6MSB: 26S proteasome non-ATPase regulatory subunit 1

Cryo-EM structures and dynamics of substrate-engaged human 26S proteasome. Determined by electron microscopy at 3.0 Å resolution. Released 5 Dec 2018.

Method: Electron microscopy
Resolution: 3.0 Å
Organism: Homo sapiens
Chains: 48
Atoms: 104,938
Mol. weight: 1681.81 kDa
Ligands: ZN, ATP, ADP, MG
Released: 5 Dec 2018

Explore 6MSB in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

6MSB contains 607 α-helices and 547 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: 21 helices, 4 β-strands

Element	Residues	Length	Sheet
α-helix	5-10	6
α-helix	20-28	9
α-helix	29-31	3
α-helix	38-41	4
α-helix	43-46	4
α-helix	52-55	4
α-helix	57-66	10
α-helix	74-87	14
α-helix	90-103	14
α-helix	111-121	11
α-helix	130-137	8
α-helix	149-164	16
α-helix	170-179	10
α-helix	190-206	17
α-helix	224-226	3
α-helix	231-241	11
α-helix	245-250	6
α-helix	265-283	19
β-strand	292	1	40
α-helix	293-299	7
α-helix	304-317	14
β-strand	320-325	6	15
β-strand	330	1	40
β-strand	331-336	6	15
α-helix	345-370	26

Chain A: 14 helices, 16 β-strands

Element	Residues	Length	Sheet
α-helix	42-60	19
α-helix	79-81	3
β-strand	97-98	2	69
β-strand	100-103	4	70
β-strand	111-114	4	70
β-strand	122-123	2	70
β-strand	124	1	71
β-strand	140-142	3	69
β-strand	148	1	71
β-strand	149-152	4	69
β-strand	165	1	72
α-helix	174-176	3
α-helix	181-191	11
α-helix	193-196	4
β-strand	212-215	4	72
α-helix	222-232	11
β-strand	236-241	6	72
α-helix	242-244	3
α-helix	252-266	15
β-strand	270-275	6	72
α-helix	293-304	12
β-strand	315-321	7	72
α-helix	324-326	3
β-strand	339-342	4	72
α-helix	349-358	10
β-strand	364	1	73
α-helix	371-374	4
α-helix	382-398	17
β-strand	404	1	73
α-helix	406-416	11

Chain b: 7 helices, 5 β-strands

Element	Residues	Length	Sheet
β-strand	4-10	7	45
α-helix	25-43	19
β-strand	51-53	3	45
α-helix	69-74	6
α-helix	89-98	10
β-strand	106-113	8	45
α-helix	121-132	12
β-strand	137-142	6	45
α-helix	154-158	5
β-strand	170-171	2	45
α-helix	172-174	3
α-helix	185-189	5

Chain B: 18 helices, 19 β-strands

Element	Residues	Length	Sheet
α-helix	59-85	27
α-helix	94-102	9
β-strand	107-108	2	76
β-strand	109-112	4	77
β-strand	116	1	78
β-strand	120	1	78
β-strand	123-124	2	77
β-strand	130-131	2	77
β-strand	134	1	78
β-strand	135	1	79
β-strand	149-151	3	77
β-strand	159	1	79
β-strand	161-162	2	77
α-helix	171-174	4
β-strand	175	1	80
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	80
α-helix	232-242	11
β-strand	246-251	6	80
α-helix	253-255	3
α-helix	262-275	14
β-strand	280-285	6	80
α-helix	288-290	3
α-helix	302-316	15
β-strand	325-331	7	80
β-strand	349-352	4	80
α-helix	354-357	4
α-helix	361-368	8
β-strand	374	1	81
α-helix	380-384	5
α-helix	392-408	17
β-strand	414	1	81
α-helix	416-426	11

Chain c: 14 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	31-32	2	49
α-helix	37-47	11
β-strand	54-60	7	49
β-strand	67-74	8	49
α-helix	75-76	2
α-helix	89-96	8
β-strand	107-114	8	49
α-helix	124-131	8
α-helix	140	1
β-strand	141-145	5	49
β-strand	160-162	3	49
α-helix	166-170	5
α-helix	186-188	3
α-helix	189-193	5
β-strand	200-202	3	49
β-strand	205-206	2	49
α-helix	211-218	8
α-helix	234-243	10
α-helix	245-255	11
α-helix	269-272	4
α-helix	273-275	3
α-helix	280-308	29

