3jct

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<table><tr><td colspan='2'>[[3jct]] is a 58 chain structure with sequence from [http://en.wikipedia.org/wiki/Saccharomyces_cerevisiae_s288c Saccharomyces cerevisiae s288c]. Full crystallographic information is available from [http://oca.weizmann.ac.il/oca-bin/ocashort?id=3JCT OCA]. For a <b>guided tour on the structure components</b> use [http://oca.weizmann.ac.il/oca-docs/fgij/fg.htm?mol=3JCT FirstGlance]. <br>
<table><tr><td colspan='2'>[[3jct]] is a 58 chain structure with sequence from [http://en.wikipedia.org/wiki/Saccharomyces_cerevisiae_s288c Saccharomyces cerevisiae s288c]. Full crystallographic information is available from [http://oca.weizmann.ac.il/oca-bin/ocashort?id=3JCT OCA]. For a <b>guided tour on the structure components</b> use [http://oca.weizmann.ac.il/oca-docs/fgij/fg.htm?mol=3JCT FirstGlance]. <br>
</td></tr><tr id='ligand'><td class="sblockLbl"><b>[[Ligand|Ligands:]]</b></td><td class="sblockDat"><scene name='pdbligand=GTP:GUANOSINE-5-TRIPHOSPHATE'>GTP</scene>, <scene name='pdbligand=MG:MAGNESIUM+ION'>MG</scene>, <scene name='pdbligand=ZN:ZINC+ION'>ZN</scene></td></tr>
</td></tr><tr id='ligand'><td class="sblockLbl"><b>[[Ligand|Ligands:]]</b></td><td class="sblockDat"><scene name='pdbligand=GTP:GUANOSINE-5-TRIPHOSPHATE'>GTP</scene>, <scene name='pdbligand=MG:MAGNESIUM+ION'>MG</scene>, <scene name='pdbligand=ZN:ZINC+ION'>ZN</scene></td></tr>
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<tr id='resources'><td class="sblockLbl"><b>Resources:</b></td><td class="sblockDat"><span class='plainlinks'>[http://oca.weizmann.ac.il/oca-docs/fgij/fg.htm?mol=3jct FirstGlance], [http://oca.weizmann.ac.il/oca-bin/ocaids?id=3jct OCA], [http://pdbe.org/3jct PDBe], [http://www.rcsb.org/pdb/explore.do?structureId=3jct RCSB], [http://www.ebi.ac.uk/pdbsum/3jct PDBsum]</span></td></tr>
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<tr id='resources'><td class="sblockLbl"><b>Resources:</b></td><td class="sblockDat"><span class='plainlinks'>[http://oca.weizmann.ac.il/oca-docs/fgij/fg.htm?mol=3jct FirstGlance], [http://oca.weizmann.ac.il/oca-bin/ocaids?id=3jct OCA], [http://pdbe.org/3jct PDBe], [http://www.rcsb.org/pdb/explore.do?structureId=3jct RCSB], [http://www.ebi.ac.uk/pdbsum/3jct PDBsum], [http://prosat.h-its.org/prosat/prosatexe?pdbcode=3jct ProSAT]</span></td></tr>
</table>
</table>
{{Large structure}}
{{Large structure}}
== Function ==
== Function ==
[[http://www.uniprot.org/uniprot/NSA2_YEAST NSA2_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit. May play a part in the quality control of pre-60S particles. Under normal, rapid growth conditions, high levels of NSA2 would allow the progression of pre-60S particles through the ITS2 processing.<ref>PMID:16544271</ref> <ref>PMID:16861225</ref> [[http://www.uniprot.org/uniprot/NOG1_YEAST NOG1_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit.<ref>PMID:12808088</ref> [[http://www.uniprot.org/uniprot/CGR1_YEAST CGR1_YEAST]] Involved in nucleolar integrity and required for processing of the pre-rRNA for the 60S ribosome subunit.<ref>PMID:11116400</ref> <ref>PMID:11932453</ref> <ref>PMID:16544271</ref> [[http://www.uniprot.org/uniprot/PESC_YEAST PESC_YEAST]] Component of the NOP7 complex, which is required for maturation of the 25S and 5.8S ribosomal RNAs and formation of the 60S ribosome.[HAMAP-Rule:MF_03028]<ref>PMID:11911362</ref> <ref>PMID:12022229</ref> <ref>PMID:12110181</ref> <ref>PMID:18448671</ref> [[http://www.uniprot.org/uniprot/RRS1_YEAST RRS1_YEAST]] Required for ribosome biogenesis.<ref>PMID:10688653</ref> [[http://www.uniprot.org/uniprot/RL25_YEAST RL25_YEAST]] This protein binds to a specific region on the 26S rRNA. [[http://www.uniprot.org/uniprot/RL11A_YEAST RL11A_YEAST]] Binds to 5S ribosomal RNA. [[http://www.uniprot.org/uniprot/CIC1_YEAST CIC1_YEAST]] An adapter protein that specifically links the 26S proteasome to its substrate CDC4 which is one of the substrate recognition subunits of the SCF E3 ubiquitin ligase complex. Required for turnover of cell cycle regulatory proteins CDC4 and GRR1. Required for synthesis and nuclear export of 60S ribosomal subunits. Required for vegetative growth.