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9U4M

Focused refinement of 19S in the substrate-engaged human 26S proteasome bound to midnolin with RPT6 at top of spiral staircase

Functional Information from GO Data
ChainGOidnamespacecontents
a0000502cellular_componentproteasome complex
a0005198molecular_functionstructural molecule activity
a0005515molecular_functionprotein binding
a0005576cellular_componentextracellular region
a0005634cellular_componentnucleus
a0005654cellular_componentnucleoplasm
a0005829cellular_componentcytosol
a0005838cellular_componentproteasome regulatory particle
a0006511biological_processubiquitin-dependent protein catabolic process
a0006979biological_processresponse to oxidative stress
a0008021cellular_componentsynaptic vesicle
a0008541cellular_componentproteasome regulatory particle, lid subcomplex
a0010498biological_processproteasomal protein catabolic process
a0016020cellular_componentmembrane
a0022624cellular_componentproteasome accessory complex
a0034774cellular_componentsecretory granule lumen
a0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
a0061136biological_processregulation of proteasomal protein catabolic process
a0071357biological_processcellular response to type I interferon
a1904813cellular_componentficolin-1-rich granule lumen
A0000502cellular_componentproteasome complex
A0000932cellular_componentP-body
A0001649biological_processosteoblast differentiation
A0005515molecular_functionprotein binding
A0005576cellular_componentextracellular region
A0005634cellular_componentnucleus
A0005654cellular_componentnucleoplasm
A0005737cellular_componentcytoplasm
A0005829cellular_componentcytosol
A0006511biological_processubiquitin-dependent protein catabolic process
A0006979biological_processresponse to oxidative stress
A0008021cellular_componentsynaptic vesicle
A0010498biological_processproteasomal protein catabolic process
A0016020cellular_componentmembrane
A0022624cellular_componentproteasome accessory complex
A0034774cellular_componentsecretory granule lumen
A0036402molecular_functionproteasome-activating activity
A0036464cellular_componentcytoplasmic ribonucleoprotein granule
A0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
A0061136biological_processregulation of proteasomal protein catabolic process
A0071357biological_processcellular response to type I interferon
A1901800biological_processpositive regulation of proteasomal protein catabolic process
A1904813cellular_componentficolin-1-rich granule lumen
b0000502cellular_componentproteasome complex
b0003723molecular_functionRNA binding
b0005515molecular_functionprotein binding
b0005634cellular_componentnucleus
b0005829cellular_componentcytosol
b0006979biological_processresponse to oxidative stress
b0008021cellular_componentsynaptic vesicle
b0010498biological_processproteasomal protein catabolic process
b0022624cellular_componentproteasome accessory complex
b0042802molecular_functionidentical protein binding
b0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
b0060090molecular_functionmolecular adaptor activity
b0061136biological_processregulation of proteasomal protein catabolic process
b0071357biological_processcellular response to type I interferon
B0000502cellular_componentproteasome complex
B0003723molecular_functionRNA binding
B0005515molecular_functionprotein binding
B0005634cellular_componentnucleus
B0005654cellular_componentnucleoplasm
B0005737cellular_componentcytoplasm
B0005829cellular_componentcytosol
B0005838cellular_componentproteasome regulatory particle
B0006979biological_processresponse to oxidative stress
B0008021cellular_componentsynaptic vesicle
B0010498biological_processproteasomal protein catabolic process
B0016020cellular_componentmembrane
B0022624cellular_componentproteasome accessory complex
B0036402molecular_functionproteasome-activating activity
B0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
B0061136biological_processregulation of proteasomal protein catabolic process
B0071357biological_processcellular response to type I interferon
c0000502cellular_componentproteasome complex
c0000724biological_processdouble-strand break repair via homologous recombination
c0004843molecular_functioncysteine-type deubiquitinase activity
c0005515molecular_functionprotein binding
c0005576cellular_componentextracellular region
c0005634cellular_componentnucleus
c0005654cellular_componentnucleoplasm
c0005829cellular_componentcytosol
