£€0€ €0€ €0€ €¡¡€0€ €4A3V£€0€¢€0€ €Protein Data Bank¡€ €¡€0€ €Ü¡€¢€¢€¡€0€ €PDB¡€¡€4A3V£€¡€0€ €Û¡€ ¢€¤€¡€0€ €0€ € €0€0€ €¡€0€ €Richet£€N.Richet¤€N.0€ €¡€0€ €Barrault£€ M.B.Barrault¤€M.B.0€ €¡€0€ €Godart£€C.Godart¤€C.0€ €¡€0€ €Murciano£€ B.Murciano¤€B.0€ €¡€0€ € Le Tallec£€ B.Le Tallec¤€B.0€ €¡€0€ €Rous£€E.Rous¤€E.0€ €¡€0€ €Ledu£€M.H.Ledu¤€M.H.0€ €¡€0€ € Charbonnier£€J.B.Charbonnier¤€J.B.0€ €¡€0€ €Legrand£€ P.Legrand¤€P.0€ €¡€0€ €Guerois£€ R.Guerois¤€R.0€ €¡€0€ €Peyroche£€ A.Peyroche¤€A.0€ €¡€0€ € Ochsenbein£€ F.Ochsenbein¤€F.¡€0€ €¡€0€ €Û¡€ ¢€¤€«€1€¤€0€ €1€ €~Dual functions of the Hsm3 protein in chaperoning and scaffolding regulatory particle subunits during the proteasome assembly.¡€0€ € €0€0€ €¡€0€ €Barrault£€ M.B.Barrault¤€M.B.0€ €¡€0€ €Richet£€N.Richet¤€N.0€ €¡€0€ €Godard£€C.Godard¤€C.0€ €¡€0€ €Murciano£€ B.Murciano¤€B.0€ €¡€0€ € Le Tallec£€ B.Le Tallec¤€B.0€ €¡€0€ €Rousseau£€ E.Rousseau¤€E.0€ €¡€0€ €Legrand£€ P.Legrand¤€P.0€ €¡€0€ € Charbonnier£€J.B.Charbonnier¤€J.B.0€ €¡€0€ €Le Du£€ M.H.Le Du¤€M.H.0€ €¡€0€ €Guerois£€ R.Guerois¤€R.0€ €¡€0€ € Ochsenbein£€ F.Ochsenbein¤€F.0€ €¡€0€ €Peyroche£€ A.Peyroche¤€A.¡€ €‚Commissariat a l'Energie Atomique (CEA), Institut de Biologie et Technologies de Saclay (iBiTEC-S), Service de Biologie Integrative et Genetique Moleculaire, Laboratoire du Metabolisme des Acides Nucleiques et Reponses aux Genotoxiques, F-91191 Gif-sur-Yvette, France.¢€ €0€ €1€¤€Proc. Natl. Acad. Sci. U.S.A.¥€Proc Natl Acad Sci U S A§€ 1091-6490 €OProceedings of the National Academy of Sciences of the United States of America¡€0€ €¡€0€ €Ü¡€¢€¡€109¢€17£€ E1001-E1010¨€eng¬€­€1€0€ € ¡€¡€0€ €Ü¡€¢€0€ €ÿ¡€¡€0€ €Ü¡€¢€¤€¥€0€ €¡€¡€0€ €Ü¡€¢€¤€¥€0€ € ¡€¡€0€ €Ü¡€¢€¤€¥€£€1€£€ 1116538109¢€10.1073/pnas.1116538109 €V¹€§€0€ €PDB Citation Status¡€¡€primary¢€0€¡€¡€0€ €Ü¡€¢€¢€0€¢€ €0€ €1€ €}Dual functions of the Hsm3 protein in chaperoning and scaffolding regulatory particle subunits during the proteasome assembly¡€0€ €¡€0€ €Ü¡€¢€£€‚&The 26S proteasome, a molecular machine responsible for regulated protein degradation, consists of a proteolytic core particle (20S CP) associated with 19S regulatory particles (19S RPs) subdivided into base and lid subcomplexes. The assembly of 19S RP base subcomplex is mediated by multiple dedicated chaperones. Among these, Hsm3 is important for normal growth and directly targets the carboxyl-terminal (C-terminal) domain of Rpt1 of the Rpt1-Rpt2-Rpn1 assembly intermediate. Here, we report crystal structures of the yeast Hsm3 chaperone free and bound to the C-terminal domain of Rpt1. Unexpectedly, the structure of the complex suggests that within the Hsm3-Rpt1-Rpt2 module, Hsm3 also contacts Rpt2. We show that in both yeast and mammals, Hsm3 actually directly binds the AAA domain of Rpt2. The Hsm3 C-terminal region involved in this interaction is required in vivo for base assembly, although it is dispensable for binding Rpt1. Although Rpt1 and Rpt2 exhibit weak affinity for each other, Hsm3 unexpectedly acts as an essential matchmaker for the Rpt1-Rpt2-Rpn1 assembly by bridging both Rpt1 and Rpt2. In addition, we provide structural and biochemical evidence on how Hsm3/S5b may regulate the 19S RP association to the 20S CP proteasome. Our data point out the diverse functions of assembly chaperones.