7W3A: 26S protease regulatory subunit 7

Structure of USP14-bound human 26S proteasome in substrate-engaged state ED4_USP14. Determined by electron microscopy at 3.5 Å resolution. Released 4 May 2022.

Method: Electron microscopy
Resolution: 3.5 Å
Organisms: Homo sapiens, Saccharomyces cerevisiae
Chains: 49
Atoms: 110,889
Mol. weight: 1735.78 kDa
Ligands: ADP, ATP, ZN
Released: 4 May 2022

Explore 7W3A in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

7W3A contains 612 α-helices and 570 β-strands across 49 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, 4 β-strands

Element	Residues	Length	Sheet
α-helix	5-10	6
α-helix	18-29	12
α-helix	35-45	11
α-helix	54-67	14
α-helix	73-87	15
α-helix	90-102	13
α-helix	109-124	16
α-helix	131-139	9
α-helix	152-161	10
α-helix	170-182	13
α-helix	190-206	17
α-helix	220-223	4
α-helix	231-242	12
α-helix	247-254	8
α-helix	265-275	11
α-helix	277-283	7
α-helix	293-299	7
α-helix	306-317	12
β-strand	320	1	108
β-strand	323-324	2	85
β-strand	331-332	2	85
β-strand	335	1	108
α-helix	340-342	3
α-helix	343-345	3
α-helix	346-370	25

Chain A: 18 helices, 13 β-strands

Element	Residues	Length	Sheet
α-helix	28-35	8
α-helix	42-46	5
α-helix	51-55	5
α-helix	78-81	4
β-strand	98-103	6	1
β-strand	111-114	4	1
β-strand	120-125	6	1
β-strand	139-142	4	1
β-strand	148-151	4	1
α-helix	153-158	6
α-helix	174-176	3
α-helix	181-196	16
α-helix	199-204	6
β-strand	211-212	2	2
β-strand	213-215	3	3
α-helix	222-231	10
β-strand	236-241	6	2
α-helix	242-245	4
β-strand	249	1	4
α-helix	253-266	14
β-strand	270-275	6	2
α-helix	277-279	3
α-helix	293-306	14
β-strand	315-318	4	2
β-strand	321	1	3
α-helix	330-332	3
β-strand	340-342	3	3
α-helix	350-360	11
α-helix	371-375	5
α-helix	382-398	17
α-helix	406-420	15

Chain b: 8 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	4-10	7	112
β-strand	13	1	113
α-helix	14-17	4
α-helix	25-43	19
β-strand	48-53	6	112
β-strand	65	1	112
α-helix	69-75	7
β-strand	82	1	113
α-helix	87-97	11
β-strand	106-113	8	112
α-helix	121-134	14
β-strand	138-142	5	112
α-helix	151-160	10
β-strand	170-171	2	112
α-helix	173-174	2
α-helix	180-183	4

Chain B: 24 helices, 11 β-strands

Element	Residues	Length	Sheet
α-helix	35-38	4
α-helix	59-76	18
α-helix	77-81	5
α-helix	95-102	8
β-strand	107-115	9	7
β-strand	120-124	5	7
β-strand	130-135	6	7
α-helix	141-143	3
β-strand	149-152	4	7
β-strand	159-162	4	7
α-helix	166-168	3
α-helix	169-172	4
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
α-helix	217-219	3
β-strand	221-225	5	8
α-helix	232-241	10
β-strand	247-250	4	8
α-helix	252-254	3
β-strand	259	1	9
α-helix	262-277	16
β-strand	281-284	4	8
α-helix	287-290	4
α-helix	302-316	15
β-strand	326-331	6	8
α-helix	339-341	3
β-strand	347-352	6	8
α-helix	354-356	3
α-helix	360-368	9
α-helix	380-383	4
α-helix	392-405	14
α-helix	416-433	18

Chain c: 15 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	30-33	4	117
α-helix	35-47	13
α-helix	49-51	3
β-strand	54-61	8	117
β-strand	66-74	9	117
α-helix	84-86	3
α-helix	89-101	13
β-strand	110-114	5	117
α-helix	124-136	13
β-strand	141-146	6	117
β-strand	156-162	7	117
α-helix	163-164	2
α-helix	166-169	4
α-helix	189-193	5
β-strand	200-203	4	117
β-strand	205-207	3	117
α-helix	214-221	8
α-helix	227-229	3
α-helix	237-261	25
α-helix	267-273	7
α-helix	286-289	4
α-helix	290-292	3
α-helix	293-308	16

