36AX: 26S proteasome regulatory subunit 7 homolog

yeast 26S proteasome base assembly intermediate, base-Nas2-Rpn14-Hsm3-Nas6 with Rpt4-EQ. Determined by electron microscopy at 4.87 Å resolution. Released 19 Aug 2026.

Method
Electron microscopy
Resolution
4.87 Å
Organism
Saccharomyces cerevisiae
Chains
13
Atoms
43,176
Mol. weight
679.52 kDa
Ligands
ATP
Released
19 Aug 2026

Explore 36AX in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

36AX contains 303 α-helices and 184 β-strands across 13 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: 21 helices, 12 β-strands

ElementResiduesLengthSheet
α-helix42-432
α-helix46-6520
β-strand7611
α-helix79-813
α-helix86-883
β-strand199-20132
α-helix208-2103
α-helix215-23016
α-helix232-2387
α-helix241-2433
β-strand245-24952
α-helix256-26712
β-strand270-27562
α-helix276-2816
α-helix287-30014
β-strand304-30962
α-helix312-3154
β-strand31613
α-helix317-3193
α-helix328-34215
β-strand349-35572
α-helix358-3603
β-strand36113
β-strand373-37642
α-helix382-39413
β-strand39814
α-helix404-4096
α-helix416-43217
β-strand43814
α-helix440-4467
α-helix447-4526
β-strand46315
α-helix464-4652
Chain B: 20 helices, 21 β-strands
ElementResiduesLengthSheet
α-helix56-8126
α-helix90-10011
β-strand104-10636
β-strand108-11477
β-strand117-12157
β-strand12218
β-strand12518
β-strand129-13137
β-strand13216
β-strand13611
α-helix138-1403
β-strand147-15046
β-strand156-15946
α-helix166-1683
β-strand172-17439
α-helix181-1833
α-helix188-1947
α-helix195-1995
α-helix200-2034
α-helix205-2117
α-helix213-2164
β-strand218-22369
α-helix229-24012
β-strand243-24869
α-helix249-2546
α-helix260-27314
β-strand277-28269
α-helix285-2884
α-helix297-31519
β-strand322-32879
α-helix331-3333
β-strand339-340210
β-strand344110
β-strand34615
β-strand347-35049
α-helix355-36713
β-strand371111
α-helix377-3804
α-helix389-40517
β-strand411111
α-helix413-43624
Chain C: 52 helices, 13 β-strands
ElementResiduesLengthSheet
α-helix41-5717
α-helix61-633
α-helix64-7916
α-helix87-915
α-helix93-953
α-helix96-10510
α-helix109-12416
α-helix133-1397
α-helix151-17727
α-helix197-21317
α-helix217-22610
α-helix230-2323
α-helix234-2363
α-helix239-25012
α-helix253-2553
α-helix260-27314
α-helix278-28811
α-helix291-2999
α-helix304-31714
α-helix327-3304
α-helix333-34614
α-helix355-3584
α-helix377-39014
α-helix405-4084
α-helix414-42512
α-helix431-4377
α-helix438-4414
α-helix445-45814
α-helix469-4746
α-helix483-49614
α-helix503-51412
α-helix520-53314
α-helix540-55213
α-helix555-5584
α-helix562-57413
α-helix578-5803
α-helix581-5888
α-helix596-60813
α-helix615-6239
α-helix724-74522
α-helix746-7494
α-helix751-76414
α-helix770-78314
α-helix790-80011
α-helix805-81814
α-helix825-83713
α-helix842-85514
α-helix857-8604
β-strand862-863212
α-helix875-88713
α-helix891-8944
α-helix898-9036
α-helix904-9074
β-strand909-910212
β-strand913113
β-strand915-916214
β-strand922114
β-strand926-932715
β-strand945-952815
β-strand955113
β-strand961-965515
β-strand970-972316
β-strand977115
β-strand981-982214
β-strand983-985316
Chain D: 40 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix3-2018
α-helix29-4113
α-helix52-6312
α-helix73-8614
α-helix89-957
α-helix98-1069
α-helix110-12112
α-helix128-1303
α-helix134-1429
α-helix150-16314
α-helix167-1715
α-helix172-1765
α-helix178-18710
α-helix190-20617
α-helix214-2174
α-helix221-2277
α-helix231-25121
α-helix254-2563
α-helix257-2604
α-helix264-2729
α-helix278-2836
α-helix285-29612
α-helix304-3096
α-helix310-3145
α-helix323-3297
α-helix332-3387
α-helix340-3467
α-helix351-3533
α-helix354-3607
α-helix364-3718
α-helix376-3805
α-helix384-39411
α-helix398-4025
α-helix403-4075
α-helix412-4154
α-helix421-4233
α-helix427-44216
α-helix445-4484
α-helix449-4513
α-helix452-46211
Chain I: 18 helices, 15 β-strands
ElementResiduesLengthSheet
α-helix19-6648
α-helix68-692
β-strand70-7896
β-strand83-8866
β-strand92-9876
β-strand112-11546
β-strand122-12546
α-helix134-1374
β-strand139117
α-helix154-16310
α-helix166-1694
α-helix171-1777
β-strand184-188517
α-helix195-20511
β-strand209-213517
α-helix215-2184
α-helix225-23915
β-strand243-249717
α-helix250-2523
β-strand255118
α-helix256-2583
α-helix263-28119
α-helix284-2863
β-strand288-294717
β-strand300118
β-strand312-315417
α-helix321-33111
β-strand337119
α-helix3381
α-helix343-3497
α-helix355-37117
β-strand377119
α-helix379-40022
Chain J: 23 helices, 13 β-strands
ElementResiduesLengthSheet
α-helix47-9044
β-strand94-104116
β-strand107-11266
β-strand117-12266
β-strand136-14056
β-strand146-14946
α-helix1601
β-strand161120
α-helix1621
α-helix163-1653
α-helix171-1733
α-helix178-1847
α-helix185-1895
α-helix190-1934
α-helix195-2006
β-strand208-212520
α-helix219-22911
β-strand233-238620
α-helix240-2423
α-helix249-26315
β-strand267-272620
α-helix274-2763
α-helix2791
α-helix2811
α-helix287-30418
β-strand312-317620
α-helix326-3294
β-strand336-339420
α-helix340-3423
α-helix346-35712
β-strand362121
α-helix369-3724
α-helix380-39617
β-strand402121
α-helix404-41411
α-helix425-4273
Chain K: 17 helices, 16 β-strands
ElementResiduesLengthSheet
α-helix54-9744
α-helix101-1022
β-strand103-1131122
β-strand116-120522
β-strand126-129422
β-strand131123
α-helix137-1393
β-strand144-148522
β-strand155123
β-strand158122
α-helix165-1717
α-helix187-19711
α-helix205-2084
β-strand217-221524
α-helix228-23912
β-strand242-247624
α-helix248-2514
α-helix258-27114
β-strand276-281624
α-helix283-2853
β-strand288125
α-helix296-31318
β-strand321-327724
α-helix330-3323
β-strand333125
α-helix335-3384
β-strand345-348424
α-helix354-36613
β-strand370126
α-helix376-3827
α-helix388-40417
β-strand410126
α-helix412-42817
Chain L: 1 helix, 11 β-strands
ElementResiduesLengthSheet
β-strand8-11427
β-strand12-17628
β-strand23-27528
β-strand32-35427
β-strand36-37229
β-strand46-48329
β-strand50127
β-strand66-67229
β-strand73-77528
β-strand86-91628
β-strand96-101628
α-helix116-1249

