7W3C: 26S protease regulatory subunit 4

Structure of USP14-bound human 26S proteasome in substrate-engaged state ED0_USP14. Determined by electron microscopy at 3.4 Å resolution. Released 4 May 2022.

Method: Electron microscopy
Resolution: 3.4 Å
Organisms: Homo sapiens, Saccharomyces cerevisiae
Chains: 49
Atoms: 110,747
Mol. weight: 1735.44 kDa
Ligands: ZN, ADP, ATP
Released: 4 May 2022

Explore 7W3C in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

7W3C contains 591 α-helices and 580 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: 20 helices, 5 β-strands

Element	Residues	Length	Sheet
α-helix	5-10	6
α-helix	18-30	13
α-helix	35-45	11
α-helix	54-68	15
α-helix	73-87	15
α-helix	90-103	14
α-helix	110-122	13
α-helix	131-140	10
α-helix	150-162	13
α-helix	170-182	13
α-helix	190-206	17
α-helix	220-226	7
α-helix	231-242	12
α-helix	247-254	8
α-helix	262-264	3
α-helix	265-282	18
β-strand	291-292	2	149
α-helix	293-299	7
α-helix	304-317	14
β-strand	320	1	150
β-strand	323-325	3	149
β-strand	330-332	3	149
β-strand	335	1	150
α-helix	344-370	27
α-helix	372-375	4

Chain A: 22 helices, 17 β-strands

Element	Residues	Length	Sheet
α-helix	27-35	9
α-helix	42-61	20
α-helix	72-74	3
α-helix	75-78	4
α-helix	81-85	5
α-helix	86-90	5
β-strand	97-99	3	33
β-strand	100-103	4	34
β-strand	111-114	4	34
β-strand	120-123	4	34
β-strand	125	1	33
β-strand	140-142	3	33
β-strand	149-152	4	33
α-helix	161-166	6
β-strand	168	1	35
α-helix	174-177	4
α-helix	181-196	16
α-helix	199-204	6
α-helix	206-209	4
β-strand	211-215	5	36
α-helix	222-231	10
β-strand	235	1	35
β-strand	237-241	5	36
α-helix	242-245	4
α-helix	250-263	14
β-strand	271-275	5	36
α-helix	279-283	5
α-helix	292-304	13
α-helix	305-309	5
β-strand	316-321	6	36
α-helix	324-326	3
β-strand	337-342	6	36
α-helix	349-360	12
β-strand	364	1	37
α-helix	370-376	7
α-helix	382-396	15
β-strand	404	1	37
α-helix	406-417	12
β-strand	432	1	38

Chain b: 7 helices, 10 β-strands

Element	Residues	Length	Sheet
β-strand	4-10	7	151
β-strand	13	1	152
α-helix	14-16	3
α-helix	25-43	19
β-strand	48-53	6	151
β-strand	55	1	153
β-strand	65	1	151
α-helix	69-75	7
β-strand	82	1	152
β-strand	84	1	153
α-helix	86-98	13
β-strand	106-113	8	151
α-helix	121-133	13
β-strand	137-142	6	151
α-helix	151-160	10
β-strand	171	1	151
α-helix	180-189	10

Chain B: 18 helices, 15 β-strands

Element	Residues	Length	Sheet
α-helix	36-38	3
α-helix	59-84	26
α-helix	94-102	9
β-strand	107-114	8	1
β-strand	120-124	5	1
β-strand	130-135	6	1
β-strand	149-152	4	1
β-strand	159-162	4	1
α-helix	169-173	5
β-strand	175	1	2
α-helix	184-186	3
α-helix	191-197	7
α-helix	198-202	5
α-helix	203-206	4
α-helix	210-214	5
α-helix	217-219	3
β-strand	221-225	5	2
α-helix	232-241	10
β-strand	247-251	5	2
α-helix	262-276	15
β-strand	281-285	5	2
α-helix	287-289	3
α-helix	300-316	17
β-strand	326-331	6	2
β-strand	349-352	4	2
α-helix	358-368	11
β-strand	374	1	3
α-helix	380-386	7
α-helix	392-408	17
β-strand	411	1	4
β-strand	414	1	3
α-helix	416-425	10
β-strand	439	1	5

Chain c: 12 helices, 10 β-strands

Element	Residues	Length	Sheet
β-strand	30-33	4	6
α-helix	35-47	13
β-strand	54-63	10	6
β-strand	66-74	9	6
α-helix	76-78	3
α-helix	89-101	13
α-helix	105-107	3
β-strand	108	1	6
β-strand	110-113	4	6
α-helix	124-136	13
β-strand	141-145	5	6
β-strand	157-162	6	6
α-helix	163-164	2
α-helix	189-194	6
β-strand	196	1	6
β-strand	200-202	3	6
β-strand	204-207	4	6
α-helix	214-221	8
α-helix	227-229	3
α-helix	237-261	25
α-helix	268-273	6
α-helix	286-308	23

