9BV3: M1B Midnolin-Proteasome

M1B Midnolin-Proteasome. Determined by electron microscopy at 2.9 Å resolution. Released 11 Jun 2025.

Method
Electron microscopy
Resolution
2.9 Å
Organism
Homo sapiens
Chains
40
Atoms
88,575
Mol. weight
1530.51 kDa
Ligands
ADP, MG, ATP
Released
11 Jun 2025

Explore 9BV3 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9BV3 contains 524 α-helices and 466 β-strands across 40 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: 19 helices, 3 β-strands

ElementResiduesLengthSheet
α-helix3-1412
α-helix19-3113
α-helix37-4610
α-helix55-6713
α-helix73-8513
α-helix90-10314
α-helix108-12518
α-helix129-14315
α-helix152-16615
α-helix169-18214
α-helix190-20617
α-helix214-2185
α-helix234-2418
α-helix245-2506
α-helix252-2554
α-helix265-28319
β-strand290-292322
α-helix293-3008
α-helix304-31714
β-strand320-325622
β-strand330-336722
α-helix343-37028
Chain A: 25 helices, 19 β-strands
ElementResiduesLengthSheet
α-helix45-6319
β-strand72162
α-helix73-742
α-helix75-773
α-helix80-8910
β-strand93-94263
β-strand97164
β-strand98-102565
β-strand110-115665
β-strand119-123565
α-helix131-1333
β-strand139164
β-strand142-143263
β-strand153164
α-helix154-1563
α-helix159-1624
β-strand165166
α-helix169-1702
α-helix174-1763
α-helix181-19616
α-helix199-2024
β-strand212-215466
α-helix222-23211
β-strand236-241666
α-helix242-2454
α-helix253-26614
β-strand270-275666
α-helix277-2793
α-helix293-30614
β-strand308167
β-strand310167
β-strand315-321766
α-helix329-3324
β-strand341-342266
α-helix344-3463
α-helix348-35912
β-strand364168
α-helix370-3767
α-helix382-39817
β-strand404168
α-helix406-4138
α-helix414-4185
α-helix419-4235
α-helix426-4294
Chain b: 8 helices, 6 β-strands
ElementResiduesLengthSheet
β-strand4-11845
α-helix14-174
α-helix25-4218
β-strand48-54745
β-strand59-65745
α-helix69-768
α-helix87-9812
β-strand106-112745
α-helix121-13313
β-strand137-143745
α-helix150-16112
β-strand169-173545
α-helix181-1844
α-helix187-1904
Chain B: 23 helices, 15 β-strands
ElementResiduesLengthSheet
α-helix60-8526
α-helix92-10312
β-strand106-1171263
β-strand120-124563
β-strand130-133463
β-strand135163
β-strand139162
β-strand149-153563
β-strand159-162463
α-helix167-1682
α-helix169-1746
β-strand175173
α-helix184-1863
α-helix191-1977
α-helix198-2025
α-helix203-2064
α-helix208-2147
α-helix217-2193
β-strand221-225573
α-helix232-24211
β-strand246-251673
α-helix252-2554
α-helix262-27615
β-strand280-285673
α-helix287-2893
α-helix302-31615
α-helix321-3222
β-strand325-331773
α-helix334-3363
α-helix339-3424
β-strand350-352373
α-helix354-3563
α-helix358-36912
β-strand374174
α-helix382-3854
α-helix392-40817
β-strand414174
α-helix416-42510
Chain c: 11 helices, 12 β-strands
ElementResiduesLengthSheet
β-strand26149
β-strand30-33450
α-helix35-4511
β-strand54-60750
β-strand66-74950
α-helix89-10113
β-strand107-114850
α-helix124-13613
β-strand141-145550
β-strand151151
β-strand154151
β-strand157-162650
α-helix163-1642
α-helix165-1695
β-strand176149
α-helix189-1946
β-strand196150
β-strand200-207850
α-helix213-2197
α-helix225-2295
α-helix230-2334
α-helix234-26330
α-helix280-30829
Chain C: 27 helices, 12 β-strands
ElementResiduesLengthSheet
α-helix20-6546
α-helix69-702
β-strand71-79963
β-strand84-89663
β-strand94-99663
α-helix1001
α-helix105-1073
β-strand112-116563
β-strand122-126563
α-helix1271
α-helix129-1324
α-helix133-1419
α-helix143-1453
α-helix155-1617
α-helix162-1665
α-helix167-1704
α-helix174-1785
α-helix181-1833
β-strand185-189580
α-helix196-20712
β-strand210-214580
α-helix218-2203
α-helix221-2244
α-helix231-2333
α-helix234-2407
β-strand244-249680
α-helix268-28013
β-strand289-295780
α-helix303-3064
β-strand313-316480
α-helix3181
α-helix3201
α-helix322-33312
β-strand338181
α-helix3391
α-helix344-3507
α-helix356-37217
β-strand378181
α-helix380-39011
Chain d: 16 helices, 2 β-strands
ElementResiduesLengthSheet
α-helix84-10421
α-helix110-12314
α-helix1401
α-helix141-1466
α-helix147-15812
α-helix161-17717
α-helix187-20216
α-helix205-21410
α-helix217-2215
α-helix224-23815
α-helix241-25010
α-helix254-2563
α-helix257-27721
β-strand282-283255
α-helix285-2917
α-helix296-3049
β-strand315-316255
α-helix332-34716
Chain D: 22 helices, 16 β-strands
ElementResiduesLengthSheet
α-helix40-5112
α-helix52-576
α-helix58-7922
α-helix80-823
β-strand87-95963
β-strand100-105663
β-strand110-115663
α-helix121-1233
β-strand128-132563
β-strand139-142463
α-helix151-1533
α-helix164-1674
α-helix171-1777
α-helix178-1825
α-helix184-19411
β-strand205182
α-helix212-22211
β-strand226-231683
α-helix232-2354
α-helix244-25613
β-strand260-265683
α-helix267-2693
β-strand272184
α-helix281-29717
β-strand305-309583
β-strand311182
α-helix314-3163
β-strand317184
β-strand332182
α-helix334-3363
α-helix338-35013
β-strand354185
α-helix361-3644
α-helix372-38817
β-strand391121
β-strand394185
α-helix396-40611
α-helix415-4173