5 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 regulatory subunit 7 homologAprotein467Saccharomyces cerevisiaeP33299 (AlphaFold model)
26S proteasome regulatory subunit 4 homologBprotein437Saccharomyces cerevisiaeP40327 (AlphaFold model)
26S proteasome regulatory subunit RPN1Cprotein993Saccharomyces cerevisiaeP38764 (AlphaFold model)
DNA mismatch repair protein HSM3Dprotein480Saccharomyces cerevisiaeP38348 (AlphaFold model)
26S proteasome regulatory subunit 8 homologIprotein405Saccharomyces cerevisiaeQ01939
26S proteasome regulatory subunit 6B homologJprotein428Saccharomyces cerevisiaeP33298
26S proteasome subunit RPT4Kprotein437Saccharomyces cerevisiaeP53549
26S proteasome regulatory subunit RPN13Lprotein156Saccharomyces cerevisiaeO13563
26S proteasome regulatory subunit RPN14Mprotein417Saccharomyces cerevisiaeP53196
Probable 26S proteasome regulatory subunit p28Nprotein228Saccharomyces cerevisiaeP50086
26S proteasome regulatory subunit 6APprotein434Saccharomyces cerevisiaeP33297
Probable 26S proteasome regulatory subunit p27Rprotein220Saccharomyces cerevisiaeP40555

1 more molecules are not listed.