Chain C: 14 helices, 13 β-strands

Element	Residues	Length	Sheet
α-helix	12-64	53
β-strand	71	1	13
β-strand	72-78	7	7
β-strand	84-88	5	7
β-strand	95-99	5	7
β-strand	112-116	5	7
β-strand	122-126	5	7
α-helix	155-170	16
α-helix	172-178	7
α-helix	180-183	4
β-strand	185-188	4	14
α-helix	196-205	10
β-strand	210-214	5	14
α-helix	226-240	15
β-strand	244-248	5	14
α-helix	267-280	14
β-strand	289-294	6	14
α-helix	298-300	3
β-strand	313-315	3	14
α-helix	318-320	3
α-helix	322-332	11
β-strand	338	1	15
α-helix	344-348	5
α-helix	357-372	16
β-strand	378	1	15
α-helix	380-385	6
α-helix	386-390	5

Chain d: 13 helices, 2 β-strands

Element	Residues	Length	Sheet
α-helix	18-28	11
α-helix	29-31	3
α-helix	42-64	23
α-helix	68-81	14
α-helix	95-108	14
α-helix	112-119	8
α-helix	131-145	15
α-helix	149-154	6
α-helix	164-182	19
β-strand	190	1	123
α-helix	191-192	2
α-helix	193-198	6
α-helix	206-211	6
β-strand	221	1	123
α-helix	239-253	15

Chain D: 13 helices, 10 β-strands

Element	Residues	Length	Sheet
α-helix	40-81	42
β-strand	88-97	10	13
β-strand	100-105	6	13
β-strand	109-113	5	13
α-helix	121-123	3
β-strand	129-132	4	13
β-strand	139-142	4	13
α-helix	171-186	16
α-helix	189-193	5
β-strand	201-205	5	20
α-helix	212-219	8
β-strand	226-230	5	20
α-helix	242-257	16
β-strand	260-264	5	20
α-helix	267-269	3
α-helix	283-296	14
β-strand	305-308	4	20
β-strand	329-332	4	20
α-helix	334-336	3
α-helix	338-349	12
α-helix	360-363	4
α-helix	372-388	17
α-helix	396-405	10

41 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
26S protease regulatory subunit 6A	F	protein	439	Homo sapiens	P17980
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 (A), FASTA

>7W3A_1 26S protease regulatory subunit 7 (chains A)
MPDYLGADQRKTKEDEKDDKPIRALDEGDIALLKTYGQSTYSRQIKQVEDDIQQLLKKIN
ELTGIKESDTGLAPPALWDLAADKQTLQSEQPLQVARCTKIINADSEDPKYIINVKQFAK
FVVDLSDQVAPTDIEEGMRVGVDRNKYQIHIPLPPKIDPTVTMMQVEEKPDVTYSDVGGC
KEQIEKLREVVETPLLHPERFVNLGIEPPKGVLLFGPPGTGKTLCARAVANRTDACFIRV
IGSELVQKYVGEGARMVRELFEMARTKKACLIFFDEIDAIGGARFDDGAGGDNEVQRTML
ELINQLDGFDPRGNIKVLMATNRPDTLDPALMRPGRLDRKIEFSLPDLEGRTHIFKIHAR
SMSVERDIRFELLARLCPNSTGAEIRSVCTEAGMFAIRARRKIATEKDFLEAVNKVIKSY
AKFSATPRYMTYN

Sequence of entity 2 (B), FASTA

>7W3A_2 26S protease regulatory subunit 4 (chains B)
MGQSQSGGHGPGGGKKDDKDKKKKYEPPVPTRVGKKKKKTKGPDAASKLPLVTPHTQCRL
KLLKLERIKDYLLMEEEFIRNQEQMKPLEEKQEEERSKVDDLRGTPMSVGTLEEIIDDNH
AIVSTSVGSEHYVSILSFVDKDLLEPGCSVLLNHKVHAVIGVLMDDTDPLVTVMKVEKAP
QETYADIGGLDNQIQEIKESVELPLTHPEYYEEMGIKPPKGVILYGPPGTGKTLLAKAVA
NQTSATFLRVVGSELIQKYLGDGPKLVRELFRVAEEHAPSIVFIDEIDAIGTKRYDSNSG
GEREIQRTMLELLNQLDGFDSRGDVKVIMATNRIETLDPALIRPGRIDRKIEFPLPDEKT
KKRIFQIHTSRMTLADDVTLDDLIMAKDDLSGADIKAICTEAGLMALRERRMKVTNEDFK
KSKENVLYKKQEGTPEGLYL

