8USB: 26S proteasome non-ATPase regulatory subunit 12

Non-substrate-engaged human 26S proteasome with Nub1/FAT10 bound to Rpn1. Determined by electron microscopy at 2.73 Å resolution. Released 6 Nov 2024.

Method
Electron microscopy
Resolution
2.73 Å
Organism
Homo sapiens
Chains
27
Atoms
70,143
Mol. weight
1211.82 kDa
Ligands
ZN, ATP, MG, ADP
Released
6 Nov 2024

Explore 8USB in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

8USB contains 507 α-helices and 274 β-strands across 27 chains. Residue ranges use author residue numbering, as in the PDB file, from PDBe. To see them in 3D, choose the Cartoon representation with Secondary structure coloring: helices, sheets and coils get different colors.

Chain a: 25 helices, 3 β-strands

ElementResiduesLengthSheet
α-helix6-138
α-helix19-3113
α-helix34-4512
α-helix55-595
α-helix60-645
α-helix65-673
α-helix68-703
α-helix73-8614
α-helix90-10314
α-helix108-12417
α-helix128-14417
α-helix152-16615
α-helix169-18214
α-helix190-20617
α-helix213-2175
α-helix220-2256
α-helix231-24111
α-helix245-2506
α-helix252-2554
α-helix259-2635
α-helix265-28319
α-helix286-2883
β-strand290-29232
α-helix293-2997
α-helix304-31714
β-strand320-32562
β-strand330-33672
α-helix343-37028
Chain A: 26 helices, 19 β-strands
ElementResiduesLengthSheet
α-helix42-6322
β-strand72117
α-helix73-742
α-helix75-773
α-helix80-8910
β-strand93-102103
β-strand110-11563
β-strand119-12353
β-strand12513
α-helix131-1333
β-strand139-14243
β-strand149-15243
α-helix154-1563
α-helix159-1624
β-strand165120
α-helix174-1763
α-helix181-19111
α-helix193-1964
α-helix199-2024
β-strand211-215520
α-helix222-23211
β-strand236-241620
α-helix242-2454
α-helix252-26413
β-strand270-275620
α-helix277-2793
β-strand282121
α-helix283-2842
β-strand287122
β-strand289122
α-helix293-30614
β-strand315-321720
α-helix324-3263
β-strand327121
β-strand339-342420
α-helix344-3463
α-helix348-35811
β-strand364123
α-helix370-3756
α-helix382-39817
β-strand404123
α-helix406-4138
α-helix414-4185
α-helix419-4235
α-helix426-4294
Chain b: 9 helices, 7 β-strands
ElementResiduesLengthSheet
β-strand416
β-strand5-1067
α-helix14-174
α-helix25-4319
β-strand48-5367
β-strand59-6577
α-helix69-768
α-helix86-9813
β-strand10616
α-helix1071
β-strand109-11357
α-helix121-13313
β-strand138-14257
α-helix146-1494
α-helix152-16110
α-helix180-1834
Chain B: 26 helices, 14 β-strands
ElementResiduesLengthSheet
α-helix37-382
α-helix45-473
α-helix59-8527
α-helix92-10312
β-strand107-11483
β-strand120-12453
β-strand130-13563
β-strand139117
α-helix141-1433
β-strand149-15353
β-strand159-16243
α-helix165-1684
α-helix169-1746
β-strand175118
α-helix184-1863
α-helix191-1977
α-helix198-2025
α-helix203-2064
α-helix208-2147
α-helix217-2193
β-strand221-225518
α-helix232-24110
β-strand246-251618
α-helix252-2554
α-helix262-27615
β-strand280-285618
α-helix289-2924
α-helix302-31615
α-helix321-3222
β-strand325-331718
β-strand349-352418
α-helix354-3563
α-helix358-36811
β-strand374119
α-helix380-3834
α-helix392-40817
β-strand414119
α-helix4151
α-helix416-4227
α-helix423-4275
Chain c: 18 helices, 10 β-strands
ElementResiduesLengthSheet
α-helix24-252
β-strand26167
β-strand30-34568
α-helix35-4713
β-strand54-631068
β-strand66-74968
α-helix75-773
α-helix84-874
α-helix89-9911
α-helix100-1023
β-strand107-114868
α-helix124-13613
β-strand141-1511168
β-strand154-162968
α-helix163-1642
α-helix165-1695
β-strand176167
α-helix181-1833
α-helix186-1883
α-helix189-1935
β-strand196168
β-strand200-208968
α-helix211-2188
α-helix225-2295
α-helix230-2334
α-helix234-26330
α-helix269-2724
α-helix280-30829
Chain C: 25 helices, 13 β-strands
ElementResiduesLengthSheet
α-helix21-6646
α-helix69-702
β-strand71-80103
β-strand84-8963
β-strand94-9743
β-strand9913
α-helix105-1073
β-strand113-11643
β-strand123-12643
α-helix127-1293
α-helix133-1419
α-helix143-1453
α-helix148-1503
α-helix155-1617
α-helix162-1665
α-helix167-1704
α-helix172-1787
α-helix181-1833
β-strand185-18954
α-helix196-20611
β-strand210-21344
α-helix221-2244
α-helix231-2344
α-helix235-2406
β-strand244-24964
α-helix268-28215
β-strand289-29574
α-helix303-3064
β-strand313-31644
α-helix318-3203
α-helix322-33312
β-strand33815
α-helix3391
α-helix344-3496
α-helix356-37217
β-strand37815
α-helix380-3878
α-helix388-3936
Chain d: 15 helices, 4 β-strands
ElementResiduesLengthSheet
α-helix87-10317
α-helix109-12517
α-helix138-15518
α-helix161-17616
α-helix178-1803
α-helix188-20114
α-helix205-2128
α-helix217-2226
α-helix224-23815
α-helix241-2466
α-helix254-2563
α-helix257-27822
β-strand27918
β-strand281-28339
α-helix287-2904
α-helix297-30610
β-strand31019
β-strand315-31739
α-helix332-34716
Chain D: 21 helices, 15 β-strands
ElementResiduesLengthSheet
α-helix40-7839
α-helix79-835
α-helix841
α-helix861
β-strand87-9483
β-strand100-10563
β-strand110-11563
α-helix121-1233
β-strand128-13363
β-strand139-14243
α-helix171-1777
α-helix178-1825
α-helix183-1864
α-helix188-1947
β-strand201-205524
α-helix212-22110
β-strand226-231624
α-helix232-2343
α-helix242-25615
β-strand260-265624
α-helix267-2693
β-strand272125
α-helix280-29718
β-strand305-311724
α-helix314-3163
β-strand317125
α-helix319-3224
β-strand329-332424
α-helix338-34912
β-strand354126
α-helix361-3644
α-helix372-38817
β-strand39111
β-strand394126
α-helix396-40712
α-helix415-4173