Chain C: 14 helices, 14 β-strands

Element	Residues	Length	Sheet
α-helix	14-66	53
β-strand	71-80	10	1
β-strand	84-88	5	1
β-strand	95-99	5	1
β-strand	112-116	5	1
β-strand	122-127	6	1
β-strand	139	1	7
α-helix	155-170	16
α-helix	173-178	6
α-helix	181-183	3
β-strand	185-189	5	7
α-helix	196-205	10
β-strand	210-215	6	7
α-helix	226-240	15
β-strand	244-249	6	7
α-helix	269-281	13
β-strand	289-294	6	7
α-helix	298-300	3
β-strand	313-316	4	7
α-helix	318-320	3
α-helix	322-332	11
β-strand	338	1	8
α-helix	344-349	6
α-helix	357-372	16
β-strand	378	1	8
α-helix	380-390	11
α-helix	402-404	3
β-strand	405	1	9

Chain d: 11 helices, 2 β-strands

Element	Residues	Length	Sheet
α-helix	16-32	17
α-helix	42-65	24
α-helix	68-82	15
α-helix	94-108	15
α-helix	112-119	8
α-helix	131-145	15
α-helix	148-154	7
α-helix	164-182	19
β-strand	190	1	10
α-helix	191-198	8
α-helix	204-212	9
β-strand	221	1	10
α-helix	239-253	15

Chain D: 16 helices, 12 β-strands

Element	Residues	Length	Sheet
α-helix	40-80	41
α-helix	81-83	3
β-strand	88-97	10	1
β-strand	100-105	6	1
β-strand	111-115	5	1
α-helix	121-123	3
β-strand	129-132	4	1
β-strand	139-142	4	1
α-helix	148-150	3
α-helix	164-166	3
α-helix	172-178	7
α-helix	183-186	4
α-helix	188-192	5
β-strand	202-205	4	11
α-helix	212-222	11
β-strand	227-230	4	11
α-helix	243-256	14
β-strand	260-266	7	11
β-strand	272	1	12
α-helix	283-297	15
β-strand	305-311	7	11
β-strand	317	1	12
α-helix	319-322	4
β-strand	329-332	4	11
α-helix	341-348	8
α-helix	360-365	6
α-helix	372-388	17
α-helix	398-407	10

40 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 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 (AlphaFold model)
26S protease regulatory subunit 6A	F	protein	439	Homo sapiens	P17980
26S protease regulatory subunit 7	A	protein	433	Homo sapiens	P35998
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

23 more molecules are not listed.

Sequence of entity 1 (B), FASTA

>7W3C_1 26S protease regulatory subunit 4 (chains B)
MGQSQSGGHGPGGGKKDDKDKKKKYEPPVPTRVGKKKKKTKGPDAASKLPLVTPHTQCRL
KLLKLERIKDYLLMEEEFIRNQEQMKPLEEKQEEERSKVDDLRGTPMSVGTLEEIIDDNH
AIVSTSVGSEHYVSILSFVDKDLLEPGCSVLLNHKVHAVIGVLMDDTDPLVTVMKVEKAP
QETYADIGGLDNQIQEIKESVELPLTHPEYYEEMGIKPPKGVILYGPPGTGKTLLAKAVA
NQTSATFLRVVGSELIQKYLGDGPKLVRELFRVAEEHAPSIVFIDEIDAIGTKRYDSNSG
GEREIQRTMLELLNQLDGFDSRGDVKVIMATNRIETLDPALIRPGRIDRKIEFPLPDEKT
KKRIFQIHTSRMTLADDVTLDDLIMAKDDLSGADIKAICTEAGLMALRERRMKVTNEDFK
KSKENVLYKKQEGTPEGLYL

Sequence of entity 2 (C), FASTA

>7W3C_2 Isoform 2 of 26S proteasome regulatory subunit 8 (chains C)
MELEEGKAGSGLRQYYLSKIEELQLIVNDKSQNLRRLQAQRNELNAKVRLLREELQLLQE
QGSYVGEVVRAMDKKKVLVKVHPEGKFVVDVDKNIDINDVTPNCRVALRNDSYTLHKILP
NKVDPLVSLMMVEKVPDSTYEMIGGLDKQIKEIKEVIELPVKHPELFEALGIAQPKGVLL
YGPPGTGKTLLARAVAHHTDCTFIRVSGSELVQKFIGEGARMVRELFVMAREHAPSIIFM
DEIDSIGSSRLEGGSGGDSEVQRTMLELLNQLDGFEATKNIKVIMATNRIDILDSALLRP
GRIDRKIEFPPPNEEARLDILKIHSRKMNLTRGINLRKIAELMPGASGAEVKGVCTEAGM
YALRERRVHVTQEDFEMAVAKVMQKDSEKNMSIKKLWK