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

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
Midnolinyprotein567Homo sapiensQ504T8 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 1Uprotein953Homo sapiensQ99460 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 3Vprotein534Homo sapiensO43242 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 12Wprotein456Homo sapiensO00232 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 11Xprotein422Homo sapiensO00231
26S proteasome non-ATPase regulatory subunit 6Yprotein389Homo sapiensQ15008
26S proteasome non-ATPase regulatory subunit 7Zprotein324Homo sapiensP51665
26S proteasome non-ATPase regulatory subunit 13aprotein376Homo sapiensQ9UNM6
26S proteasome non-ATPase regulatory subunit 4bprotein377Homo sapiensP55036
26S proteasome non-ATPase regulatory subunit 14cprotein310Homo sapiensO00487
26S proteasome non-ATPase regulatory subunit 8dprotein350Homo sapiensP48556
26S proteasome complex subunit SEM1eprotein70Homo sapiensP60896

21 more molecules are not listed.

Sequence of entity 1 (y), FASTA
>9BV3_1 Midnolin (chains y)
MDYKDDDDKDYKDDDDKGGGGSGGGGFETSLYKKAGLATMEKVLVETSYPSQTTRLPPIT
YTGRFSLEPAPNSGNTLWPEPLFSLVSGLVSMTNPPASSSEPQPGGARSCRRGAPGGACE
LGPAAEAAPMSLAIHSTTGTRYDLAVPPDETVEGLRKRLSQRLKVPKERLALLHKDTRLS
SGKLQEFGVGDGSKLTLVPTVEAGLMSQASRPEQSVMQALESLTETQVSDFLSGRSPLTL
ALRVGDHMMFVQLQLAAQHAPLQHRHVLAAAAAAAAARGDPSIASPVSSPCRPVSSAARV
PPVPTSPSPASPSPITAGSFRSHAASTTCPEQMDCSPTASSSASPGASTTSTPGASPAPR
SRKPGAVIESFVNHAPGVFSGTFSGTLHPNCQDSSGRPRRDIGTILQILNDLLSATRHYQ
GMPPSLAQLRCHAQCSPASPAPDLAPRTTSCEKLTAAPSASLLQGQSQIRMCKPPGDRLR
QTENRATRCKVERLQLLLQQKRLRRKARRDARGPYHWSPSRKAGRSDSSSSGGGGSPSEA
SGLGLDFEDSVWKPEANPDIKSEFVVA
Sequence of entity 2 (U), FASTA
>9BV3_2 26S proteasome non-ATPase regulatory subunit 1 (chains U)
MITSAAGIISLLDEDEPQLKEFALHKLNAVVNDFWAEISESVDKIEVLYEDEGFRSRQFA
ALVASKVFYHLGAFEESLNYALGAGDLFNVNDNSEYVETIIAKCIDHYTKQCVENADLPE
GEKKPIDQRLEGIVNKMFQRCLDDHKYKQAIGIALETRRLDVFEKTILESNDVPGMLAYS