Sequence of entity 1 (A), FASTA
>36AX_1 26S proteasome regulatory subunit 7 homolog (chains A)
MPPKEDWEKYKAPLEDDDKKPDDDKIVPLTEGDIQVLKSYGAAPYAAKLKQTENDLKDIE
ARIKEKAGVKESDTGLAPSHLWDIMGDRQRLGEEHPLQVARCTKIIKGNGESDETTTDNN
NSGNSNSNSNQQSTDADEDDEDAKYVINLKQIAKFVVGLGERVSPTDIEEGMRVGVDRSK
YNIELPLPPRIDPSVTMMTVEEKPDVTYSDVGGCKDQIEKLREVVELPLLSPERFATLGI
DPPKGILLYGPPGTGKTLCARAVANRTDATFIRVIGSELVQKYVGEGARMVRELFEMART
KKACIIFFDEIDAVGGARFDDGAGGDNEVQRTMLELITQLDGFDPRGNIKVMFATNRPNT
LDPALLRPGRIDRKVEFSLPDLEGRANIFRIHSKSMSVERGIRWELISRLCPNSTGAELR
SVCTEAGMFAIRARRKVATEKDFLKAVDKVISGYKKFSSTSRYMQYN
Sequence of entity 2 (B), FASTA
>36AX_2 26S proteasome regulatory subunit 4 homolog (chains B)
MGQGVSSGQDKKKKKGSNQKPKYEPPVQSKFGRKKRKGGPATAEKLPNIYPSTRCKLKLL
RMERIKDHLLLEEEFVSNSEILKPFEKKQEEEKKQLEEIRGNPLSIGTLEEIIDDDHAIV
TSPTMPDYYVSILSFVDKELLEPGCSVLLHHKTMSIVGVLQDDADPMVSVMKMDKSPTES
YSDIGGLESQIQEIKESVELPLTHPELYEEMGIKPPKGVILYGAPGTGKTLLAKAVANQT
SATFLRIVGSELIQKYLGDGPRLCRQIFKVAGENAPSIVFIDEIDAIGTKRYDSNSGGER
EIQRTMLELLNQLDGFDDRGDVKVIMATNKIETLDPALIRPGRIDRKILFENPDLSTKKK
ILGIHTSKMNLSEDVNLETLVTTKDDLSGADIQAMCTEAGLLALRERRMQVTAEDFKQAK
ERVMKNKVEENLEGLYL
Sequence of entity 3 (C), FASTA
>36AX_3 26S proteasome regulatory subunit RPN1 (chains C)
MVDESDKKQQTIDEQSQISPEKQTPNKKDKKKEEEEQLSEEDAKLKTDLELLVERLKEDD
SSLYEASLNALKESIKNSTSSMTAVPKPLKFLRPTYPDLCSIYDKWTDPNLKSSLADVLS
ILAMTYSENGKHDSLRYRLLSDVSDFEGWGHEYIRHLALEIGEVYNDQVEKDAEDETSSD
GSKSDGSAATSGFEFSKEDTLRLCLDIVPYFLKHNGEEDAVDLLLEIESIDKLPQFVDEN
TFQRVCQYMVACVPLLPPPEDVAFLKTAYSIYLSQNELTDAIALAVRLGEEDMIRSVFDA
TSDPVMHKQLAYILAAQKTSFEYEGVQDIIGNGKLSEHFLYLAKELNLTGPKVPEDIYKS
HLDNSKSVFSSAGLDSAQQNLASSFVNGFLNLGYCNDKLIVDNDNWVYKTKGDGMTSAVA
SIGSIYQWNLDGLQQLDKYLYVDEPEVKAGALLGIGISASGVHDGEVEPALLLLQDYVTN
PDTKISSAAILGLGIAFAGSKNDEVLGLLLPIAASTDLPIETAAMASLALAHVFVGTCNG
DITTSIMDNFLERTAIELKTDWVRFLALALGILYMGQGEQVDDVLETISAIEHPMTSAIE
VLVGSCAYTGTGDVLLIQDLLHRLTPKNVKGEEDADEEETAEGQTNSISDFLGEQVNEPT
KNEEAEIEVDEMEVDAEGEEVEVKAEITEKKNGESLEGEEIKSEEKKGKSSDKDATTDGK
NDDEEEEKEAGIVDELAYAVLGIALIALGEDIGKEMSLRHFGHLMHYGNEHIRRMVPLAM
GIVSVSDPQMKVFDTLTRFSHDADLEVSMNSIFAMGLCGAGTNNARLAQLLRQLASYYSR
EQDALFITRLAQGLLHLGKGTMTMDVFNDAHVLNKVTLASILTTAVGLVSPSFMLKHHQL
FYMLNAGIRPKFILALNDEGEPIKVNVRVGQAVETVGQAGRPKKITGWITQSTPVLLNHG
ERAELETDEYISYTSHIEGVVILKKNPDYREEE
Sequence of entity 4 (D), FASTA