<ref>PMID:11500370</ref> <ref>PMID:14623999</ref> [[http://www.uniprot.org/uniprot/RPF2_YEAST RPF2_YEAST]] Required for biogenesis of the 60S ribosomal subunit.<ref>PMID:12702244</ref> [[http://www.uniprot.org/uniprot/RLP7_YEAST RLP7_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit. May act as a specificity factor that binds precursor rRNAs and tethers the enzymes that carry out the early 5' to 3' exonucleolytic reactions that generate the mature rRNAs.<ref>PMID:11087857</ref> [[http://www.uniprot.org/uniprot/MRT4_YEAST MRT4_YEAST]] Involved in mRNA turnover and ribosome assembly. [[http://www.uniprot.org/uniprot/NLE1_YEAST NLE1_YEAST]] Involved in processing and efficient intra-nuclear transport or pre-60S ribosomal subunits. Forms a complex with REA1 which is essential for ATP-dependent dissociation of a group of nonribosomal factors from the pre-60S particle.<ref>PMID:16221974</ref> <ref>PMID:19737519</ref> <ref>PMID:20542003</ref> [[http://www.uniprot.org/uniprot/NOG2_YEAST NOG2_YEAST]] GTPase that associates with pre-60S ribosomal subunits in the nucleolus and is required for their nuclear export and maturation.<ref>PMID:11707418</ref> [[http://www.uniprot.org/uniprot/RL4A_YEAST RL4A_YEAST]] Participates in the regulation of the accumulation of its own mRNA.<ref>PMID:2065661</ref> [[http://www.uniprot.org/uniprot/RL37A_YEAST RL37A_YEAST]] Binds to the 23S rRNA (By similarity). [[http://www.uniprot.org/uniprot/BUD20_YEAST BUD20_YEAST]] Involved in positioning the proximal bud pole signal.<ref>PMID:11452010</ref> [[http://www.uniprot.org/uniprot/IF6_YEAST IF6_YEAST]] Binds to the 60S ribosomal subunit and prevents its association with the 40S ribosomal subunit to form the 80S initiation complex in the cytoplasm. Is also involved in ribosome biogenesis. Associates with pre-60S subunits in the nucleus and is involved in its nuclear export. Cytoplasmic release of TIF6 from 60S subunits and nuclear relocalization is promoted by the GTPase RIA1/EFL1 and by SDO1. Also required for pre-rRNA processing.<ref>PMID:10085284</ref> <ref>PMID:11779510</ref> <ref>PMID:11238882</ref> <ref>PMID:17353896</ref> <ref>PMID:18256024</ref> <ref>PMID:11101899</ref> [[http://www.uniprot.org/uniprot/NOP15_YEAST NOP15_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit. Required for pre-rRNA processing and cytokinesis. Associates with the precursors of the 25S and 5.8S rRNAs.<ref>PMID:11583614</ref> <ref>PMID:14657029</ref> [[http://www.uniprot.org/uniprot/RLP24_YEAST RLP24_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit. Ensures the docking of NOG1 to pre-60S particles.<ref>PMID:12808088</ref> [[http://www.uniprot.org/uniprot/RL5_YEAST RL5_YEAST]] Binds 5S RNA and is required for 60S subunit assembly. [[http://www.uniprot.org/uniprot/ARX1_YEAST ARX1_YEAST]] Probable metalloprotease involved in proper assembly of pre-ribosomal particles during the biogenesis of the 60S ribosomal subunit. Accompanies the pre-60S particles to the cytoplasm.<ref>PMID:12374754</ref> <ref>PMID:16648468</ref> [[http://www.uniprot.org/uniprot/NUG1_YEAST NUG1_YEAST]] GTPase required for 60S ribosomal subunit export to the cytoplasm.<ref>PMID:11583615</ref> [[http://www.uniprot.org/uniprot/NOP53_YEAST NOP53_YEAST]] Late-acting factor in the nuclear maturation of 60S ribosomal subunits, which is required for normal acquisition of export competence. Required for the export of the large subunit. Acts to stimulate the RNase activity of the exosome complex, and may recruit the exosome to 7S pre-rRNA for processing. Associates with numerous RNAs including the 27S and 7S pre-rRNAs and the box H/ACA snoRNA snR37. Also interacts (via N-terminal region) with the mature 25S rRNA and the mature 5.8S rRNA.<ref>PMID:15686447</ref> <ref>PMID:16043506</ref> <ref>PMID:16128814</ref> <ref>PMID:18631361</ref>
