7W38: 26S protease regulatory subunit 7

Structure of USP14-bound human 26S proteasome in state EA2.0_UBL. Determined by electron microscopy at 3.1 Å resolution. Released 4 May 2022.

Method: Electron microscopy
Resolution: 3.1 Å
Organism: Homo sapiens
Chains: 48
Atoms: 106,178
Mol. weight: 1733.01 kDa
Ligands: ATP, MG, ADP, ZN
Released: 4 May 2022

Explore 7W38 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

7W38 contains 589 α-helices and 536 β-strands across 48 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, 5 β-strands

Element	Residues	Length	Sheet
α-helix	5-10	6
α-helix	18-30	13
α-helix	40-46	7
α-helix	54-66	13
α-helix	74-87	14
α-helix	90-103	14
β-strand	105	1	133
β-strand	109	1	133
α-helix	110-121	12
α-helix	131-137	7
α-helix	152-164	13
α-helix	170-182	13
α-helix	190-205	16
α-helix	223-226	4
α-helix	231-242	12
α-helix	246-251	6
α-helix	265-283	19
β-strand	290-292	3	130
α-helix	293-299	7
α-helix	304-306	3
α-helix	307-317	11
β-strand	320-325	6	130
β-strand	330-336	7	130
α-helix	345-370	26

Chain A: 22 helices, 13 β-strands

Element	Residues	Length	Sheet
α-helix	27-35	9
α-helix	39-63	25
α-helix	72-74	3
α-helix	75-78	4
α-helix	84-90	7
β-strand	97-103	7	1
β-strand	111-114	4	1
β-strand	120-123	4	1
β-strand	139-142	4	1
β-strand	149-152	4	1
α-helix	159-162	4
β-strand	165	1	2
α-helix	174-176	3
α-helix	181-191	11
α-helix	193-196	4
α-helix	200-204	5
β-strand	212-215	4	2
α-helix	222-232	11
β-strand	236-241	6	2
α-helix	242-245	4
α-helix	252-266	15
β-strand	270-275	6	2
α-helix	293-306	14
β-strand	315-321	7	2
α-helix	324-326	3
α-helix	329-332	4
β-strand	339-342	4	2
α-helix	349-359	11
β-strand	364	1	3
α-helix	370-376	7
α-helix	382-398	17
β-strand	404	1	3
α-helix	406-413	8
α-helix	414-418	5
α-helix	419-422	4

Chain b: 8 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	4	1	4
β-strand	5-10	6	5
α-helix	14-17	4
α-helix	25-43	19
β-strand	48-53	6	5
β-strand	65	1	5
α-helix	69-75	7
α-helix	86-99	14
β-strand	106	1	4
β-strand	108-113	6	5
α-helix	121-134	14
β-strand	137-142	6	5
α-helix	153-161	9
β-strand	171	1	5
α-helix	172-174	3
α-helix	181-184	4

Chain B: 20 helices, 17 β-strands

Element	Residues	Length	Sheet
α-helix	59-84	26
α-helix	94-102	9
β-strand	107-112	6	6
β-strand	116	1	7
β-strand	120	1	7
β-strand	123-124	2	6
β-strand	130	1	6
β-strand	134	1	7
β-strand	135	1	6
β-strand	149-152	4	6
β-strand	159-162	4	6
α-helix	163-168	6
α-helix	169-174	6
β-strand	175	1	8
α-helix	184-186	3
α-helix	193-197	5
α-helix	198-202	5
α-helix	203-206	4
α-helix	210-214	5
β-strand	221-225	5	8
α-helix	232-242	11
β-strand	246-251	6	8
α-helix	252-255	4
α-helix	263-277	15
β-strand	280-285	6	8
α-helix	289-292	4
α-helix	302-317	16
β-strand	325-331	7	8
α-helix	339-341	3
β-strand	349-352	4	8
α-helix	354-356	3
α-helix	358-368	11
β-strand	374	1	9
α-helix	380-386	7
α-helix	392-408	17
β-strand	414	1	9
α-helix	416-426	11