>36AX_4 DNA mismatch repair protein HSM3 (chains D)
MSEKETNYVENLLTQLENELNEDNLPEDINTLLRKCSLNLVTVVSLPDMDVKPLLATIKR
FLTSNVSYDSLNYDYLLDVVDKLVPMADFDDVLEVYSAEDLVKALRSEIDPLKVAACRVI
ENSQPKGLFATSNIIDILLDILFDEKVENDKLITAIEKALERLSTDELIRRRLFDNNLPY
LVSVKGRMETVSFVRLIDFLTIEFQFISGPEFKDIIFCFTKEEILKSVEDILVFIELVNY
YTKFLLEIRNQDKYWALRHVKKILPVFAQLFEDTENYPDVRAFSTNCLLQLFAEVSRIEE
DEYSLFKTMDKDSLKIGSEAKLITEWLELINPQYLVKYHKDVVENYFHVSGYSIGMLRNL
SADEECFNAIRNKFSAEIVLRLPYLEQMQVVETLTRYEYTSKFLLNEMPKVMGSLIGDGS
AGAIIDLETVHYRNSALRNLLDKGEEKLSVWYEPLLREYSKAVNGKNYSTGSETKIADCR
Sequence of entity 5 (I), FASTA
>36AX_5 26S proteasome regulatory subunit 8 homolog (chains I)
MTAAVTSSNIVLETHESGIKPYFEQKIQETELKIRSKTENVRRLEAQRNALNDKVRFIKD
ELRLLQEPGSYVGEVIKIVSDKKVLVKVQPEGKYIVDVAKDINVKDLKASQRVCLRSDSY
MLHKVLENKADPLVSLMMVEKVPDSTYDMVGGLTKQIKEIKEVIELPVKHPELFESLGIA
QPKGVILYGPPGTGKTLLARAVAHHTDCKFIRVSGAELVQKYIGEGSRMVRELFVMAREH
APSIIFMDEIDSIGSTRVEGSGGGDSEVQRTMLELLNQLDGFETSKNIKIIMATNRLDIL
DPALLRPGRIDRKIEFPPPSVAARAEILRIHSRKMNLTRGINLRKVAEKMNGCSGADVKG
VCTEAGMYALRERRIHVTQEDFELAVGKVMNKNQETAISVAKLFK
Sequence of entity 6 (J), FASTA
>36AX_6 26S proteasome regulatory subunit 6B homolog (chains J)
MEELGIVTPVEKAVEEKPAVKSYASLLAQLNGTVNNNSALSNVNSDIYFKLKKLEKEYEL
LTLQEDYIKDEQRHLKRELKRAQEEVKRIQSVPLVIGQFLEPIDQNTGIVSSTTGMSYVV
RILSTLDRELLKPSMSVALHRHSNALVDILPPDSDSSISVMGENEKPDVTYADVGGLDMQ
KQEIREAVELPLVQADLYEQIGIDPPRGVLLYGPPGTGKTMLVKAVANSTKAAFIRVNGS
EFVHKYLGEGPRMVRDVFRLARENAPSIIFIDEVDSIATKRFDAQTGSDREVQRILIELL
TQMDGFDQSTNVKVIMATNRADTLDPALLRPGRLDRKIEFPSLRDRRERRLIFGTIASKM
SLAPEADLDSLIIRNDSLSGAVIAAIMQEAGLRAVRKNRYVILQSDLEEAYATQVKTDNT
VDKFDFYK
Sequence of entity 7 (K), FASTA
>36AX_7 26S proteasome subunit RPT4 (chains K)
MSEEQDPLLAGLGETSGDNHTQQSHEQQPEQPQETEEHHEEEPSRVDPEQEAHNKALNQF
KRKLLEHRRYDDQLKQRRQNIRDLEKLYDKTENDIKALQSIGQLIGEVMKELSEEKYIVK
ASSGPRYIVGVRNSVDRSKLKKGVRVTLDITTLTIMRILPRETDPLVYNMTSFEQGEITF
DGIGGLTEQIRELREVIELPLKNPEIFQRVGIKPPKGVLLYGPPGTGKTLLAKAVAATIG
ANFIFSPASGIVDKYIGESARIIREMFAYAKEHEPCIIFMDQVDAIGGRRFSEGTSADRE
IQRTLMELLTQMDGFDNLGQTKIIMATNRPDTLDPALLRPGRLDRKVEIPLPNEAGRLEI
FKIHTAKVKKTGEFDFEAAVKMSDGFNGADIRNCATEAGFFAIRDDRDHINPDDLMKAVR
KVAEVKKLEGTIEYQKL
Sequence of entity 8 (L), FASTA
>36AX_8 26S proteasome regulatory subunit RPN13 (chains L)
MSMSSTVIKFRAGVCEYNEDSRLCTPIPVQGEIEIKPNEEEELGFWDFEWRPTEKPVGRE
LDPISLILIPGETMWVPIKSSKSGRIFALVFSSNERYFFWLQEKNSGNLPLNELSAKDKE