[[http://www.uniprot.org/uniprot/NSA2_YEAST NSA2_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit. May play a part in the quality control of pre-60S particles. Under normal, rapid growth conditions, high levels of NSA2 would allow the progression of pre-60S particles through the ITS2 processing.<ref>PMID:16544271</ref> <ref>PMID:16861225</ref> [[http://www.uniprot.org/uniprot/NOG1_YEAST NOG1_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit.<ref>PMID:12808088</ref> [[http://www.uniprot.org/uniprot/CGR1_YEAST CGR1_YEAST]] Involved in nucleolar integrity and required for processing of the pre-rRNA for the 60S ribosome subunit.<ref>PMID:11116400</ref> <ref>PMID:11932453</ref> <ref>PMID:16544271</ref> [[http://www.uniprot.org/uniprot/PESC_YEAST PESC_YEAST]] Component of the NOP7 complex, which is required for maturation of the 25S and 5.8S ribosomal RNAs and formation of the 60S ribosome.[HAMAP-Rule:MF_03028]<ref>PMID:11911362</ref> <ref>PMID:12022229</ref> <ref>PMID:12110181</ref> <ref>PMID:18448671</ref> [[http://www.uniprot.org/uniprot/RRS1_YEAST RRS1_YEAST]] Required for ribosome biogenesis.<ref>PMID:10688653</ref> [[http://www.uniprot.org/uniprot/RL25_YEAST RL25_YEAST]] This protein binds to a specific region on the 26S rRNA. [[http://www.uniprot.org/uniprot/RL11A_YEAST RL11A_YEAST]] Binds to 5S ribosomal RNA. [[http://www.uniprot.org/uniprot/CIC1_YEAST CIC1_YEAST]] An adapter protein that specifically links the 26S proteasome to its substrate CDC4 which is one of the substrate recognition subunits of the SCF E3 ubiquitin ligase complex. Required for turnover of cell cycle regulatory proteins CDC4 and GRR1. Required for synthesis and nuclear export of 60S ribosomal subunits. Required for vegetative growth.<ref>PMID:11500370</ref> <ref>PMID:14623999</ref> [[http://www.uniprot.org/uniprot/RPF2_YEAST RPF2_YEAST]] Required for biogenesis of the 60S ribosomal subunit.<ref>PMID:12702244</ref> [[http://www.uniprot.org/uniprot/RLP7_YEAST RLP7_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit. May act as a specificity factor that binds precursor rRNAs and tethers the enzymes that carry out the early 5' to 3' exonucleolytic reactions that generate the mature rRNAs.<ref>PMID:11087857</ref> [[http://www.uniprot.org/uniprot/MRT4_YEAST MRT4_YEAST]] Involved in mRNA turnover and ribosome assembly. [[http://www.uniprot.org/uniprot/NLE1_YEAST NLE1_YEAST]] Involved in processing and efficient intra-nuclear transport or pre-60S ribosomal subunits. Forms a complex with REA1 which is essential for ATP-dependent dissociation of a group of nonribosomal factors from the pre-60S particle.<ref>PMID:16221974</ref> <ref>PMID:19737519</ref> <ref>PMID:20542003</ref> [[http://www.uniprot.org/uniprot/NOG2_YEAST NOG2_YEAST]] GTPase that associates with pre-60S ribosomal subunits in the nucleolus and is required for their nuclear export and maturation.<ref>PMID:11707418</ref> [[http://www.uniprot.org/uniprot/RL4A_YEAST RL4A_YEAST]] Participates in the regulation of the accumulation of its own mRNA.<ref>PMID:2065661</ref> [[http://www.uniprot.org/uniprot/RL37A_YEAST RL37A_YEAST]] Binds to the 23S rRNA (By similarity). [[http://www.uniprot.org/uniprot/BUD20_YEAST BUD20_YEAST]] Involved in positioning the proximal bud pole signal.<ref>PMID:11452010</ref> [[http://www.uniprot.org/uniprot/IF6_YEAST IF6_YEAST]] Binds to the 60S ribosomal subunit and prevents its association with the 40S ribosomal subunit to form the 80S initiation complex in the cytoplasm. Is also involved in ribosome biogenesis. Associates with pre-60S subunits in the nucleus and is involved in its nuclear export. Cytoplasmic release of TIF6 from 60S subunits and nuclear relocalization is promoted by the GTPase RIA1/EFL1 and by SDO1. Also required for pre-rRNA processing.<ref>PMID:10085284</ref> <ref>PMID:11779510</ref> <ref>PMID:11238882</ref> <ref>PMID:17353896</ref> <ref>PMID:18256024</ref> <ref>PMID:11101899</ref> [[http://www.uniprot.org/uniprot/NOP15_YEAST NOP15_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit. Required for pre-rRNA processing and cytokinesis. Associates with the precursors of the 25S and 5.8S rRNAs.<ref>PMID:11583614</ref> <ref>PMID:14657029</ref> [[http://www.uniprot.org/uniprot/RLP24_YEAST RLP24_YEAST]] Involved in the biogenesis of the 60S ribosomal subunit. Ensures the docking of NOG1 to pre-60S particles.