Sequence of entity 3 (C), FASTA

>7W3A_3 Isoform 2 of 26S proteasome regulatory subunit 8 (chains C)
MELEEGKAGSGLRQYYLSKIEELQLIVNDKSQNLRRLQAQRNELNAKVRLLREELQLLQE
QGSYVGEVVRAMDKKKVLVKVHPEGKFVVDVDKNIDINDVTPNCRVALRNDSYTLHKILP
NKVDPLVSLMMVEKVPDSTYEMIGGLDKQIKEIKEVIELPVKHPELFEALGIAQPKGVLL
YGPPGTGKTLLARAVAHHTDCTFIRVSGSELVQKFIGEGARMVRELFVMAREHAPSIIFM
DEIDSIGSSRLEGGSGGDSEVQRTMLELLNQLDGFEATKNIKVIMATNRIDILDSALLRP
GRIDRKIEFPPPNEEARLDILKIHSRKMNLTRGINLRKIAELMPGASGAEVKGVCTEAGM
YALRERRVHVTQEDFEMAVAKVMQKDSEKNMSIKKLWK

Sequence of entity 4 (D), FASTA

>7W3A_4 26S protease regulatory subunit 6B (chains D)
MEEIGILVEKAQDEIPALSVSRPQTGLSFLGPEPEDLEDLYSRYKKLQQELEFLEVQEEY
IKDEQKNLKKEFLHAQEEVKRIQSIPLVIGQFLEAVDQNTAIVGSTTGSNYYVRILSTID
RELLKPNASVALHKHSNALVDVLPPEADSSIMMLTSDQKPDVMYADIGGMDIQKQEVREA
VELPLTHFELYKQIGIDPPRGVLMYGPPGCGKTMLAKAVAHHTTAAFIRVVGSEFVQKYL
GEGPRMVRDVFRLAKENAPAIIFIDEIDAIATKRFDAQTGADREVQRILLELLNQMDGFD
QNVNVKVIMATNRADTLDPALLRPGRLDRKIEFPLPDRRQKRLIFSTITSKMNLSEEVDL
EDYVARPDKISGADINSICQESGMLAVRENRYIVLAKDFEKAYKTVIKKDEQEHEFYK

Sequence of entity 5 (E), FASTA

>7W3A_5 26S proteasome regulatory subunit 10B (chains E)
MAIPGIPYERRLLIMADPRDKALQDYRKKLLEHKEIDGRLKELREQLKELTKQYEKSEND
LKALQSVGQIVGEVLKQLTEEKFIVKATNGPRYVVGCRRQLDKSKLKPGTRVALDMTTLT
IMRYLPREVDPLVYNMSHEDPGNVSYSEIGGLSEQIRELREVIELPLTNPELFQRVGIIP
PKGCLLYGPPGTGKTLLARAVASQLDCNFLKVVSSSIVDKYIGESARLIREMFNYARDHQ
PCIIFMDEIDAIGGRRFSEGTSADREIQRTLMELLNQMDGFDTLHRVKMIMATNRPDTLD
PALLRPGRLDRKIHIDLPNEQARLDILKIHAGPITKHGEIDYEAIVKLSDGFNGADLRNV
CTEAGMFAIRADHDFVVQEDFMKAVRKVADSKKLESKLDYKPV

Sequence of entity 6 (F), FASTA

>7W3A_6 26S protease regulatory subunit 6A (chains F)
MNLLPNIESPVTRQEKMATVWDEAEQDGIGEEVLKMSTEEIIQRTRLLDSEIKIMKSEVL
RVTHELQAMKDKIKENSEKIKVNKTLPYLVSNVIELLDVDPNDQEEDGANIDLDSQRKGK
CAVIKTSTRQTYFLPVIGLVDAEKLKPGDLVGVNKDSYLILETLPTEYDSRVKAMEVDER
PTEQYSDIGGLDKQIQELVEAIVLPMNHKEKFENLGIQPPKGVLMYGPPGTGKTLLARAC
AAQTKATFLKLAGPQLVQMFIGDGAKLVRDAFALAKEKAPSIIFIDELDAIGTKRFDSEK
AGDREVQRTMLELLNQLDGFQPNTQVKVIAATNRVDILDPALLRSGRLDRKIEFPMPNEE
ARARIMQIHSRKMNVSPDVNYEELARCTDDFNGAQCKAVCVEAGMIALRRGATELTHEDY
MEGILEVQAKKKANLQYYA