19 more chain groups are not listed. Open the entry in the viewer and use the sequence panel to see them.

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
26S proteasome non-ATPase regulatory subunit 12Wprotein456Homo sapiensO00232 (AlphaFold model)
26S protease regulatory subunit 8Cprotein406Homo sapiensP62195 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 4bprotein377Homo sapiensP55036 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 8dprotein350Homo sapiensP48556 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 2fprotein908Homo sapiensQ13200
NEDD8 ultimate buster 1gprotein605Homo sapiensH3BM14
26S proteasome regulatory subunit 4Bprotein440Homo sapiensP62191
26S proteasome regulatory subunit 7Aprotein433Homo sapiensP35998
26S proteasome regulatory subunit 6BDprotein418Homo sapiensP43686
26S protease regulatory subunit 10BEprotein389Homo sapiensP62333
26S proteasome regulatory subunit 6AFprotein439Homo sapiensP17980
Proteasome subunit alpha type-6Gprotein246Homo sapiensP60900

15 more molecules are not listed.

Sequence of entity 1 (W), FASTA
>8USB_1 26S proteasome non-ATPase regulatory subunit 12 (chains W)
MADGGSERADGRIVKMEVDYSATVDQRLPECAKLAKEGRLQEVIETLLSLEKQTRTASDM
VSTSRILVAVVKMCYEAKEWDLLNENIMLLSKRRSQLKQAVAKMVQQCCTYVEEITDLPI
KLRLIDTLRMVTEGKIYVEIERARLTKTLATIKEQNGDVKEAASILQELQVETYGSMEKK
ERVEFILEQMRLCLAVKDYIRTQIISKKINTKFFQEENTEKLKLKYYNLMIQLDQHEGSY
LSICKHYRAIYDTPCIQAESEKWQQALKSVVLYVILAPFDNEQSDLVHRISGDKKLEEIP
KYKDLLKLFTTMELMRWSTLVEDYGMELRKGSLESPATDVFGSTEEGEKRWKDLKNRVVE
HNIRIMAKYYTRITMKRMAQLLDLSVDESEAFLSNLVVNKTIFAKVDRLAGIINFQRPKD
PNNLLNDWSQKLNSLMSLVNKTTHLIAKEEMIHNLQ
Sequence of entity 2 (C), FASTA
>8USB_2 26S protease regulatory subunit 8 (chains C)
MALDGPEQMELEEGKAGSGLRQYYLSKIEELQLIVNDKSQNLRRLQAQRNELNAKVRLLR
EELQLLQEQGSYVGEVVRAMDKKKVLVKVHPEGKFVVDVDKNIDINDVTPNCRVALRNDS
YTLHKILPNKVDPLVSLMMVEKVPDSTYEMIGGLDKQIKEIKEVIELPVKHPELFEALGI
AQPKGVLLYGPPGTGKTLLARAVAHHTDCTFIRVSGSELVQKFIGEGARMVRELFVMARE
HAPSIIFMDEIDSIGSSRLEGGSGGDSEVQRTMLELLNQLDGFEATKNIKVIMATNRIDI
LDSALLRPGRIDRKIEFPPPNEEARLDILKIHSRKMNLTRGINLRKIAELMPGASGAEVK
