35ZW: 26S proteasome regulatory subunit 7 homolog

yeast 26S proteasome base assembly intermediate, Hsm3-Rpt1-Rpt2-Rpt3-Rpt4-Rpt5 (base-Hsm3-Nas6 with Rpt4-EQ). Determined by electron microscopy at 3.31 Å resolution. Released 19 Aug 2026.

Method: Electron microscopy
Resolution: 3.31 Å
Organism: Saccharomyces cerevisiae
Chains: 7
Atoms: 15,764
Mol. weight: 330.49 kDa
Ligands: ATP
Released: 19 Aug 2026

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

Secondary structure: helices and β-sheets

35ZW contains 141 α-helices and 56 β-strands across 7 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: 17 helices, 11 β-strands

Element	Residues	Length	Sheet
β-strand	198-201	4	1
α-helix	208-210	3
α-helix	215-225	11
α-helix	227-230	4
α-helix	232-238	7
α-helix	240-243	4
β-strand	245-249	5	1
α-helix	256-266	11
β-strand	270-275	6	1
α-helix	276-279	4
α-helix	288-299	12
β-strand	304-309	6	1
α-helix	312-315	4
β-strand	316	1	2
α-helix	317-318	2
β-strand	327	1	3
α-helix	328-342	15
β-strand	349-355	7	1
α-helix	358-360	3
β-strand	361	1	2
β-strand	373-376	4	1
α-helix	378-380	3
α-helix	382-393	12
β-strand	398	1	4
α-helix	404-409	6
α-helix	416-432	17
β-strand	438	1	4
α-helix	440-450	11

Chain B: 17 helices, 8 β-strands

Element	Residues	Length	Sheet
α-helix	167-170	4
β-strand	171-174	4	5
α-helix	181-183	3
α-helix	188-194	7
α-helix	195-199	5
α-helix	200-203	4
α-helix	205-211	7
α-helix	214-216	3
β-strand	218-223	6	5
α-helix	229-240	12
β-strand	243-248	6	5
α-helix	249-254	6
α-helix	260-273	14
β-strand	277-282	6	5
α-helix	285-291	7
α-helix	301-315	15
β-strand	322-328	7	5
α-helix	331-333	3
β-strand	347-350	4	5
α-helix	355-366	12
β-strand	371	1	6
α-helix	389-406	18
β-strand	411	1	6
α-helix	412	1
α-helix	413-427	15

Chain D: 36 helices, 3 β-strands

Element	Residues	Length	Sheet
α-helix	8-21	14
α-helix	29-42	14
α-helix	52-64	13
α-helix	73-83	11
α-helix	89-95	7
α-helix	98-106	9
α-helix	110-121	12
α-helix	134-142	9
α-helix	150-163	14
α-helix	167-171	5
α-helix	172-176	5
α-helix	178-187	10
α-helix	190-206	17
α-helix	214-217	4
α-helix	219-220	2
α-helix	221-227	7
α-helix	231-250	20
α-helix	254-256	3
α-helix	257-260	4
α-helix	264-272	9
α-helix	278-283	6
α-helix	285-296	12
α-helix	304-309	6
α-helix	310-314	5
α-helix	323-329	7
α-helix	332-346	15
α-helix	354-360	7
α-helix	364-370	7
α-helix	376-381	6
α-helix	384-396	13
α-helix	398-407	10
α-helix	409-416	8
α-helix	427-441	15
α-helix	445-448	4
α-helix	449-451	3
α-helix	452-463	12
β-strand	467	1	3
β-strand	473	1	7
β-strand	475	1	8

Chain G: 20 helices, 10 β-strands

Element	Residues	Length	Sheet
α-helix	167-171	5
α-helix	180-182	3
α-helix	187-193	7
α-helix	194-198	5
α-helix	199-202	4
α-helix	205-210	6
α-helix	213-215	3
β-strand	217-221	5	9
α-helix	228-239	12
β-strand	242-245	4	9
α-helix	248-251	4
β-strand	255	1	8
α-helix	258-272	15
β-strand	276-281	6	9
α-helix	283-285	3
β-strand	288	1	10
α-helix	289-291	3
α-helix	296-313	18
β-strand	322-327	6	9
α-helix	330-332	3
β-strand	333	1	10
α-helix	335-338	4
β-strand	345-348	4	9
α-helix	350-352	3
α-helix	354-366	13
β-strand	370	1	11
α-helix	376-382	7
α-helix	388-404	17
β-strand	410	1	11
α-helix	412-426	15