Chain c: 11 helices, 9 β-strands

Element	Residues	Length	Sheet
β-strand	30-33	4	115
α-helix	35-47	13
β-strand	54-60	7	115
β-strand	66-74	9	115
β-strand	78-79	2	116
β-strand	84-85	2	117
α-helix	90-101	12
β-strand	107-114	8	115
α-helix	128-136	9
β-strand	142-145	4	115
β-strand	157-162	6	115
α-helix	166-169	4
α-helix	186-188	3
α-helix	189-192	4
β-strand	200-207	8	115
α-helix	211-218	8
α-helix	227-229	3
α-helix	236-261	26
α-helix	267-273	7
α-helix	283-308	26

Chain C: 20 helices, 12 β-strands

Element	Residues	Length	Sheet
α-helix	22-65	44
β-strand	71-79	9	6
β-strand	84-88	5	6
β-strand	94-98	5	6
β-strand	113-116	4	6
β-strand	123-126	4	6
α-helix	127	1
α-helix	129-132	4
α-helix	133-141	9
α-helix	148-150	3
α-helix	155-161	7
α-helix	162-166	5
α-helix	167-170	4
α-helix	172-178	7
α-helix	181-183	3
β-strand	185-189	5	10
α-helix	196-207	12
β-strand	210-213	4	10
α-helix	218-224	7
α-helix	234-240	7
β-strand	244-249	6	10
β-strand	254	1	11
β-strand	257	1	11
α-helix	268-279	12
β-strand	289-295	7	10
α-helix	298-300	3
β-strand	313-316	4	10
α-helix	318-320	3
α-helix	322-332	11
α-helix	344-350	7
α-helix	356-372	17
α-helix	380-395	16

Chain d: 11 helices, 0 β-strands

Element	Residues	Length
α-helix	19-32	14
α-helix	46-64	19
α-helix	69-83	15
α-helix	95-108	14
α-helix	112-118	7
α-helix	131-144	14
α-helix	149-156	8
α-helix	164-181	18
α-helix	193-196	4
α-helix	205-212	8
α-helix	236-254	19

Chain D: 19 helices, 10 β-strands

Element	Residues	Length	Sheet
α-helix	42-76	35
α-helix	79-82	4
β-strand	87-97	11	6
β-strand	100-105	6	6
β-strand	110-114	5	6
α-helix	121-123	3
β-strand	129-133	5	6
β-strand	139-142	4	6
α-helix	171-177	7
α-helix	178-182	5
α-helix	183-186	4
α-helix	188-194	7
β-strand	201-205	5	16
α-helix	212-222	11
β-strand	226-231	6	16
α-helix	232-235	4
α-helix	242-257	16
β-strand	260-265	6	16
α-helix	267-269	3
α-helix	281-297	17
β-strand	305-310	6	16
α-helix	319-322	4
β-strand	330-332	3	16
α-helix	334-336	3
α-helix	339-349	11
α-helix	360-365	6
α-helix	373-388	16
α-helix	396-406	11
α-helix	415-417	3

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

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
26S protease regulatory subunit 7	A	protein	433	Homo sapiens	P35998 (AlphaFold model)
26S protease regulatory subunit 4	B	protein	440	Homo sapiens	P62191 (AlphaFold model)
Isoform 2 of 26S proteasome regulatory subunit 8	C	protein	398	Homo sapiens	P62195 (AlphaFold model)
26S protease regulatory subunit 6B	D	protein	418	Homo sapiens	P43686 (AlphaFold model)
26S proteasome regulatory subunit 10B	E	protein	403	Homo sapiens	A0A087X2I1
Proteasome subunit alpha type-6	G, g	protein	246	Homo sapiens	P60900
Proteasome subunit alpha type-2	H, h	protein	234	Homo sapiens	P25787
Proteasome subunit alpha type-4	I, i	protein	261	Homo sapiens	P25789
Proteasome subunit alpha type-7	J, j	protein	248	Homo sapiens	O14818
Proteasome subunit alpha type-5	K, k	protein	241	Homo sapiens	P28066
Isoform Long of Proteasome subunit alpha type-1	L, l	protein	269	Homo sapiens	P25786
Proteasome subunit alpha type-3	M, m	protein	255	Homo sapiens	P25788

22 more molecules are not listed.