c0006303biological_processdouble-strand break repair via nonhomologous end joining
c0006511biological_processubiquitin-dependent protein catabolic process
c0006890biological_processretrograde vesicle-mediated transport, Golgi to endoplasmic reticulum
c0006979biological_processresponse to oxidative stress
c0008021cellular_componentsynaptic vesicle
c0008237molecular_functionmetallopeptidase activity
c0008541cellular_componentproteasome regulatory particle, lid subcomplex
c0010498biological_processproteasomal protein catabolic process
c0016241biological_processregulation of macroautophagy
c0016579biological_processprotein deubiquitination
c0022624cellular_componentproteasome accessory complex
c0031597cellular_componentcytosolic proteasome complex
c0034774cellular_componentsecretory granule lumen
c0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
c0045471biological_processresponse to ethanol
c0061133molecular_functionendopeptidase activator activity
c0061136biological_processregulation of proteasomal protein catabolic process
c0061578molecular_functionK63-linked deubiquitinase activity
c0070536biological_processprotein K63-linked deubiquitination
c0070628molecular_functionproteasome binding
c0071357biological_processcellular response to type I interferon
c0140492molecular_functionmetal-dependent deubiquitinase activity
c1904813cellular_componentficolin-1-rich granule lumen
C0000502cellular_componentproteasome complex
C0005102molecular_functionsignaling receptor binding
C0005515molecular_functionprotein binding
C0005634cellular_componentnucleus
C0005654cellular_componentnucleoplasm
C0005737cellular_componentcytoplasm
C0005829cellular_componentcytosol
C0005838cellular_componentproteasome regulatory particle
C0005886cellular_componentplasma membrane
C0006357biological_processregulation of transcription by RNA polymerase II
C0006979biological_processresponse to oxidative stress
C0008021cellular_componentsynaptic vesicle
C0008134molecular_functiontranscription factor binding
C0010498biological_processproteasomal protein catabolic process
C0016020cellular_componentmembrane
C0016234cellular_componentinclusion body
C0016887molecular_functionATP hydrolysis activity
C0017025molecular_functionTBP-class protein binding
C0022624cellular_componentproteasome accessory complex
C0031410cellular_componentcytoplasmic vesicle
C0031531molecular_functionthyrotropin-releasing hormone receptor binding
C0031595cellular_componentnuclear proteasome complex
C0031597cellular_componentcytosolic proteasome complex
C0036402molecular_functionproteasome-activating activity
C0043069biological_processnegative regulation of programmed cell death
C0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
C0045893biological_processpositive regulation of DNA-templated transcription
C0061136biological_processregulation of proteasomal protein catabolic process
C0070062cellular_componentextracellular exosome
C0071357biological_processcellular response to type I interferon
C0072562cellular_componentblood microparticle
C0090261biological_processpositive regulation of inclusion body assembly
C0140296molecular_functiongeneral transcription initiation factor binding
C0140297molecular_functionDNA-binding transcription factor binding
C1901800biological_processpositive regulation of proteasomal protein catabolic process
d0000502cellular_componentproteasome complex
d0005515molecular_functionprotein binding
d0005634cellular_componentnucleus
d0005654cellular_componentnucleoplasm
d0005829cellular_componentcytosol
d0005838cellular_componentproteasome regulatory particle
d0006979biological_processresponse to oxidative stress
d0008021cellular_componentsynaptic vesicle
d0008541cellular_componentproteasome regulatory particle, lid subcomplex
d0010498biological_processproteasomal protein catabolic process
d0022624cellular_componentproteasome accessory complex
d0032991cellular_componentprotein-containing complex
d0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
d0061136biological_processregulation of proteasomal protein catabolic process
d0071357biological_processcellular response to type I interferon
D0000502cellular_componentproteasome complex
D0005515molecular_functionprotein binding
D0005634cellular_componentnucleus
D0005654cellular_componentnucleoplasm
D0005737cellular_componentcytoplasm
D0005829cellular_componentcytosol
D0006508biological_processproteolysis
D0006979biological_processresponse to oxidative stress