Chain H: 19 helices, 11 β-strands

Element	Residues	Length	Sheet
α-helix	165-170	6
β-strand	171	1	12
α-helix	175-176	2
α-helix	180-182	3
α-helix	187-193	7
α-helix	194-198	5
α-helix	199-202	4
α-helix	204-209	6
α-helix	212-215	4
β-strand	217-221	5	12
α-helix	228-237	10
β-strand	242-247	6	12
α-helix	248-251	4
β-strand	255	1	7
α-helix	258-272	15
β-strand	276-281	6	12
α-helix	283-285	3
β-strand	288	1	13
α-helix	296-312	17
β-strand	321-327	7	12
β-strand	333	1	13
α-helix	335-338	4
β-strand	347-348	2	12
α-helix	350-352	3
α-helix	354-365	12
β-strand	370	1	14
α-helix	376-382	7
α-helix	388-404	17
β-strand	410	1	14
α-helix	412-424	13

Chain L: 12 helices, 9 β-strands

Element	Residues	Length	Sheet
α-helix	178-188	11
β-strand	209-212	4	15
α-helix	219-229	11
β-strand	233-238	6	15
α-helix	239-242	4
α-helix	249-263	15
β-strand	267-272	6	15
α-helix	274-276	3
β-strand	279	1	16
α-helix	288-302	15
β-strand	312-318	7	15
α-helix	321-323	3
β-strand	324	1	16
β-strand	336-339	4	15
α-helix	340-342	3
α-helix	347-357	11
β-strand	362	1	17
α-helix	369-372	4
α-helix	380-396	17
β-strand	402	1	17
α-helix	404-415	12

Chain O: 20 helices, 4 β-strands

Element	Residues	Length	Sheet
α-helix	5-12	8
α-helix	15-24	10
α-helix	26-29	4
α-helix	39-45	7
α-helix	49-56	8
α-helix	59-61	3
α-helix	64-66	3
α-helix	75-82	8
α-helix	85-92	8
α-helix	97-99	3
β-strand	103	1	18
β-strand	109	1	18
α-helix	110-116	7
α-helix	120-128	9
α-helix	143-150	8
α-helix	153-157	5
α-helix	158-162	5
α-helix	177-183	7
α-helix	187-191	5
α-helix	192-196	5
α-helix	197	1
β-strand	204	1	19
β-strand	210	1	19
α-helix	218-225	8

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
26S proteasome regulatory subunit 7 homolog	A	protein	467	Saccharomyces cerevisiae	P33299 (AlphaFold model)
26S proteasome regulatory subunit 4 homolog	B	protein	437	Saccharomyces cerevisiae	P40327 (AlphaFold model)
DNA mismatch repair protein HSM3	D	protein	480	Saccharomyces cerevisiae	P38348 (AlphaFold model)
26S proteasome subunit RPT4	G	protein	437	Saccharomyces cerevisiae	P53549 (AlphaFold model)
26S proteasome regulatory subunit 6A	H	protein	434	Saccharomyces cerevisiae	P33297
26S proteasome regulatory subunit 6B homolog	L	protein	428	Saccharomyces cerevisiae	P33298
Probable 26S proteasome regulatory subunit p28	O	protein	228	Saccharomyces cerevisiae	P50086