Sequence of entity 1 (A), FASTA

>7W38_1 26S protease regulatory subunit 7 (chains A)
MPDYLGADQRKTKEDEKDDKPIRALDEGDIALLKTYGQSTYSRQIKQVEDDIQQLLKKIN
ELTGIKESDTGLAPPALWDLAADKQTLQSEQPLQVARCTKIINADSEDPKYIINVKQFAK
FVVDLSDQVAPTDIEEGMRVGVDRNKYQIHIPLPPKIDPTVTMMQVEEKPDVTYSDVGGC
KEQIEKLREVVETPLLHPERFVNLGIEPPKGVLLFGPPGTGKTLCARAVANRTDACFIRV
IGSELVQKYVGEGARMVRELFEMARTKKACLIFFDEIDAIGGARFDDGAGGDNEVQRTML
ELINQLDGFDPRGNIKVLMATNRPDTLDPALMRPGRLDRKIEFSLPDLEGRTHIFKIHAR
SMSVERDIRFELLARLCPNSTGAEIRSVCTEAGMFAIRARRKIATEKDFLEAVNKVIKSY
AKFSATPRYMTYN

Sequence of entity 2 (B), FASTA

>7W38_2 26S protease regulatory subunit 4 (chains B)
MGQSQSGGHGPGGGKKDDKDKKKKYEPPVPTRVGKKKKKTKGPDAASKLPLVTPHTQCRL
KLLKLERIKDYLLMEEEFIRNQEQMKPLEEKQEEERSKVDDLRGTPMSVGTLEEIIDDNH
AIVSTSVGSEHYVSILSFVDKDLLEPGCSVLLNHKVHAVIGVLMDDTDPLVTVMKVEKAP
QETYADIGGLDNQIQEIKESVELPLTHPEYYEEMGIKPPKGVILYGPPGTGKTLLAKAVA
NQTSATFLRVVGSELIQKYLGDGPKLVRELFRVAEEHAPSIVFIDEIDAIGTKRYDSNSG
GEREIQRTMLELLNQLDGFDSRGDVKVIMATNRIETLDPALIRPGRIDRKIEFPLPDEKT
KKRIFQIHTSRMTLADDVTLDDLIMAKDDLSGADIKAICTEAGLMALRERRMKVTNEDFK
KSKENVLYKKQEGTPEGLYL

Sequence of entity 3 (C), FASTA

>7W38_3 Isoform 2 of 26S proteasome regulatory subunit 8 (chains C)
MELEEGKAGSGLRQYYLSKIEELQLIVNDKSQNLRRLQAQRNELNAKVRLLREELQLLQE
QGSYVGEVVRAMDKKKVLVKVHPEGKFVVDVDKNIDINDVTPNCRVALRNDSYTLHKILP
NKVDPLVSLMMVEKVPDSTYEMIGGLDKQIKEIKEVIELPVKHPELFEALGIAQPKGVLL
YGPPGTGKTLLARAVAHHTDCTFIRVSGSELVQKFIGEGARMVRELFVMAREHAPSIIFM
DEIDSIGSSRLEGGSGGDSEVQRTMLELLNQLDGFEATKNIKVIMATNRIDILDSALLRP
GRIDRKIEFPPPNEEARLDILKIHSRKMNLTRGINLRKIAELMPGASGAEVKGVCTEAGM
YALRERRVHVTQEDFEMAVAKVMQKDSEKNMSIKKLWK

Sequence of entity 4 (D), FASTA

>7W38_4 26S protease regulatory subunit 6B (chains D)
MEEIGILVEKAQDEIPALSVSRPQTGLSFLGPEPEDLEDLYSRYKKLQQELEFLEVQEEY
IKDEQKNLKKEFLHAQEEVKRIQSIPLVIGQFLEAVDQNTAIVGSTTGSNYYVRILSTID
RELLKPNASVALHKHSNALVDVLPPEADSSIMMLTSDQKPDVMYADIGGMDIQKQEVREA
VELPLTHFELYKQIGIDPPRGVLMYGPPGCGKTMLAKAVAHHTTAAFIRVVGSEFVQKYL
GEGPRMVRDVFRLAKENAPAIIFIDEIDAIATKRFDAQTGADREVQRILLELLNQMDGFD
QNVNVKVIMATNRADTLDPALLRPGRLDRKIEFPLPDRRQKRLIFSTITSKMNLSEEVDL
EDYVARPDKISGADINSICQESGMLAVRENRYIVLAKDFEKAYKTVIKKDEQEHEFYK

