1 Puf6 Nsa2 Mrt4 RanGTP Grc3 Bud20 Sdo1
Rsa3 Nop12 uL18 Rpf2 Erb1 Loc1 Spb1 Mex67 Rat1 Ecm1 Efl1
Nop8 Nop13 uL5 Rrs1 Mak15 Rlp24 Mtr2 Rai1 Nmd3

Scenes from the video Eukaryotic pre-60S ribosome biogenesis

90S-pre-60S assembly Nucleolar Nuclear Nuclear Pre-60S Pre-60S Cytoplasmic Mature 60S
pre-60S pre-60S with pre-60S with in NPC following pre-60S subunit
Mtr4-exosome Rea1-Rixosome nuclear export

See online version for supplemental information,


1314 Cell 187, February 29, 2024 © 2024 Elsevier Inc. DOI https://doi.org/10.1016/j.cell.2024.01.043 legends, references, and declaration of interests

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