Sequence of entity 1 (A), FASTA

>35ZW_1 26S proteasome regulatory subunit 7 homolog (chains A)
MPPKEDWEKYKAPLEDDDKKPDDDKIVPLTEGDIQVLKSYGAAPYAAKLKQTENDLKDIE
ARIKEKAGVKESDTGLAPSHLWDIMGDRQRLGEEHPLQVARCTKIIKGNGESDETTTDNN
NSGNSNSNSNQQSTDADEDDEDAKYVINLKQIAKFVVGLGERVSPTDIEEGMRVGVDRSK
YNIELPLPPRIDPSVTMMTVEEKPDVTYSDVGGCKDQIEKLREVVELPLLSPERFATLGI
DPPKGILLYGPPGTGKTLCARAVANRTDATFIRVIGSELVQKYVGEGARMVRELFEMART
KKACIIFFDEIDAVGGARFDDGAGGDNEVQRTMLELITQLDGFDPRGNIKVMFATNRPNT
LDPALLRPGRIDRKVEFSLPDLEGRANIFRIHSKSMSVERGIRWELISRLCPNSTGAELR
SVCTEAGMFAIRARRKVATEKDFLKAVDKVISGYKKFSSTSRYMQYN

Sequence of entity 2 (B), FASTA

>35ZW_2 26S proteasome regulatory subunit 4 homolog (chains B)
MGQGVSSGQDKKKKKGSNQKPKYEPPVQSKFGRKKRKGGPATAEKLPNIYPSTRCKLKLL
RMERIKDHLLLEEEFVSNSEILKPFEKKQEEEKKQLEEIRGNPLSIGTLEEIIDDDHAIV
TSPTMPDYYVSILSFVDKELLEPGCSVLLHHKTMSIVGVLQDDADPMVSVMKMDKSPTES
YSDIGGLESQIQEIKESVELPLTHPELYEEMGIKPPKGVILYGAPGTGKTLLAKAVANQT
SATFLRIVGSELIQKYLGDGPRLCRQIFKVAGENAPSIVFIDEIDAIGTKRYDSNSGGER
EIQRTMLELLNQLDGFDDRGDVKVIMATNKIETLDPALIRPGRIDRKILFENPDLSTKKK
ILGIHTSKMNLSEDVNLETLVTTKDDLSGADIQAMCTEAGLLALRERRMQVTAEDFKQAK
ERVMKNKVEENLEGLYL

Sequence of entity 3 (D), FASTA

>35ZW_3 DNA mismatch repair protein HSM3 (chains D)
MSEKETNYVENLLTQLENELNEDNLPEDINTLLRKCSLNLVTVVSLPDMDVKPLLATIKR
FLTSNVSYDSLNYDYLLDVVDKLVPMADFDDVLEVYSAEDLVKALRSEIDPLKVAACRVI
ENSQPKGLFATSNIIDILLDILFDEKVENDKLITAIEKALERLSTDELIRRRLFDNNLPY
LVSVKGRMETVSFVRLIDFLTIEFQFISGPEFKDIIFCFTKEEILKSVEDILVFIELVNY
YTKFLLEIRNQDKYWALRHVKKILPVFAQLFEDTENYPDVRAFSTNCLLQLFAEVSRIEE
DEYSLFKTMDKDSLKIGSEAKLITEWLELINPQYLVKYHKDVVENYFHVSGYSIGMLRNL
SADEECFNAIRNKFSAEIVLRLPYLEQMQVVETLTRYEYTSKFLLNEMPKVMGSLIGDGS
AGAIIDLETVHYRNSALRNLLDKGEEKLSVWYEPLLREYSKAVNGKNYSTGSETKIADCR

Sequence of entity 4 (G), FASTA

>35ZW_4 26S proteasome subunit RPT4 (chains G)
MSEEQDPLLAGLGETSGDNHTQQSHEQQPEQPQETEEHHEEEPSRVDPEQEAHNKALNQF
KRKLLEHRRYDDQLKQRRQNIRDLEKLYDKTENDIKALQSIGQLIGEVMKELSEEKYIVK
ASSGPRYIVGVRNSVDRSKLKKGVRVTLDITTLTIMRILPRETDPLVYNMTSFEQGEITF
DGIGGLTEQIRELREVIELPLKNPEIFQRVGIKPPKGVLLYGPPGTGKTLLAKAVAATIG
ANFIFSPASGIVDKYIGESARIIREMFAYAKEHEPCIIFMDQVDAIGGRRFSEGTSADRE
IQRTLMELLTQMDGFDNLGQTKIIMATNRPDTLDPALLRPGRLDRKVEIPLPNEAGRLEI
FKIHTAKVKKTGEFDFEAAVKMSDGFNGADIRNCATEAGFFAIRDDRDHINPDDLMKAVR
KVAEVKKLEGTIEYQKL