LKLCMSLMQNKQFRNKVLRVLVKIYMNLEKPDFINVCQCLIFLDDPQAVSDILEKLVKED
NLLMAYQICFDLYESASQQFLSSVIQNLRTVGTPIASVPGSTNTGTVPGSEKDSDSMETE
EKTSSAFVGKTPEASPEPKDQTLKMIKILSGEMAIELHLQFLIRNNNTDLMILKNTKDAV
RNSVCHTATVIANSFMHCGTTSDQFLRDNLEWLARATNWAKFTATASLGVIHKGHEKEAL
QLMATYLPKDTSPGSAYQEGGGLYALGLIHANHGGDIIDYLLNQLKNASNDIVRHGGSLG
LGLAAMGTARQDVYDLLKTNLYQDDAVTGEAAGLALGLVMLGSKNAQAIEDMVGYAQETQ
HEKILRGLAVGIALVMYGRMEEADALIESLCRDKDPILRRSGMYTVAMAYCGSGNNKAIR
RLLHVAVSDVNDDVRRAAVESLGFILFRTPEQCPSVVSLLSESYNPHVRYGAAMALGICC
AGTGNKEAINLLEPMTNDPVNYVRQGALIASALIMIQQTEITCPKVNQFRQLYSKVINDK
HDDVMAKFGAILAQGILDAGGHNVTISLQSRTGHTHMPSVVGVLVFTQFWFWFPLSHFLS
LAYTPTCVIGLNKDLKMPKVQYKSNCKPSTFAYPAPLEVPKEKEKEKVSTAVLSITAKAK
KKEKEKEKKEEEKMEVDEAEKKEEKEKKKEPEPNFQLLDNPARVMPAQLKVLTMPETCRY
QPFKPLSIGGIIILKDTSEDIEELVEPVAAHGPKIEEEEQEPEPPEPFEYIDD
Sequence of entity 3 (V), FASTA
>9BV3_3 26S proteasome non-ATPase regulatory subunit 3 (chains V)
MKQEGSARRRGADKAKPPPGGGEQEPPPPPAPQDVEMKEEAATGGGSTGEADGKTAAAAA
EHSQRELDTVTLEDIKEHVKQLEKAVSGKEPRFVLRALRMLPSTSRRLNHYVLYKAVQGF
FTSNNATRDFLLPFLEEPMDTEADLQFRPRTGKAASTPLLPEVEAYLQLLVVIFMMNSKR
YKEAQKISDDLMQKISTQNRRALDLVAAKCYYYHARVYEFLDKLDVVRSFLHARLRTATL
RHDADGQATLLNLLLRNYLHYSLYDQAEKLVSKSVFPEQANNNEWARYLYYTGRIKAIQL
EYSEARRTMTNALRKAPQHTAVGFKQTVHKLLIVVELLLGEIPDRLQFRQPSLKRSLMPY
FLLTQAVRTGNLAKFNQVLDQFGEKFQADGTYTLIIRLRHNVIKTGVRMISLSYSRISLA
DIAQKLQLDSPEDAEFIVAKAIRDGVIEASINHEKGYVQSKEMIDIYSTREPQLAFHQRI
SFCLDIHNMSVKAMRFPPKSYNKDLESAEERREREQQDLEFAKEMAEDDDDSFP
Sequence of entity 4 (W), FASTA
>9BV3_4 26S proteasome non-ATPase regulatory subunit 12 (chains W)
MADGGSERADGRIVKMEVDYSATVDQRLPECAKLAKEGRLQEVIETLLSLEKQTRTASDM
VSTSRILVAVVKMCYEAKEWDLLNENIMLLSKRRSQLKQAVAKMVQQCCTYVEEITDLPI
KLRLIDTLRMVTEGKIYVEIERARLTKTLATIKEQNGDVKEAASILQELQVETYGSMEKK
ERVEFILEQMRLCLAVKDYIRTQIISKKINTKFFQEENTEKLKLKYYNLMIQLDQHEGSY
LSICKHYRAIYDTPCIQAESEKWQQALKSVVLYVILAPFDNEQSDLVHRISGDKKLEEIP
KYKDLLKLFTTMELMRWSTLVEDYGMELRKGSLESPATDVFGSTEEGEKRWKDLKNRVVE
HNIRIMAKYYTRITMKRMAQLLDLSVDESEAFLSNLVVNKTIFAKVDRLAGIINFQRPKD
PNNLLNDWSQKLNSLMSLVNKTTHLIAKEEMIHNLQ
Sequence of entity 5 (X), FASTA