Sequence of entity 5 (E), FASTA

>7W38_5 26S proteasome regulatory subunit 10B (chains E)
MAIPGIPYERRLLIMADPRDKALQDYRKKLLEHKEIDGRLKELREQLKELTKQYEKSEND
LKALQSVGQIVGEVLKQLTEEKFIVKATNGPRYVVGCRRQLDKSKLKPGTRVALDMTTLT
IMRYLPREVDPLVYNMSHEDPGNVSYSEIGGLSEQIRELREVIELPLTNPELFQRVGIIP
PKGCLLYGPPGTGKTLLARAVASQLDCNFLKVVSSSIVDKYIGESARLIREMFNYARDHQ
PCIIFMDEIDAIGGRRFSEGTSADREIQRTLMELLNQMDGFDTLHRVKMIMATNRPDTLD
PALLRPGRLDRKIHIDLPNEQARLDILKIHAGPITKHGEIDYEAIVKLSDGFNGADLRNV
CTEAGMFAIRADHDFVVQEDFMKAVRKVADSKKLESKLDYKPV

Sequence of entity 6 (G, g), FASTA

>7W38_6 Proteasome subunit alpha type-6 (chains G, g)
MSRGSSAGFDRHITIFSPEGRLYQVEYAFKAINQGGLTSVAVRGKDCAVIVTQKKVPDKL
LDSSTVTHLFKITENIGCVMTGMTADSRSQVQRARYEAANWKYKYGYEIPVDMLCKRIAD
ISQVYTQNAEMRPLGCCMILIGIDEEQGPQVYKCDPAGYYCGFKATAAGVKQTESTSFLE
KKVKKKFDWTFEQTVETAITCLSTVLSIDFKPSEIEVGVVTVENPKFRILTEAEIDAHLV
ALAERD

Sequence of entity 7 (H, h), FASTA

>7W38_7 Proteasome subunit alpha type-2 (chains H, h)
MAERGYSFSLTTFSPSGKLVQIEYALAAVAGGAPSVGIKAANGVVLATEKKQKSILYDER
SVHKVEPITKHIGLVYSGMGPDYRVLVHRARKLAQQYYLVYQEPIPTAQLVQRVASVMQE
YTQSGGVRPFGVSLLICGWNEGRPYLFQSDPSGAYFAWKATAMGKNYVNGKTFLEKRYNE
DLELEDAIHTAILTLKESFEGQMTEDNIEVGICNEAGFRRLTPTEVKDYLAAIA

Sequence of entity 8 (I, i), FASTA

>7W38_8 Proteasome subunit alpha type-4 (chains I, i)
MSRRYDSRTTIFSPEGRLYQVEYAMEAIGHAGTCLGILANDGVLLAAERRNIHKLLDEVF
FSEKIYKLNEDMACSVAGITSDANVLTNELRLIAQRYLLQYQEPIPCEQLVTALCDIKQA
YTQFGGKRPFGVSLLYIGWDKHYGFQLYQSDPSGNYGGWKATCIGNNSAAAVSMLKQDYK
EGEMTLKSALALAIKVLNKTMDVSKLSAEKVEIATLTRENGKTVIRVLKQKEVEQLIKKH
EEEEAKAEREKKEKEQKEKDK

Sequence of entity 9 (J, j), FASTA

>7W38_9 Proteasome subunit alpha type-7 (chains J, j)
MSYDRAITVFSPDGHLFQVEYAQEAVKKGSTAVGVRGRDIVVLGVEKKSVAKLQDERTVR
KICALDDNVCMAFAGLTADARIVINRARVECQSHRLTVEDPVTVEYITRYIASLKQRYTQ
SNGRRPFGISALIVGFDFDGTPRLYQTDPSGTYHAWKANAIGRGAKSVREFLEKNYTDEA
IETDDLTIKLVIKALLEVVQSGGKNIELAVMRRDQSLKILNPEEIEKYVAEIEKEKEENE
KKKQKKAS