Sequence of entity 5 (H), FASTA

>35ZW_5 26S proteasome regulatory subunit 6A (chains H)
MATLEELDAQTLPGDDELDQEILNLSTQELQTRAKLLDNEIRIFRSELQRLSHENNVMLE
KIKDNKEKIKNNRQLPYLVANVVEVMDMNEIEDKENSESTTQGGNVNLDNTAVGKAAVVK
TSSRQTVFLPMVGLVDPDKLKPNDLVGVNKDSYLILDTLPSEFDSRVKAMEVDEKPTETY
SDVGGLDKQIEELVEAIVLPMKRADKFKDMGIRAPKGALMYGPPGTGKTLLARACAAQTN
ATFLKLAAPQLVQMYIGEGAKLVRDAFALAKEKAPTIIFIDELDAIGTKRFDSEKSGDRE
VQRTMLELLNQLDGFSSDDRVKVLAATNRVDVLDPALLRSGRLDRKIEFPLPSEDSRAQI
LQIHSRKMTTDDDINWQELARSTDEFNGAQLKAVTVEAGMIALRNGQSSVKHEDFVEGIS
EVQARKSKSVSFYA

Sequence of entity 6 (L), FASTA

>35ZW_6 26S proteasome regulatory subunit 6B homolog (chains L)
MEELGIVTPVEKAVEEKPAVKSYASLLAQLNGTVNNNSALSNVNSDIYFKLKKLEKEYEL
LTLQEDYIKDEQRHLKRELKRAQEEVKRIQSVPLVIGQFLEPIDQNTGIVSSTTGMSYVV
RILSTLDRELLKPSMSVALHRHSNALVDILPPDSDSSISVMGENEKPDVTYADVGGLDMQ
KQEIREAVELPLVQADLYEQIGIDPPRGVLLYGPPGTGKTMLVKAVANSTKAAFIRVNGS
EFVHKYLGEGPRMVRDVFRLARENAPSIIFIDEVDSIATKRFDAQTGSDREVQRILIELL
TQMDGFDQSTNVKVIMATNRADTLDPALLRPGRLDRKIEFPSLRDRRERRLIFGTIASKM
SLAPEADLDSLIIRNDSLSGAVIAAIMQEAGLRAVRKNRYVILQSDLEEAYATQVKTDNT
VDKFDFYK

Sequence of entity 7 (O), FASTA

>35ZW_7 Probable 26S proteasome regulatory subunit p28 (chains O)
MSNYPLHQACMENEFFKVQELLHSKPSLLLQKDQDGRIPLHWSVSFQAHEITSFLLSKME
NVNLDDYPDDSGWTPFHIACSVGNLEVVKSLYDRPLKPDLNKITNQGVTCLHLAVGKKWF
EVSQFLIENGASVRIKDKFNQIPLHRAASVGSLKLIELLCGLGKSAVNWQDKQGWTPLFH
ALAEGHGDAAVLLVEKYGAEYDLVDNKGAKAEDVALNEQVKKFFLNNV

Ligands and cofactors

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

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:

35ZR 2.99 Å, yeast 26S proteasome base assembly intermediate, Hsm3-Rpt1-Rpt2 (base-Hsm3-Nas6 with…
9CGC 3.61 Å, Yeast 26S proteasome non-substrate-engaged (S1 state)
3VLF 3.8 Å, Crystal structure of yeast proteasome interacting protein
4A3V 3.8 Å, yeast regulatory particle proteasome assembly chaperone Hsm3 in complex with Rpt1…
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
5MP9 4.1 Å, 26S proteasome in presence of ATP (s1)
6FVT 4.1 Å, 26S proteasome, s1 state
6EF3 4.17 Å, Yeast 26S proteasome bound to ubiquitinated substrate (4D motor state)

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