D0008021cellular_componentsynaptic vesicle
D0010498biological_processproteasomal protein catabolic process
D0016020cellular_componentmembrane
D0022624cellular_componentproteasome accessory complex
D0036402molecular_functionproteasome-activating activity
D0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
D0061136biological_processregulation of proteasomal protein catabolic process
D0071357biological_processcellular response to type I interferon
e0000502cellular_componentproteasome complex
e0005515molecular_functionprotein binding
e0005634cellular_componentnucleus
e0005654cellular_componentnucleoplasm
e0005829cellular_componentcytosol
e0006406biological_processmRNA export from nucleus
e0006979biological_processresponse to oxidative stress
e0008021cellular_componentsynaptic vesicle
e0010498biological_processproteasomal protein catabolic process
e0032039cellular_componentintegrator complex
e0032991cellular_componentprotein-containing complex
e0043130molecular_functionubiquitin binding
e0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
e0045944biological_processpositive regulation of transcription by RNA polymerase II
e0061136biological_processregulation of proteasomal protein catabolic process
e0070390cellular_componenttranscription export complex 2
e0071357biological_processcellular response to type I interferon
E0005634cellular_componentnucleus
E0005737cellular_componentcytoplasm
F0000502cellular_componentproteasome complex
F0000932cellular_componentP-body
F0005515molecular_functionprotein binding
F0005576cellular_componentextracellular region
F0005634cellular_componentnucleus
F0005654cellular_componentnucleoplasm
F0005737cellular_componentcytoplasm
F0005829cellular_componentcytosol
F0006979biological_processresponse to oxidative stress
F0008021cellular_componentsynaptic vesicle
F0010498biological_processproteasomal protein catabolic process
F0016020cellular_componentmembrane
F0022624cellular_componentproteasome accessory complex
F0034774cellular_componentsecretory granule lumen
F0036402molecular_functionproteasome-activating activity
F0042802molecular_functionidentical protein binding
F0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
F0043921biological_processhost-mediated perturbation of viral transcription
F0045944biological_processpositive regulation of transcription by RNA polymerase II
F0061136biological_processregulation of proteasomal protein catabolic process
F0071357biological_processcellular response to type I interferon
F1901800biological_processpositive regulation of proteasomal protein catabolic process
F1904813cellular_componentficolin-1-rich granule lumen
U0000502cellular_componentproteasome complex
U0005515molecular_functionprotein binding
U0005576cellular_componentextracellular region
U0005654cellular_componentnucleoplasm
U0005829cellular_componentcytosol
U0005838cellular_componentproteasome regulatory particle
U0006979biological_processresponse to oxidative stress
U0008021cellular_componentsynaptic vesicle
U0010498biological_processproteasomal protein catabolic process
U0016020cellular_componentmembrane
U0022624cellular_componentproteasome accessory complex
U0031625molecular_functionubiquitin protein ligase binding
U0034515cellular_componentproteasome storage granule
U0035578cellular_componentazurophil granule lumen
U0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
U0061136biological_processregulation of proteasomal protein catabolic process
U0071357biological_processcellular response to type I interferon
V0000502cellular_componentproteasome complex
V0005515molecular_functionprotein binding
V0005576cellular_componentextracellular region
V0005634cellular_componentnucleus
V0005654cellular_componentnucleoplasm
V0005829cellular_componentcytosol
V0005838cellular_componentproteasome regulatory particle
V0006511biological_processubiquitin-dependent protein catabolic process
V0006979biological_processresponse to oxidative stress
V0008021cellular_componentsynaptic vesicle
V0008541cellular_componentproteasome regulatory particle, lid subcomplex
V0010498biological_processproteasomal protein catabolic process
V0016020cellular_componentmembrane
V0022624cellular_componentproteasome accessory complex
V0034774cellular_componentsecretory granule lumen
V0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
V0061136biological_processregulation of proteasomal protein catabolic process
V0070062cellular_componentextracellular exosome