>9BV3_5 26S proteasome non-ATPase regulatory subunit 11 (chains X)
MAAAAVVEFQRAQSLLSTDREASIDILHSIVKRDIQENDEEAVQVKEQSILELGSLLAKT
GQAAELGGLLKYVRPFLNSISKAKAARLVRSLLDLFLDMEAATGQEVELCLECIEWAKSE
KRTFLRQALEARLVSLYFDTKRYQEALHLGSQLLRELKKMDDKALLVEVQLLESKTYHAL
SNLPKARAALTSARTTANAIYCPPKLQATLDMQSGIIHAAEEKDWKTAYSYFYEAFEGYD
SIDSPKAITSLKYMLLCKIMLNTPEDVQALVSGKLALRYAGRQTEALKCVAQASKNRSLA
DFEKALTDYRAELRDDPIISTHLAKLYDNLLEQNLIRVIEPFSRVQIEHISSLIKLSKAD
VERKLSQMILDKKFHGILDQGEGVLIIFDEPPVDKTYEAALETIQNMSKVVDSLYNKAKK
LT
Sequence of entity 6 (Y), FASTA
>9BV3_6 26S proteasome non-ATPase regulatory subunit 6 (chains Y)
MPLENLEEEGLPKNPDLRIAQLRFLLSLPEHRGDAAVRDELMAAVRDNNMAPYYEALCKS
LDWQIDVDLLNKMKKANEDELKRLDEELEDAEKNLGESEIRDAMMAKAEYLCRIGDKEGA
LTAFRKTYDKTVALGHRLDIVFYLLRIGLFYMDNDLITRNTEKAKSLIEEGGDWDRRNRL
KVYQGLYCVAIRDFKQAAELFLDTVSTFTSYELMDYKTFVTYTVYVSMIALERPDLREKV
IKGAEILEVLHSLPAVRQYLFSLYECRYSVFFQSLAVVEQEMKKDWLFAPHYRYYVREMR
IHAYSQLLESYRSLTLGYMAEAFGVGVEFIDQELSRFIAAGRLHCKIDKVNEIVETNRPD
SKNWQYQETIKKGDLLLNRVQKLSRVINM
Sequence of entity 7 (Z), FASTA
>9BV3_7 26S proteasome non-ATPase regulatory subunit 7 (chains Z)
MPELAVQKVVVHPLVLLSVVDHFNRIGKVGNQKRVVGVLLGSWQKKVLDVSNSFAVPFDE
DDKDDSVWFLDHDYLENMYGMFKKVNARERIVGWYHTGPKLHKNDIAINELMKRYCPNSV
LVIIDVKPKDLGLPTEAYISVEEVHDDGTPTSKTFEHVTSEIGAEEAEEVGVEHLLRDIK
DTTVGTLSQRITNQVHGLKGLNSKLLDIRSYLEKVATGKLPINHQIIYQLQDVFNLLPDV
SLQEFVKAFYLKTNDQMVVVYLASLIRSVVALHNLINNKIANRDAEKKEGQEKEESKKDR
KEDKEKDKDKEKSDVKKEEKKEKK
Sequence of entity 8 (a), FASTA
>9BV3_8 26S proteasome non-ATPase regulatory subunit 13 (chains a)
MKDVPGFLQQSQNSGPGQPAVWHRLEELYTKKLWHQLTLQVLDFVQDPCFAQGDGLIKLY
ENFISEFEHRVNPLSLVEIILHVVRQMTDPNVALTFLEKTREKVKSSDEAVILCKTAIGA
LKLNIGDLQVTKETIEDVEEMLNNLPGVTSVHSRFYDLSSKYYQTIGNHASYYKDALRFL
GCVDIKDLPVSEQQERAFTLGLAGLLGEGVFNFGELLMHPVLESLRNTDRQWLIDTLYAF
NSGNVERFQTLKTAWGQQPDLAANEAQLLRKIQLLCLMEMTFTRPANHRQLTFEEIAKSA
KITVNEVELLVMKALSVGLVKGSIDEVDKRVHMTWVQPRVLDLQQIKGMKDRLEFWCTDV
KSMEMLVEHQAHDILT
Sequence of entity 9 (b), FASTA
>9BV3_9 26S proteasome non-ATPase regulatory subunit 4 (chains b)