GVCTEAGMYALRERRVHVTQEDFEMAVAKVMQKDSEKNMSIKKLWK
Sequence of entity 3 (b), FASTA
>8USB_3 26S proteasome non-ATPase regulatory subunit 4 (chains b)
MVLESTMVCVDNSEYMRNGDFLPTRLQAQQDAVNIVCHSKTRSNPENNVGLITLANDCEV
LTTLTPDTGRILSKLHTVQPKGKITFCTGIRVAHLALKHRQGKNHKMRIIAFVGSPVEDN
EKDLVKLAKRLKKEKVNVDIINFGEEEVNTEKLTAFVNTLNGKDGTGSHLVTVPPGPSLA
DALISSPILAGEGGAMLGLGASDFEFGVDPSADPELALALRVSMEEQRQRQEEEARRAAA
ASAAEAGIATTGTEDSDDALLKMTISQQEFGRTGLPDLSSMTEEEQIAYAMQMSLQGAEF
GQAESADIDASSAMDTSEPAKEEDDYDVMQDPEFLQSVLENLPGVDPNNEAIRNAMGSLA
SQATKDGKKDKKEEDKK
Sequence of entity 4 (d), FASTA
>8USB_4 26S proteasome non-ATPase regulatory subunit 8 (chains d)
MFIKGRAPRAPPRERRRATRGGLRQVVAPPRALGSTSRPHFRRASVCRRRCRKSGGLLAA
SRKMAAAAVNGAAGFSSSGPAATSGAVLQAATGMYEQLKGEWNRKSPNLSKCGEELGRLK
LVLLELNFLPTTGTKLTKQQLILARDILEIGAQWSILRKDIPSFERYMAQLKCYYFDYKE
QLPESAYMHQLLGLNLLFLLSQNRVAEFHTELERLPAKDIQTNVYIKHPVSLEQYLMEGS
YNKVFLAKGNIPAESYTFFIDILLDTIRDEIAGCIEKAYEKILFTEATRILFFNTPKKMT
DYAKKRGWVLGPNNYYSFASQQQKPEDTTIPSTELAKQVIEYARQLEMIV
Sequence of entity 5 (f), FASTA
>8USB_5 26S proteasome non-ATPase regulatory subunit 2 (chains f)
MEEGGRDKAPVQPQQSPAAAPGGTDEKPSGKERRDAGDKDKEQELSEEDKQLQDELEMLV
ERLGEKDTSLYRPALEELRRQIRSSTTSMTSVPKPLKFLRPHYGKLKEIYENMAPGENKR
FAADIISVLAMTMSGERECLKYRLVGSQEELASWGHEYVRHLAGEVAKEWQELDDAEKVQ
REPLLTLVKEIVPYNMAHNAEHEACDLLMEIEQVDMLEKDIDENAYAKVCLYLTSCVNYV
PEPENSALLRCALGVFRKFSRFPEALRLALMLNDMELVEDIFTSCKDVVVQKQMAFMLGR
HGVFLELSEDVEEYEDLTEIMSNVQLNSNFLALARELDIMEPKVPDDIYKTHLENNRFGG
SGSQVDSARMNLASSFVNGFVNAAFGQDKLLTDDGNKWLYKNKDHGMLSAAASLGMILLW
DVDGGLTQIDKYLYSSEDYIKSGALLACGIVNSGVRNECDPALALLSDYVLHNSNTMRLG
SIFGLGLAYAGSNREDVLTLLLPVMGDSKSSMEVAGVTALACGMIAVGSCNGDVTSTILQ
TIMEKSETELKDTYARWLPLGLGLNHLGKGEAIEAILAALEVVSEPFRSFANTLVDVCAY
AGSGNVLKVQQLLHICSEHFDSKEKEEDKDKKEKKDKDKKEAPADMGAHQGVAVLGIALI
AMGEEIGAEMALRTFGHLLRYGEPTLRRAVPLALALISVSNPRLNILDTLSKFSHDADPE
VSYNSIFAMGMVGSGTNNARLAAMLRQLAQYHAKDPNNLFMVRLAQGLTHLGKGTLTLCP
YHSDRQLMSQVAVAGLLTVLVSFLDVRNIILGKSHYVLYGLVAAMQPRMLVTFDEELRPL
PVSVRVGQAVDVVGQAGKPKTITGFQTHTTPVLLAHGERAELATEEFLPVTPILEGFVIL
RKNPNYDL
Sequence of entity 6 (g), FASTA
>8USB_6 NEDD8 ultimate buster 1 (chains g)