Sequence of entity 7 (G, g), FASTA

>7W3A_7 Proteasome subunit alpha type-6 (chains G, g)
MSRGSSAGFDRHITIFSPEGRLYQVEYAFKAINQGGLTSVAVRGKDCAVIVTQKKVPDKL
LDSSTVTHLFKITENIGCVMTGMTADSRSQVQRARYEAANWKYKYGYEIPVDMLCKRIAD
ISQVYTQNAEMRPLGCCMILIGIDEEQGPQVYKCDPAGYYCGFKATAAGVKQTESTSFLE
KKVKKKFDWTFEQTVETAITCLSTVLSIDFKPSEIEVGVVTVENPKFRILTEAEIDAHLV
ALAERD

Sequence of entity 8 (H, h), FASTA

>7W3A_8 Proteasome subunit alpha type-2 (chains H, h)
MAERGYSFSLTTFSPSGKLVQIEYALAAVAGGAPSVGIKAANGVVLATEKKQKSILYDER
SVHKVEPITKHIGLVYSGMGPDYRVLVHRARKLAQQYYLVYQEPIPTAQLVQRVASVMQE
YTQSGGVRPFGVSLLICGWNEGRPYLFQSDPSGAYFAWKATAMGKNYVNGKTFLEKRYNE
DLELEDAIHTAILTLKESFEGQMTEDNIEVGICNEAGFRRLTPTEVKDYLAAIA

Sequence of entity 9 (I, i), FASTA

>7W3A_9 Proteasome subunit alpha type-4 (chains I, i)
MSRRYDSRTTIFSPEGRLYQVEYAMEAIGHAGTCLGILANDGVLLAAERRNIHKLLDEVF
FSEKIYKLNEDMACSVAGITSDANVLTNELRLIAQRYLLQYQEPIPCEQLVTALCDIKQA
YTQFGGKRPFGVSLLYIGWDKHYGFQLYQSDPSGNYGGWKATCIGNNSAAAVSMLKQDYK
EGEMTLKSALALAIKVLNKTMDVSKLSAEKVEIATLTRENGKTVIRVLKQKEVEQLIKKH
EEEEAKAEREKKEKEQKEKDK

Sequence of entity 10 (J, j), FASTA

>7W3A_10 Proteasome subunit alpha type-7 (chains J, j)
MSYDRAITVFSPDGHLFQVEYAQEAVKKGSTAVGVRGRDIVVLGVEKKSVAKLQDERTVR
KICALDDNVCMAFAGLTADARIVINRARVECQSHRLTVEDPVTVEYITRYIASLKQRYTQ
SNGRRPFGISALIVGFDFDGTPRLYQTDPSGTYHAWKANAIGRGAKSVREFLEKNYTDEA
IETDDLTIKLVIKALLEVVQSGGKNIELAVMRRDQSLKILNPEEIEKYVAEIEKEKEENE
KKKQKKAS

Sequence of entity 11 (K, k), FASTA

>7W3A_11 Proteasome subunit alpha type-5 (chains K, k)
MFLTRSEYDRGVNTFSPEGRLFQVEYAIEAIKLGSTAIGIQTSEGVCLAVEKRITSPLME
PSSIEKIVEIDAHIGCAMSGLIADAKTLIDKARVETQNHWFTYNETMTVESVTQAVSNLA
LQFGEEDADPGAMSRPFGVALLFGGVDEKGPQLFHMDPSGTFVQCDARAIGSASEGAQSS
LQEVYHKSMTLKEAIKSSLIILKQVMEEKLNATNIELATVQPGQNFHMFTKEELEEVIKD
I

Sequence of entity 12 (L, l), FASTA

>7W3A_12 Isoform Long of Proteasome subunit alpha type-1 (chains L, l)
MQLSKVKFRNQYDNDVTVWSPQGRIHQIEYAMEAVKQGSATVGLKSKTHAVLVALKRAQS
ELAAHQKKILHVDNHIGISIAGLTADARLLCNFMRQECLDSRFVFDRPLPVSRLVSLIGS
KTQIPTQRYGRRPYGVGLLIAGYDDMGPHIFQTCPSANYFDCRAMSIGARSQSARTYLER
HMSEFMECNLNELVKHGLRALRETLPAEQDLTTKNVSIGIVGKDLEFTIYDDDDVSPFLE
GLEERPQRKAQPAQPADEPAEKADEPMEH

Ligands and cofactors

ID	Name	Formula	Copies
ADP	Adenosine-5'-diphosphate	C10 H15 N5 O10 P2	2
ATP	Adenosine-5'-triphosphate	C10 H16 N5 O13 P3	4
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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