Sequence of entity 10 (K, k), FASTA

>7W38_10 Proteasome subunit alpha type-5 (chains K, k)
MFLTRSEYDRGVNTFSPEGRLFQVEYAIEAIKLGSTAIGIQTSEGVCLAVEKRITSPLME
PSSIEKIVEIDAHIGCAMSGLIADAKTLIDKARVETQNHWFTYNETMTVESVTQAVSNLA
LQFGEEDADPGAMSRPFGVALLFGGVDEKGPQLFHMDPSGTFVQCDARAIGSASEGAQSS
LQEVYHKSMTLKEAIKSSLIILKQVMEEKLNATNIELATVQPGQNFHMFTKEELEEVIKD
I

Sequence of entity 11 (L, l), FASTA

>7W38_11 Isoform Long of Proteasome subunit alpha type-1 (chains L, l)
MQLSKVKFRNQYDNDVTVWSPQGRIHQIEYAMEAVKQGSATVGLKSKTHAVLVALKRAQS
ELAAHQKKILHVDNHIGISIAGLTADARLLCNFMRQECLDSRFVFDRPLPVSRLVSLIGS
KTQIPTQRYGRRPYGVGLLIAGYDDMGPHIFQTCPSANYFDCRAMSIGARSQSARTYLER
HMSEFMECNLNELVKHGLRALRETLPAEQDLTTKNVSIGIVGKDLEFTIYDDDDVSPFLE
GLEERPQRKAQPAQPADEPAEKADEPMEH

Sequence of entity 12 (M, m), FASTA

>7W38_12 Proteasome subunit alpha type-3 (chains M, m)
MSSIGTGYDLSASTFSPDGRVFQVEYAMKAVENSSTAIGIRCKDGVVFGVEKLVLSKLYE
EGSNKRLFNVDRHVGMAVAGLLADARSLADIAREEASNFRSNFGYNIPLKHLADRVAMYV
HAYTLYSAVRPFGCSFMLGSYSVNDGAQLYMIDPSGVSYGYWGCAIGKARQAAKTEIEKL
QMKEMTCRDIVKEVAKIIYIVHDEVKDKAFELELSWVGELTNGRHEIVPKDIREEAEKYA
KESLKEEDESDDDNM

Ligands and cofactors

ID	Name	Formula	Copies
ATP	Adenosine-5'-triphosphate	C10 H16 N5 O13 P3	4
MG	Magnesium ion	Mg	6
ADP	Adenosine-5'-diphosphate	C10 H15 N5 O10 P2	2
ZN	Zinc ion	Zn	1

Primary citation

USP14-regulated allostery of the human proteasome by time-resolved cryo-EM. Zhang, S., Zou, S., Yin, D. et al. Nature (2022) 605:567-574. DOI 10.1038/s41586-022-04671-8 · PubMed

Other PDB entries of the same protein (UniProt P35998 (AlphaFold model), which also has an AlphaFold model), best resolution first:

9K53 2.5 Å, Structure of substrate-engaged human 26S proteasome RP-CP subcomplex in state ED2.1
8USD 2.7 Å, Rpn1/Nub1UBL-focused alignment of the non-substrate-engaged human 26S proteasome
8USB 2.73 Å, Non-substrate-engaged human 26S proteasome with Nub1/FAT10 bound to Rpn1
9MBP 2.75 Å, Substrate-engaged human 26S proteasome bound to midnolin with RPT1 at top of spiral…
9PDL 2.76 Å, Nub1/Fat10-processing human 26S proteasome with Rpt5 at top of spiral staircase (AAA+…
9NKG 2.8 Å, Structure of substrate engaged MIDN-bound human 26S proteasome, EB-MIDN (Composite map)
9E8I 2.87 Å, Human proteasome in resting state conformation bound to TXNL1 in Forward conformation
9BV3 2.9 Å, M1B Midnolin-Proteasome
9E8H 2.9 Å, Human proteasome in resting state conformation bound to TXNL1 in backward conformation
9K4J 2.9 Å, Structure of substrate-engaged human 26S proteasome RP-CP subcomplex in state EA1.0
9NKF 2.9 Å, Structure of human substrate-free 26S proteasome in the presence of ATPgS and…
9U3L 2.91 Å, Substrate-engaged human 26S proteasome bound to midnolin with RPT2 at top of spiral…

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