¬€V¹€¢€0€ €0€ €eYeast Regulatory Particle Proteasome Assembly Chaperone Hsm3 In Complex With Rpt1 C-Terminal Fragment¡€ChaperoneATP BINDING PROTEIN¢€‚‘Mol_id: 1; Organism_scientific: Saccharomyces Cerevisiae; Organism_common: Baker's Yeast; Organism_taxid: 580240; Strain: W303; Expression_system: Escherichia Coli; Expression_system_taxid: 511693; Expression_system_strain: Bl21; Expression_system_vector_type: Plasmid; Expression_system_vector: Petduet; Mol_id: 2; Organism_scientific: Saccharomyces Cerevisiae; Organism_common: Baker's Yeast; Organism_taxid: 580240; Strain: W303; Expression_system: Escherichia Coli; Expression_system_taxid: 511693; Expression_system_strain: Bl21; Expression_system_vector_type: Plasmid; Expression_system_vector: Petduet; Mol_id: 3; Organism_scientific: Saccharomyces Cerevisiae; Organism_common: Baker's Yeast; Organism_taxid: 580240; Strain: W303; Expression_system: Escherichia Coli; Expression_system_taxid: 511693; Expression_system_strain: Bl21; Expression_system_vector_type: Plasmid; Expression_system_vector: Petduet£€‚bYeast Regulatory Particle Proteasome Assembly Chaperone Hsm3 Complex With Rpt1 C-Terminal Fragment Chaperone-Atp Binding Protein Complex, 19s Mol_id: 1; Molecule: Dna Mismatch Repair Protein Hsm3; Chain: A, C; Synonym: Hsm3, Enhanced Spontaneous Mutability Protein 3; Engineered: Yes; Mol_id: 2; Molecule: 26s Protease Regulatory Subunit 7 Homolog; Chain: B, D; Synonym: Rpt1-Cter, Protein Cim5, Tat-Binding Homolog 3; Engineered: Yes; Mol_id: 3; Molecule: Linker; Chain: E; Engineered: Yes; Other_details: Chain E Is Believed To Be Made Of Residues F C-Terminus Of Either Chain A Or C, But The Identity Is Unc§€ÿ¡€0€0€ €¡€0€ €A£€"1 Dna Mismatch Repair Protein Hsm3¨€¥€0€¢€0€ €Saccharomyces cerevisiae W303£€1€0€ €taxon¡€ €Ú¥€0€ € €0€ € Saccharomyces¡€ cerevisiae£€~Eukaryota; Fungi; Dikarya; Ascomycota; Saccharomycotina; Saccharomycetes; Saccharomycetales; 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dictionary¡€ €¡€:0€ €x¡€ 115 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €y¡€ 116 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €z¡€ 117 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€ 0€ €{¡€ 118 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €|¡€ 119 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€:0€ €}¡€ 120 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €~¡€ 121 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €¡€ 122 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €€¡€ 123 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€.0€ €¡€ 124 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €‚¡€ 125 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€+0€ €ƒ¡€ 126 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€"0€ €„¡€ 127 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €…¡€ 128 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €†¡€ 129 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€(0€ €‡¡€ 130 ¢€¢€0€ €¡€0€ €Standard residue dictionary¡€ €¡€0€ €ˆ¡€ 131 ¢€¢€0€ €¡€0€ €Standard 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