V0071357biological_processcellular response to type I interferon
V1904813cellular_componentficolin-1-rich granule lumen
W0000502cellular_componentproteasome complex
W0005515molecular_functionprotein binding
W0005576cellular_componentextracellular region
W0005654cellular_componentnucleoplasm
W0005829cellular_componentcytosol
W0005838cellular_componentproteasome regulatory particle
W0006979biological_processresponse to oxidative stress
W0008021cellular_componentsynaptic vesicle
W0008541cellular_componentproteasome regulatory particle, lid subcomplex
W0010498biological_processproteasomal protein catabolic process
W0016020cellular_componentmembrane
W0022624cellular_componentproteasome accessory complex
W0034774cellular_componentsecretory granule lumen
W0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
W0061136biological_processregulation of proteasomal protein catabolic process
W0070062cellular_componentextracellular exosome
W0071357biological_processcellular response to type I interferon
W1904813cellular_componentficolin-1-rich granule lumen
X0000502cellular_componentproteasome complex
X0005198molecular_functionstructural molecule activity
X0005515molecular_functionprotein binding
X0005576cellular_componentextracellular region
X0005634cellular_componentnucleus
X0005654cellular_componentnucleoplasm
X0005829cellular_componentcytosol
X0005838cellular_componentproteasome regulatory particle
X0006511biological_processubiquitin-dependent protein catabolic process
X0006979biological_processresponse to oxidative stress
X0008021cellular_componentsynaptic vesicle
X0008541cellular_componentproteasome regulatory particle, lid subcomplex
X0010498biological_processproteasomal protein catabolic process
X0016020cellular_componentmembrane
X0022624cellular_componentproteasome accessory complex
X0032991cellular_componentprotein-containing complex
X0034774cellular_componentsecretory granule lumen
X0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
X0043248biological_processproteasome assembly
X0048863biological_processstem cell differentiation
X0061136biological_processregulation of proteasomal protein catabolic process
X0071357biological_processcellular response to type I interferon
X1904813cellular_componentficolin-1-rich granule lumen
Y0000502cellular_componentproteasome complex
Y0005515molecular_functionprotein binding
Y0005576cellular_componentextracellular region
Y0005654cellular_componentnucleoplasm
Y0005829cellular_componentcytosol
Y0005838cellular_componentproteasome regulatory particle
Y0006979biological_processresponse to oxidative stress
Y0008021cellular_componentsynaptic vesicle
Y0010498biological_processproteasomal protein catabolic process
Y0022624cellular_componentproteasome accessory complex
Y0032991cellular_componentprotein-containing complex
Y0034774cellular_componentsecretory granule lumen
Y0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
Y0061136biological_processregulation of proteasomal protein catabolic process
Y0071357biological_processcellular response to type I interferon
Y1904813cellular_componentficolin-1-rich granule lumen
Z0000502cellular_componentproteasome complex
Z0005515molecular_functionprotein binding
Z0005576cellular_componentextracellular region
Z0005634cellular_componentnucleus
Z0005654cellular_componentnucleoplasm
Z0005829cellular_componentcytosol
Z0005838cellular_componentproteasome regulatory particle
Z0006979biological_processresponse to oxidative stress
Z0008021cellular_componentsynaptic vesicle
Z0008233molecular_functionpeptidase activity
Z0008541cellular_componentproteasome regulatory particle, lid subcomplex
Z0010498biological_processproteasomal protein catabolic process
Z0016020cellular_componentmembrane
Z0034774cellular_componentsecretory granule lumen
Z0042803molecular_functionprotein homodimerization activity
Z0043161biological_processproteasome-mediated ubiquitin-dependent protein catabolic process
Z0061136biological_processregulation of proteasomal protein catabolic process
Z0070062cellular_componentextracellular exosome
Z0071357biological_processcellular response to type I interferon
Z0140492molecular_functionmetal-dependent deubiquitinase activity
Z1904813cellular_componentficolin-1-rich granule lumen
Functional Information from PROSITE/UniProt
site_idPS00012
Number of Residues16
DetailsPHOSPHOPANTETHEINE Phosphopantetheine attachment site. AAGFSSSGPAATSGAV
ChainResidueDetails
dALA72-VAL87