MVLESTMVCVDNSEYMRNGDFLPTRLQAQQDAVNIVCHSKTRSNPENNVGLITLANDCEV
LTTLTPDTGRILSKLHTVQPKGKITFCTGIRVAHLALKHRQGKNHKMRIIAFVGSPVEDN
EKDLVKLAKRLKKEKVNVDIINFGEEEVNTEKLTAFVNTLNGKDGTGSHLVTVPPGPSLA
DALISSPILAGEGGAMLGLGASDFEFGVDPSADPELALALRVSMEEQRQRQEEEARRAAA
ASAAEAGIATTGTEDSDDALLKMTISQQEFGRTGLPDLSSMTEEEQIAYAMQMSLQGAEF
GQAESADIDASSAMDTSEPAKEEDDYDVMQDPEFLQSVLENLPGVDPNNEAIRNAMGSLA
SQATKDGKKDKKEEDKK
Sequence of entity 10 (c), FASTA
>9BV3_10 26S proteasome non-ATPase regulatory subunit 14 (chains c)
MDRLLRLGGGMPGLGQGPPTDAPAVDTAEQVYISSLALLKMLKHGRAGVPMEVMGLMLGE
FVDDYTVRVIDVFAMPQSGTGVSVEAVDPVFQAKMLDMLKQTGRPEMVVGWYHSHPGFGC
WLSGVDINTQQSFEALSERAVAVVVDPIQSVKGKVVIDAFRLINANMMVLGHEPRQTTSN
LGHLNKPSIQALIHGLNRHYYSITINYRKNELEQKMLLNLHKKSWMEGLTLQDYSEHCKH
NESVVKEMLELAKNYNKAVEEEDKMTPEQLAIKNVGKQDPKRHLEEHVDVLMTSNIVQCL
AAMLDTVVFK
Sequence of entity 11 (d), FASTA
>9BV3_11 26S proteasome non-ATPase regulatory subunit 8 (chains d)
MFIKGRAPRAPPRERRRATRGGLRQVVAPPRALGSTSRPHFRRASVCRRRCRKSGGLLAA
SRKMAAAAVNGAAGFSSSGPAATSGAVLQAATGMYEQLKGEWNRKSPNLSKCGEELGRLK
LVLLELNFLPTTGTKLTKQQLILARDILEIGAQWSILRKDIPSFERYMAQLKCYYFDYKE
QLPESAYMHQLLGLNLLFLLSQNRVAEFHTELERLPAKDIQTNVYIKHPVSLEQYLMEGS
YNKVFLAKGNIPAESYTFFIDILLDTIRDEIAGCIEKAYEKILFTEATRILFFNTPKKMT
DYAKKRGWVLGPNNYYSFASQQQKPEDTTIPSTELAKQVIEYARQLEMIV
Sequence of entity 12 (e), FASTA
>9BV3_12 26S proteasome complex subunit SEM1 (chains e)
MSEKKQPVDLGLLEEDDEFEEFPAEDWAGLDEDEDAHVWEDNWDDDNVEDDFSNQLRAEL
EKHGYKMETS

Ligands and cofactors

IDNameFormulaCopies
ADPAdenosine-5'-diphosphateC10 H15 N5 O10 P22
MGMagnesium ionMg5
ATPAdenosine-5'-triphosphateC10 H16 N5 O13 P34

Primary citation

Structural basis for the midnolin-proteasome pathway and its role in suppressing myeloma. Nardone, C., Gao, J., Seo, H.S. et al. Mol Cell (2025) 85:2597-2609.e11. DOI 10.1016/j.molcel.2025.05.030 · PubMed

Other PDB entries of the same protein (UniProt Q504T8 (AlphaFold model), which also has an AlphaFold model), best resolution first:

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