Sequence of entity 3 (D), FASTA

>7W3C_3 26S protease regulatory subunit 6B (chains D)
MEEIGILVEKAQDEIPALSVSRPQTGLSFLGPEPEDLEDLYSRYKKLQQELEFLEVQEEY
IKDEQKNLKKEFLHAQEEVKRIQSIPLVIGQFLEAVDQNTAIVGSTTGSNYYVRILSTID
RELLKPNASVALHKHSNALVDVLPPEADSSIMMLTSDQKPDVMYADIGGMDIQKQEVREA
VELPLTHFELYKQIGIDPPRGVLMYGPPGCGKTMLAKAVAHHTTAAFIRVVGSEFVQKYL
GEGPRMVRDVFRLAKENAPAIIFIDEIDAIATKRFDAQTGADREVQRILLELLNQMDGFD
QNVNVKVIMATNRADTLDPALLRPGRLDRKIEFPLPDRRQKRLIFSTITSKMNLSEEVDL
EDYVARPDKISGADINSICQESGMLAVRENRYIVLAKDFEKAYKTVIKKDEQEHEFYK

Sequence of entity 4 (E), FASTA

>7W3C_4 26S proteasome regulatory subunit 10B (chains E)
MAIPGIPYERRLLIMADPRDKALQDYRKKLLEHKEIDGRLKELREQLKELTKQYEKSEND
LKALQSVGQIVGEVLKQLTEEKFIVKATNGPRYVVGCRRQLDKSKLKPGTRVALDMTTLT
IMRYLPREVDPLVYNMSHEDPGNVSYSEIGGLSEQIRELREVIELPLTNPELFQRVGIIP
PKGCLLYGPPGTGKTLLARAVASQLDCNFLKVVSSSIVDKYIGESARLIREMFNYARDHQ
PCIIFMDEIDAIGGRRFSEGTSADREIQRTLMELLNQMDGFDTLHRVKMIMATNRPDTLD
PALLRPGRLDRKIHIDLPNEQARLDILKIHAGPITKHGEIDYEAIVKLSDGFNGADLRNV
CTEAGMFAIRADHDFVVQEDFMKAVRKVADSKKLESKLDYKPV

Sequence of entity 5 (F), FASTA

>7W3C_5 26S protease regulatory subunit 6A (chains F)
MNLLPNIESPVTRQEKMATVWDEAEQDGIGEEVLKMSTEEIIQRTRLLDSEIKIMKSEVL
RVTHELQAMKDKIKENSEKIKVNKTLPYLVSNVIELLDVDPNDQEEDGANIDLDSQRKGK
CAVIKTSTRQTYFLPVIGLVDAEKLKPGDLVGVNKDSYLILETLPTEYDSRVKAMEVDER
PTEQYSDIGGLDKQIQELVEAIVLPMNHKEKFENLGIQPPKGVLMYGPPGTGKTLLARAC
AAQTKATFLKLAGPQLVQMFIGDGAKLVRDAFALAKEKAPSIIFIDELDAIGTKRFDSEK
AGDREVQRTMLELLNQLDGFQPNTQVKVIAATNRVDILDPALLRSGRLDRKIEFPMPNEE
ARARIMQIHSRKMNVSPDVNYEELARCTDDFNGAQCKAVCVEAGMIALRRGATELTHEDY
MEGILEVQAKKKANLQYYA

Sequence of entity 6 (A), FASTA

>7W3C_6 26S protease regulatory subunit 7 (chains A)
MPDYLGADQRKTKEDEKDDKPIRALDEGDIALLKTYGQSTYSRQIKQVEDDIQQLLKKIN
ELTGIKESDTGLAPPALWDLAADKQTLQSEQPLQVARCTKIINADSEDPKYIINVKQFAK
FVVDLSDQVAPTDIEEGMRVGVDRNKYQIHIPLPPKIDPTVTMMQVEEKPDVTYSDVGGC
KEQIEKLREVVETPLLHPERFVNLGIEPPKGVLLFGPPGTGKTLCARAVANRTDACFIRV
IGSELVQKYVGEGARMVRELFEMARTKKACLIFFDEIDAIGGARFDDGAGGDNEVQRTML
ELINQLDGFDPRGNIKVLMATNRPDTLDPALMRPGRLDRKIEFSLPDLEGRTHIFKIHAR
SMSVERDIRFELLARLCPNSTGAEIRSVCTEAGMFAIRARRKIATEKDFLEAVNKVIKSY
AKFSATPRYMTYN