GPLGMAQKKYLQAKLTQFLREDRIQLWKPPYTDENKKVGLALKDLAKQYSDRLECCENEV
EKVIEEIRCKAIERGTGNDNYRTTGIATIEVFLPPRLKKDRKNLLETRLHITGRELRSKI
AETFGLQENYIKIVINKKQLQLGKTLEEQGVAHNVKAMVLELKQSEEDARKNFQLEEEEQ
NEAKLKEKQIQRTKRGLEILAKRAAETVVDPEMTPYLDIANQTGRSIRIPPSERKALMLA
MGYHEKGRAFLKRKEYGIALPCLLDADKYFCECCRELLDTVDNYAVLQLDIVWCYFRLEQ
LECLDDAEKKLNLAQKCFKNCYGENHQRLVHIKGNCGKEKVLFLRLYLLQGIRNYHSGND
VEAYEYLNKARQLFKELYIDPSKVDNLLQLGFTAQEARLGLRACDGNVDHAATHITNRRE
ELAQIRKEEKEKKRRRLENIRFLKGMGYSTHAAQQILLSNPQMWWLNDSNPETDNRQESP
SQENIDRLVYMGFDALVAEAALRVFRGNVQLAAQTLAHNGGSLPPELPLSPEDSLSPPAT
SPSDSAGTSSASTDEDMETEAVNEILEDIPEHEEDYLDSTLEDEEIIIAEYLSYVENRKS
ATKKN
Sequence of entity 7 (B), FASTA
>8USB_7 26S proteasome regulatory subunit 4 (chains B)
MGQSQSGGHGPGGGKKDDKDKKKKYEPPVPTRVGKKKKKTKGPDAASKLPLVTPHTQCRL
KLLKLERIKDYLLMEEEFIRNQEQMKPLEEKQEEERSKVDDLRGTPMSVGTLEEIIDDNH
AIVSTSVGSEHYVSILSFVDKDLLEPGCSVLLNHKVHAVIGVLMDDTDPLVTVMKVEKAP
QETYADIGGLDNQIQEIKESVELPLTHPEYYEEMGIKPPKGVILYGPPGTGKTLLAKAVA
NQTSATFLRVVGSELIQKYLGDGPKLVRELFRVAEEHAPSIVFIDEIDAIGTKRYDSNSG
GEREIQRTMLELLNQLDGFDSRGDVKVIMATNRIETLDPALIRPGRIDRKIEFPLPDEKT
KKRIFQIHTSRMTLADDVTLDDLIMAKDDLSGADIKAICTEAGLMALRERRMKVTNEDFK
KSKENVLYKKQEGTPEGLYL
Sequence of entity 8 (A), FASTA
>8USB_8 26S proteasome regulatory subunit 7 (chains A)
MPDYLGADQRKTKEDEKDDKPIRALDEGDIALLKTYGQSTYSRQIKQVEDDIQQLLKKIN
ELTGIKESDTGLAPPALWDLAADKQTLQSEQPLQVARCTKIINADSEDPKYIINVKQFAK
FVVDLSDQVAPTDIEEGMRVGVDRNKYQIHIPLPPKIDPTVTMMQVEEKPDVTYSDVGGC
KEQIEKLREVVETPLLHPERFVNLGIEPPKGVLLFGPPGTGKTLCARAVANRTDACFIRV
IGSELVQKYVGEGARMVRELFEMARTKKACLIFFDEIDAIGGARFDDGAGGDNEVQRTML
ELINQLDGFDPRGNIKVLMATNRPDTLDPALMRPGRLDRKIEFSLPDLEGRTHIFKIHAR
SMSVERDIRFELLARLCPNSTGAEIRSVCTEAGMFAIRARRKIATEKDFLEAVNKVIKSY
AKFSATPRYMTYN
Sequence of entity 9 (D), FASTA
>8USB_9 26S proteasome regulatory subunit 6B (chains D)
MEEIGILVEKAQDEIPALSVSRPQTGLSFLGPEPEDLEDLYSRYKKLQQELEFLEVQEEY
IKDEQKNLKKEFLHAQEEVKRIQSIPLVIGQFLEAVDQNTAIVGSTTGSNYYVRILSTID
RELLKPNASVALHKHSNALVDVLPPEADSSIMMLTSDQKPDVMYADIGGMDIQKQEVREA
VELPLTHFELYKQIGIDPPRGVLMYGPPGCGKTMLAKAVAHHTTAAFIRVVGSEFVQKYL
GEGPRMVRDVFRLAKENAPAIIFIDEIDAIATKRFDAQTGADREVQRILLELLNQMDGFD