IYNKMIGVLNNSSESDEEESNDEKQKAQDVDVSMQD
Sequence of entity 9 (M), FASTA
>36AX_9 26S proteasome regulatory subunit RPN14 (chains M)
MTKTITVAHIQYDFKAVLEENDENDDEFYINVDKNLNEIKEHKIVVLGNSRGVDAGKGNT
FEKVGSHLYKARLDGHDFLFNTIIRDGSKMLKRADYTAVDTAKLQMRRFILGTTEGDIKV
LDSNFNLQREIDQAHVSEITKLKFFPSGEALISSSQDMQLKIWSVKDGSNPRTLIGHRAT
VTDIAIIDRGRNVLSASLDGTIRLWECGTGTTIHTFNRKENPHDGVNSIALFVGTDRQLH
EISTSKKNNLEFGTYGKYVIAGHVSGVITVHNVFSKEQTIQLPSKFTCSCNSLTVDGNNA
NYIYAGYENGMLAQWDLRSPECPVGEFLINEGTPINNVYFAAGALFVSSGFDTSIKLDII
SDPESERPAIEFETPTFLVSNDDEVSQFCYVSDDESNGEVLEVGKNNFCALYNLSNP
Sequence of entity 10 (N), FASTA
>36AX_10 Probable 26S proteasome regulatory subunit p28 (chains N)
MSNYPLHQACMENEFFKVQELLHSKPSLLLQKDQDGRIPLHWSVSFQAHEITSFLLSKME
NVNLDDYPDDSGWTPFHIACSVGNLEVVKSLYDRPLKPDLNKITNQGVTCLHLAVGKKWF
EVSQFLIENGASVRIKDKFNQIPLHRAASVGSLKLIELLCGLGKSAVNWQDKQGWTPLFH
ALAEGHGDAAVLLVEKYGAEYDLVDNKGAKAEDVALNEQVKKFFLNNV
Sequence of entity 11 (P), FASTA
>36AX_11 26S proteasome regulatory subunit 6A (chains P)
MATLEELDAQTLPGDDELDQEILNLSTQELQTRAKLLDNEIRIFRSELQRLSHENNVMLE
KIKDNKEKIKNNRQLPYLVANVVEVMDMNEIEDKENSESTTQGGNVNLDNTAVGKAAVVK
TSSRQTVFLPMVGLVDPDKLKPNDLVGVNKDSYLILDTLPSEFDSRVKAMEVDEKPTETY
SDVGGLDKQIEELVEAIVLPMKRADKFKDMGIRAPKGALMYGPPGTGKTLLARACAAQTN
ATFLKLAAPQLVQMYIGEGAKLVRDAFALAKEKAPTIIFIDELDAIGTKRFDSEKSGDRE
VQRTMLELLNQLDGFSSDDRVKVLAATNRVDVLDPALLRSGRLDRKIEFPLPSEDSRAQI
LQIHSRKMTTDDDINWQELARSTDEFNGAQLKAVTVEAGMIALRNGQSSVKHEDFVEGIS
EVQARKSKSVSFYA
Sequence of entity 12 (R), FASTA
>36AX_12 Probable 26S proteasome regulatory subunit p27 (chains R)
MEEEELSKLLANVKIDPSLTSRISQIDSFKLSELMVLKTDIETQLEAYFSVLEQQGIGMD
SALVTPDGYPRSDVDVLQVTMIRKNVNMLKNDLNHLLQRSHVLLNQHFDNMNVKSNQDAR
RNNDDQAIQYTIPFAFISEVVPGSPSDKADIKVDDKLISIGNVHAANHSKLQNIQMVVMK
NEDRPLPVLLLREGQILKTSLTPSRNWNGRGLLGCRIQEL

Ligands and cofactors

IDNameFormulaCopies
ATPAdenosine-5'-triphosphateC10 H16 N5 O13 P36

Primary citation

Chaperones shape the conformational landscape of 26S-proteasome-base assembly for allosteric ATPase motor activation. Hsieh, H.H., Martin, A. bioRxiv (2026). DOI 10.64898/2026.06.01.729410 · PubMed

Other PDB entries of the same protein (UniProt P33299 (AlphaFold model), which also has an AlphaFold model), best resolution first:

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