Sequence of entity 7 (G, g), FASTA

>7W3C_7 Proteasome subunit alpha type-6 (chains G, g)
MSRGSSAGFDRHITIFSPEGRLYQVEYAFKAINQGGLTSVAVRGKDCAVIVTQKKVPDKL
LDSSTVTHLFKITENIGCVMTGMTADSRSQVQRARYEAANWKYKYGYEIPVDMLCKRIAD
ISQVYTQNAEMRPLGCCMILIGIDEEQGPQVYKCDPAGYYCGFKATAAGVKQTESTSFLE
KKVKKKFDWTFEQTVETAITCLSTVLSIDFKPSEIEVGVVTVENPKFRILTEAEIDAHLV
ALAERD

Sequence of entity 8 (H, h), FASTA

>7W3C_8 Proteasome subunit alpha type-2 (chains H, h)
MAERGYSFSLTTFSPSGKLVQIEYALAAVAGGAPSVGIKAANGVVLATEKKQKSILYDER
SVHKVEPITKHIGLVYSGMGPDYRVLVHRARKLAQQYYLVYQEPIPTAQLVQRVASVMQE
YTQSGGVRPFGVSLLICGWNEGRPYLFQSDPSGAYFAWKATAMGKNYVNGKTFLEKRYNE
DLELEDAIHTAILTLKESFEGQMTEDNIEVGICNEAGFRRLTPTEVKDYLAAIA

Sequence of entity 9 (I, i), FASTA

>7W3C_9 Proteasome subunit alpha type-4 (chains I, i)
MSRRYDSRTTIFSPEGRLYQVEYAMEAIGHAGTCLGILANDGVLLAAERRNIHKLLDEVF
FSEKIYKLNEDMACSVAGITSDANVLTNELRLIAQRYLLQYQEPIPCEQLVTALCDIKQA
YTQFGGKRPFGVSLLYIGWDKHYGFQLYQSDPSGNYGGWKATCIGNNSAAAVSMLKQDYK
EGEMTLKSALALAIKVLNKTMDVSKLSAEKVEIATLTRENGKTVIRVLKQKEVEQLIKKH
EEEEAKAEREKKEKEQKEKDK

Sequence of entity 10 (J, j), FASTA

>7W3C_10 Proteasome subunit alpha type-7 (chains J, j)
MSYDRAITVFSPDGHLFQVEYAQEAVKKGSTAVGVRGRDIVVLGVEKKSVAKLQDERTVR
KICALDDNVCMAFAGLTADARIVINRARVECQSHRLTVEDPVTVEYITRYIASLKQRYTQ
SNGRRPFGISALIVGFDFDGTPRLYQTDPSGTYHAWKANAIGRGAKSVREFLEKNYTDEA
IETDDLTIKLVIKALLEVVQSGGKNIELAVMRRDQSLKILNPEEIEKYVAEIEKEKEENE
KKKQKKAS

Sequence of entity 11 (K, k), FASTA

>7W3C_11 Proteasome subunit alpha type-5 (chains K, k)
MFLTRSEYDRGVNTFSPEGRLFQVEYAIEAIKLGSTAIGIQTSEGVCLAVEKRITSPLME
PSSIEKIVEIDAHIGCAMSGLIADAKTLIDKARVETQNHWFTYNETMTVESVTQAVSNLA
LQFGEEDADPGAMSRPFGVALLFGGVDEKGPQLFHMDPSGTFVQCDARAIGSASEGAQSS
LQEVYHKSMTLKEAIKSSLIILKQVMEEKLNATNIELATVQPGQNFHMFTKEELEEVIKD
I

Sequence of entity 12 (L, l), FASTA

>7W3C_12 Isoform Long of Proteasome subunit alpha type-1 (chains L, l)
MQLSKVKFRNQYDNDVTVWSPQGRIHQIEYAMEAVKQGSATVGLKSKTHAVLVALKRAQS
ELAAHQKKILHVDNHIGISIAGLTADARLLCNFMRQECLDSRFVFDRPLPVSRLVSLIGS
KTQIPTQRYGRRPYGVGLLIAGYDDMGPHIFQTCPSANYFDCRAMSIGARSQSARTYLER
HMSEFMECNLNELVKHGLRALRETLPAEQDLTTKNVSIGIVGKDLEFTIYDDDDVSPFLE
GLEERPQRKAQPAQPADEPAEKADEPMEH

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	1
ADP	Adenosine-5'-diphosphate	C10 H15 N5 O10 P2	2
ATP	Adenosine-5'-triphosphate	C10 H16 N5 O13 P3	3

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 P62191 (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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