35ZV: 26S proteasome regulatory subunit RPN1

yeast 26S proteasome base assembly intermediate, Rpn14-Rpt6 (base-Rpn14-Hsm3-Nas6(2) with Rpt4-EQ). Determined by electron microscopy at 3.11 Å resolution. Released 19 Aug 2026.

Method: Electron microscopy
Resolution: 3.11 Å
Organism: Saccharomyces cerevisiae
Chains: 4
Atoms: 6,397
Mol. weight: 249.84 kDa
Ligands: ATP
Released: 19 Aug 2026

Explore 35ZV in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

35ZV contains 34 α-helices and 48 β-strands across 3 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 C: 13 helices, 0 β-strands

Element	Residues	Length
α-helix	41-57	17
α-helix	61-63	3
α-helix	64-76	13
α-helix	88-92	5
α-helix	96-105	10
α-helix	109-125	17
α-helix	133-140	8
α-helix	147-149	3
α-helix	151-169	19
α-helix	197-213	17
α-helix	217-226	10
α-helix	230-232	3
α-helix	234-236	3

Chain I: 17 helices, 10 β-strands

Element	Residues	Length	Sheet
β-strand	139-140	2	1
α-helix	147-149	3
α-helix	154-160	7
α-helix	161-165	5
α-helix	166-169	4
α-helix	172-177	6
α-helix	179-182	4
β-strand	184-188	5	1
α-helix	195-206	12
β-strand	209-214	6	1
α-helix	216-218	3
α-helix	226-239	14
β-strand	243-248	6	1
α-helix	267-281	15
β-strand	288-294	7	1
α-helix	302-305	4
β-strand	307	1	2
β-strand	309	1	2
β-strand	312-315	4	1
α-helix	317-319	3
α-helix	321-331	11
β-strand	337	1	3
α-helix	338	1
α-helix	343-349	7
α-helix	355-371	17
β-strand	377	1	3
α-helix	379-394	16

Chain M: 4 helices, 38 β-strands

Element	Residues	Length	Sheet
β-strand	3-5	3	4
α-helix	6	1
β-strand	8-10	3	5
α-helix	14-19	6
β-strand	26-33	8	5
β-strand	39-47	9	5
α-helix	48-50	3
β-strand	52-54	3	5
β-strand	60-63	4	6
β-strand	68-73	6	6
β-strand	76-81	6	6
β-strand	85-86	2	7
α-helix	87-90	4
β-strand	96-102	7	8
β-strand	108-113	6	8
β-strand	118-121	4	8
β-strand	127-131	5	8
β-strand	139-144	6	9
β-strand	150-155	6	9
β-strand	159-164	6	9
β-strand	172-175	4	9
β-strand	181-187	7	10
β-strand	192-197	6	10
β-strand	201-206	6	10
β-strand	211-217	7	10
β-strand	226-232	7	11
β-strand	258-263	6	11
β-strand	268-272	5	11
β-strand	277-282	6	11
β-strand	290-295	6	4
β-strand	302-307	6	4
β-strand	311-316	6	4
β-strand	325-329	5	4
β-strand	335-341	7	6
β-strand	344-349	6	6
β-strand	353-358	6	6
β-strand	359-360	2	12
β-strand	369-370	2	12
β-strand	376-378	3	6
β-strand	385-392	8	7
β-strand	398-404	7	7
β-strand	408-413	6	7

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
26S proteasome regulatory subunit RPN1	C	protein	993	Saccharomyces cerevisiae	P38764 (AlphaFold model)
26S proteasome regulatory subunit 8 homolog	I	protein	405	Saccharomyces cerevisiae	Q01939 (AlphaFold model)
26S proteasome regulatory subunit 6B homolog	J	protein	428	Saccharomyces cerevisiae	P33298 (AlphaFold model)
26S proteasome regulatory subunit RPN14	M	protein	417	Saccharomyces cerevisiae	P53196 (AlphaFold model)