<ref>PMID:12808088</ref> [[http://www.uniprot.org/uniprot/RL5_YEAST RL5_YEAST]] Binds 5S RNA and is required for 60S subunit assembly. [[http://www.uniprot.org/uniprot/ARX1_YEAST ARX1_YEAST]] Probable metalloprotease involved in proper assembly of pre-ribosomal particles during the biogenesis of the 60S ribosomal subunit. Accompanies the pre-60S particles to the cytoplasm.<ref>PMID:12374754</ref> <ref>PMID:16648468</ref> [[http://www.uniprot.org/uniprot/NUG1_YEAST NUG1_YEAST]] GTPase required for 60S ribosomal subunit export to the cytoplasm.<ref>PMID:11583615</ref> [[http://www.uniprot.org/uniprot/NOP53_YEAST NOP53_YEAST]] Late-acting factor in the nuclear maturation of 60S ribosomal subunits, which is required for normal acquisition of export competence. Required for the export of the large subunit. Acts to stimulate the RNase activity of the exosome complex, and may recruit the exosome to 7S pre-rRNA for processing. Associates with numerous RNAs including the 27S and 7S pre-rRNAs and the box H/ACA snoRNA snR37. Also interacts (via N-terminal region) with the mature 25S rRNA and the mature 5.8S rRNA.<ref>PMID:15686447</ref> <ref>PMID:16043506</ref> <ref>PMID:16128814</ref> <ref>PMID:18631361</ref>
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<div style="background-color:#fffaf0;">
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== Publication Abstract from PubMed ==
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Ribosome biogenesis is a highly complex process in eukaryotes, involving temporally and spatially regulated ribosomal protein (r-protein) binding and ribosomal RNA remodelling events in the nucleolus, nucleoplasm and cytoplasm. Hundreds of assembly factors, organized into sequential functional groups, facilitate and guide the maturation process into productive assembly branches in and across different cellular compartments. However, the precise mechanisms by which these assembly factors function are largely unknown. Here we use cryo-electron microscopy to characterize the structures of yeast nucleoplasmic pre-60S particles affinity-purified using the epitope-tagged assembly factor Nog2. Our data pinpoint the locations and determine the structures of over 20 assembly factors, which are enriched in two areas: an arc region extending from the central protuberance to the polypeptide tunnel exit, and the domain including the internal transcribed spacer 2 (ITS2) that separates 5.8S and 25S ribosomal RNAs. In particular, two regulatory GTPases, Nog2 and Nog1, act as hub proteins to interact with multiple, distant assembly factors and functional ribosomal RNA elements, manifesting their critical roles in structural remodelling checkpoints and nuclear export. Moreover, our snapshots of compositionally and structurally different pre-60S intermediates provide essential mechanistic details for three major remodelling events before nuclear export: rotation of the 5S ribonucleoprotein, construction of the active centre and ITS2 removal. The rich structural information in our structures provides a framework to dissect molecular roles of diverse assembly factors in eukaryotic ribosome assembly.
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Diverse roles of assembly factors revealed by structures of late nuclear pre-60S ribosomes.,Wu S, Tutuncuoglu B, Yan K, Brown H, Zhang Y, Tan D, Gamalinda M, Yuan Y, Li Z, Jakovljevic J, Ma C, Lei J, Dong MQ, Woolford JL Jr, Gao N Nature. 2016 Jun 2;534(7605):133-7. doi: 10.1038/nature17942. PMID:27251291<ref>PMID:27251291</ref>
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From MEDLINE&reg;/PubMed&reg;, a database of the U.S. National Library of Medicine.<br>
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</div>
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<div class="pdbe-citations 3jct" style="background-color:#fffaf0;"></div>
== References ==
== References ==
<references/>
<references/>

Revision as of 10:44, 1 February 2017

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Cryo-em structure of eukaryotic pre-60S ribosomal subunits

3jct, resolution 3.08Å

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