QNVNVKVIMATNRADTLDPALLRPGRLDRKIEFPLPDRRQKRLIFSTITSKMNLSEEVDL
EDYVARPDKISGADINSICQESGMLAVRENRYIVLAKDFEKAYKTVIKKDEQEHEFYK
Sequence of entity 10 (E), FASTA
>8USB_10 26S protease regulatory subunit 10B (chains E)
MADPRDKALQDYRKKLLEHKEIDGRLKELREQLKELTKQYEKSENDLKALQSVGQIVGEV
LKQLTEEKFIVKATNGPRYVVGCRRQLDKSKLKPGTRVALDMTTLTIMRYLPREVDPLVY
NMSHEDPGNVSYSEIGGLSEQIRELREVIELPLTNPELFQRVGIIPPKGCLLYGPPGTGK
TLLARAVASQLDCNFLKVVSSSIVDKYIGESARLIREMFNYARDHQPCIIFMDEIDAIGG
RRFSEGTSADREIQRTLMELLNQMDGFDTLHRVKMIMATNRPDTLDPALLRPGRLDRKIH
IDLPNEQARLDILKIHAGPITKHGEIDYEAIVKLSDGFNGADLRNVCTEAGMFAIRADHD
FVVQEDFMKAVRKVADSKKLESKLDYKPV
Sequence of entity 11 (F), FASTA
>8USB_11 26S proteasome regulatory subunit 6A (chains F)
MNLLPNIESPVTRQEKMATVWDEAEQDGIGEEVLKMSTEEIIQRTRLLDSEIKIMKSEVL
RVTHELQAMKDKIKENSEKIKVNKTLPYLVSNVIELLDVDPNDQEEDGANIDLDSQRKGK
CAVIKTSTRQTYFLPVIGLVDAEKLKPGDLVGVNKDSYLILETLPTEYDSRVKAMEVDER
PTEQYSDIGGLDKQIQELVEAIVLPMNHKEKFENLGIQPPKGVLMYGPPGTGKTLLARAC
AAQTKATFLKLAGPQLVQMFIGDGAKLVRDAFALAKEKAPSIIFIDELDAIGTKRFDSEK
AGDREVQRTMLELLNQLDGFQPNTQVKVIAATNRVDILDPALLRSGRLDRKIEFPMPNEE
ARARIMQIHSRKMNVSPDVNYEELARCTDDFNGAQCKAVCVEAGMIALRRGATELTHEDY
MEGILEVQAKKKANLQYYA
Sequence of entity 12 (G), FASTA
>8USB_12 Proteasome subunit alpha type-6 (chains G)
MSRGSSAGFDRHITIFSPEGRLYQVEYAFKAINQGGLTSVAVRGKDCAVIVTQKKVPDKL
LDSSTVTHLFKITENIGCVMTGMTADSRSQVQRARYEAANWKYKYGYEIPVDMLCKRIAD
ISQVYTQNAEMRPLGCCMILIGIDEEQGPQVYKCDPAGYYCGFKATAAGVKQTESTSFLE
KKVKKKFDWTFEQTVETAITCLSTVLSIDFKPSEIEVGVVTVENPKFRILTEAEIDAHLV
ALAERD

Ligands and cofactors

IDNameFormulaCopies
ZNZinc ionZn1
ATPAdenosine-5'-triphosphateC10 H16 N5 O13 P35
MGMagnesium ionMg6
ADPAdenosine-5'-diphosphateC10 H15 N5 O10 P21

Primary citation

NUB1 traps unfolded FAT10 for ubiquitin-independent degradation by the 26S proteasome. Arkinson, C., Dong, K.C., Gee, C.L. et al. Nat Struct Mol Biol (2025) 32:1752-1765. DOI 10.1038/s41594-025-01527-3 · PubMed

Other PDB entries of the same protein (UniProt O00232 (AlphaFold model), which also has an AlphaFold model), best resolution first:

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