Sequence of entity 1 (C), FASTA

>35ZV_1 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 2 (I), FASTA

>35ZV_2 26S proteasome regulatory subunit 8 homolog (chains I)
MTAAVTSSNIVLETHESGIKPYFEQKIQETELKIRSKTENVRRLEAQRNALNDKVRFIKD
ELRLLQEPGSYVGEVIKIVSDKKVLVKVQPEGKYIVDVAKDINVKDLKASQRVCLRSDSY
MLHKVLENKADPLVSLMMVEKVPDSTYDMVGGLTKQIKEIKEVIELPVKHPELFESLGIA
QPKGVILYGPPGTGKTLLARAVAHHTDCKFIRVSGAELVQKYIGEGSRMVRELFVMAREH
APSIIFMDEIDSIGSTRVEGSGGGDSEVQRTMLELLNQLDGFETSKNIKIIMATNRLDIL
DPALLRPGRIDRKIEFPPPSVAARAEILRIHSRKMNLTRGINLRKVAEKMNGCSGADVKG
VCTEAGMYALRERRIHVTQEDFELAVGKVMNKNQETAISVAKLFK

Sequence of entity 3 (J), FASTA

>35ZV_3 26S proteasome regulatory subunit 6B homolog (chains J)
MEELGIVTPVEKAVEEKPAVKSYASLLAQLNGTVNNNSALSNVNSDIYFKLKKLEKEYEL
LTLQEDYIKDEQRHLKRELKRAQEEVKRIQSVPLVIGQFLEPIDQNTGIVSSTTGMSYVV
RILSTLDRELLKPSMSVALHRHSNALVDILPPDSDSSISVMGENEKPDVTYADVGGLDMQ
KQEIREAVELPLVQADLYEQIGIDPPRGVLLYGPPGTGKTMLVKAVANSTKAAFIRVNGS
EFVHKYLGEGPRMVRDVFRLARENAPSIIFIDEVDSIATKRFDAQTGSDREVQRILIELL
TQMDGFDQSTNVKVIMATNRADTLDPALLRPGRLDRKIEFPSLRDRRERRLIFGTIASKM
SLAPEADLDSLIIRNDSLSGAVIAAIMQEAGLRAVRKNRYVILQSDLEEAYATQVKTDNT
VDKFDFYK

Sequence of entity 4 (M), FASTA

>35ZV_4 26S proteasome regulatory subunit RPN14 (chains M)
MTKTITVAHIQYDFKAVLEENDENDDEFYINVDKNLNEIKEHKIVVLGNSRGVDAGKGNT
FEKVGSHLYKARLDGHDFLFNTIIRDGSKMLKRADYTAVDTAKLQMRRFILGTTEGDIKV
LDSNFNLQREIDQAHVSEITKLKFFPSGEALISSSQDMQLKIWSVKDGSNPRTLIGHRAT
VTDIAIIDRGRNVLSASLDGTIRLWECGTGTTIHTFNRKENPHDGVNSIALFVGTDRQLH
EISTSKKNNLEFGTYGKYVIAGHVSGVITVHNVFSKEQTIQLPSKFTCSCNSLTVDGNNA
NYIYAGYENGMLAQWDLRSPECPVGEFLINEGTPINNVYFAAGALFVSSGFDTSIKLDII
SDPESERPAIEFETPTFLVSNDDEVSQFCYVSDDESNGEVLEVGKNNFCALYNLSNP

Ligands and cofactors

ID	Name	Formula	Copies
ATP	Adenosine-5'-triphosphate	C10 H16 N5 O13 P3	1

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 P38764 (AlphaFold model), which also has an AlphaFold model), best resolution first:

9CGC 3.61 Å, Yeast 26S proteasome non-substrate-engaged (S1 state)
6J2Q 3.8 Å, Yeast proteasome in Ub-accepted state (C1-b)
6J2X 3.8 Å, Yeast proteasome in resting state (C1-a)
36BM 3.82 Å, yeast 26S proteasome base assembly intermediate, base-Hsm3-Nas6 with Rpt4-EQ
36BL 3.91 Å, yeast 26S proteasome base assembly intermediate, base-Rpn14-Hsm3-Nas6(2) with Rpt4-EQ
36BD 3.93 Å, yeast 26S proteasome base assembly intermediate, base-Rpn14-Hsm3-Nas6 with Rpt4-EQ
5MPD 4.1 Å, 26S proteasome in presence of ATP (s1)
6FVT 4.1 Å, 26S proteasome, s1 state
5WVK 4.2 Å, Yeast proteasome-ADP-AlFx
5MPE 4.5 Å, 26S proteasome in presence of ATP (s2)
6FVU 4.5 Å, 26S proteasome, s2 state
6FVW 4.5 Å, 26S proteasome, s4 state

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