site_idPS00674
Number of Residues19
DetailsAAA AAA-protein family signature. VkVImATNradtLDpALl.R
ChainResidueDetails
DVAL305-ARG323
BVAL325-ARG343
CILE289-ARG307
FVAL326-ARG344
EVAL273-ARG291
AILE315-ARG333

Functional Information from SwissProt/UniProt
site_idSWS_FT_FI1
Number of Residues1029
DetailsDomain: {"description":"PCI","evidences":[{"source":"PROSITE-ProRule","id":"PRU01185","evidenceCode":"ECO:0000255"}]}
ChainResidueDetails

site_idSWS_FT_FI2
Number of Residues11
DetailsModified residue: {"description":"N6-acetyllysine","evidences":[{"source":"PubMed","id":"19608861","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

site_idSWS_FT_FI3
Number of Residues35
DetailsBinding site: {"evidences":[{"evidenceCode":"ECO:0000255"}]}
ChainResidueDetails

site_idSWS_FT_FI4
Number of Residues2
DetailsCross-link: {"description":"Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in ubiquitin)","evidences":[{"source":"PubMed","id":"17370265","evidenceCode":"ECO:0000269"}]}
ChainResidueDetails

site_idSWS_FT_FI5
Number of Residues13
DetailsMotif: {"description":"JAMM motif","evidences":[{"source":"PROSITE-ProRule","id":"PRU01182","evidenceCode":"ECO:0000255"}]}
ChainResidueDetails

site_idSWS_FT_FI6
Number of Residues3
DetailsBinding site: {"evidences":[{"source":"PROSITE-ProRule","id":"PRU01182","evidenceCode":"ECO:0000255"}]}
ChainResidueDetails

site_idSWS_FT_FI7
Number of Residues1
DetailsModified residue: {"description":"Phosphoserine","evidences":[{"source":"PubMed","id":"17693683","evidenceCode":"ECO:0007744"},{"source":"PubMed","id":"24275569","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

site_idSWS_FT_FI8
Number of Residues1
DetailsModified residue: {"description":"Phosphoserine","evidences":[{"source":"PubMed","id":"17323924","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

site_idSWS_FT_FI9
Number of Residues1
DetailsModified residue: {"description":"Phosphothreonine","evidences":[{"source":"PubMed","id":"23186163","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

site_idSWS_FT_FI10
Number of Residues2
DetailsCross-link: {"description":"Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in ubiquitin)"}
ChainResidueDetails

site_idSWS_FT_FI11
Number of Residues33
DetailsRepeat: {"description":"PC 2"}
ChainResidueDetails

site_idSWS_FT_FI12
Number of Residues34
DetailsRepeat: {"description":"PC 3"}
ChainResidueDetails

site_idSWS_FT_FI13
Number of Residues34
DetailsRepeat: {"description":"PC 4"}
ChainResidueDetails

site_idSWS_FT_FI14
Number of Residues33
DetailsRepeat: {"description":"PC 5"}
ChainResidueDetails

site_idSWS_FT_FI15
Number of Residues35
DetailsRepeat: {"description":"PC 6"}
ChainResidueDetails

site_idSWS_FT_FI16
Number of Residues32
DetailsRepeat: {"description":"PC 7"}
ChainResidueDetails

site_idSWS_FT_FI17
Number of Residues34
DetailsRepeat: {"description":"PC 8"}
ChainResidueDetails

site_idSWS_FT_FI18
Number of Residues40
DetailsRepeat: {"description":"PC 9"}
ChainResidueDetails

site_idSWS_FT_FI19
Number of Residues32
DetailsRepeat: {"description":"PC 10"}
ChainResidueDetails

site_idSWS_FT_FI20
Number of Residues1
DetailsModified residue: {"description":"N6-acetyllysine","evidences":[{"source":"UniProtKB","id":"Q3TXS7","evidenceCode":"ECO:0000250"}]}
ChainResidueDetails

site_idSWS_FT_FI21
Number of Residues2
DetailsCross-link: {"description":"Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternate","evidences":[{"source":"PubMed","id":"25114211","evidenceCode":"ECO:0007744"},{"source":"PubMed","id":"25218447","evidenceCode":"ECO:0007744"},{"source":"PubMed","id":"25772364","evidenceCode":"ECO:0007744"},{"source":"PubMed","id":"28112733","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

site_idSWS_FT_FI22
Number of Residues5
DetailsModified residue: {"description":"Phosphoserine","evidences":[{"source":"PubMed","id":"23186163","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

site_idSWS_FT_FI32
Number of Residues2
DetailsCross-link: {"description":"Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)","evidences":[{"source":"PubMed","id":"25772364","evidenceCode":"ECO:0007744"},{"source":"PubMed","id":"28112733","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

site_idSWS_FT_FI33
Number of Residues2
DetailsCross-link: {"description":"Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2); alternate","evidences":[{"source":"PubMed","id":"28112733","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

site_idSWS_FT_FI34
Number of Residues4
DetailsCross-link: {"description":"Glycyl lysine isopeptide (Lys-Gly) (interchain with G-Cter in SUMO2)","evidences":[{"source":"PubMed","id":"28112733","evidenceCode":"ECO:0007744"}]}
ChainResidueDetails

253795

PDB entries from 2026-05-20

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