Chain C: 23 helices, 15 β-strands

Element	Residues	Length	Sheet
α-helix	20-23	4
α-helix	25-65	41
α-helix	69-70	2
β-strand	71-73	3	84
β-strand	75-80	6	76
β-strand	84-89	6	76
β-strand	92-98	7	76
β-strand	112	1	76
β-strand	113-116	4	84
β-strand	122	1	76
β-strand	123-127	5	84
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	173-176	4
α-helix	181-183	3
β-strand	185-189	5	85
α-helix	196-206	11
β-strand	210-214	5	85
α-helix	221-224	4
α-helix	232-234	3
α-helix	235-238	4
β-strand	244-248	5	85
α-helix	268-279	12
β-strand	289-294	6	85
α-helix	298-300	3
β-strand	313-316	4	85
α-helix	318-320	3
α-helix	322-332	11
α-helix	333-335	3
β-strand	338	1	86
α-helix	339	1
α-helix	344-348	5
α-helix	358-372	15
β-strand	378	1	86
α-helix	380-394	15

Chain d: 11 helices, 2 β-strands

Element	Residues	Length	Sheet
α-helix	19-32	14
α-helix	49-64	16
α-helix	71-79	9
α-helix	96-108	13
α-helix	112-116	5
α-helix	118-120	3
α-helix	131-144	14
α-helix	149-153	5
α-helix	164-181	18
α-helix	205-212	8
β-strand	217	1	55
β-strand	222	1	55
α-helix	236-251	16

Chain D: 19 helices, 11 β-strands

Element	Residues	Length	Sheet
α-helix	42-67	26
α-helix	69-78	10
α-helix	79-83	5
β-strand	87-91	5	90
β-strand	92-97	6	84
β-strand	100-104	5	84
β-strand	110-114	5	84
α-helix	121-123	3
β-strand	129-132	4	90
β-strand	139-142	4	90
α-helix	164-166	3
α-helix	171-181	11
α-helix	183-186	4
α-helix	188-190	3
α-helix	191-195	5
β-strand	201-205	5	91
α-helix	212-219	8
β-strand	227-231	5	91
α-helix	242-256	15
β-strand	261-265	5	91
α-helix	267-269	3
α-helix	281-295	15
β-strand	306-310	5	91
β-strand	329-332	4	91
α-helix	334-336	3
α-helix	341-349	9
α-helix	361-364	4
α-helix	374-388	15
α-helix	396-406	11
α-helix	415-417	3

29 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 proteasome non-ATPase regulatory subunit 1	U	protein	953	Homo sapiens	Q99460 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 3	V	protein	534	Homo sapiens	O43242 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 12	W	protein	456	Homo sapiens	O00232 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 11	X	protein	422	Homo sapiens	O00231 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 6	Y	protein	389	Homo sapiens	Q15008
26S proteasome non-ATPase regulatory subunit 7	Z	protein	324	Homo sapiens	P51665
26S proteasome non-ATPase regulatory subunit 13	a	protein	376	Homo sapiens	Q9UNM6
26S proteasome non-ATPase regulatory subunit 4	b	protein	377	Homo sapiens	P55036
26S proteasome non-ATPase regulatory subunit 14	c	protein	309	Homo sapiens	O00487
26S proteasome non-ATPase regulatory subunit 8	d	protein	349	Homo sapiens	P48556
26S proteasome complex subunit SEM1	e	protein	70	Homo sapiens	P60896
26S proteasome non-ATPase regulatory subunit 2	f	protein	908	Homo sapiens	Q13200

21 more molecules are not listed.

Sequence of entity 1 (U), FASTA

>6MSB_1 26S proteasome non-ATPase regulatory subunit 1 (chains U)
MITSAAGIISLLDEDEPQLKEFALHKLNAVVNDFWAEISESVDKIEVLYEDEGFRSRQFA
ALVASKVFYHLGAFEESLNYALGAGDLFNVNDNSEYVETIIAKCIDHYTKQCVENADLPE
GEKKPIDQRLEGIVNKMFQRCLDDHKYKQAIGIALETRRLDVFEKTILESNDVPGMLAYS
LKLCMSLMQNKQFRNKVLRVLVKIYMNLEKPDFINVCQCLIFLDDPQAVSDILEKLVKED
NLLMAYQICFDLYESASQQFLSSVIQNLRTVGTPIASVPGSTNTGTVPGSEKDSDSMETE
EKTSSAFVGKTPEASPEPKDQTLKMIKILSGEMAIELHLQFLIRNNNTDLMILKNTKDAV
RNSVCHTATVIANSFMHCGTTSDQFLRDNLEWLARATNWAKFTATASLGVIHKGHEKEAL
QLMATYLPKDTSPGSAYQEGGGLYALGLIHANHGGDIIDYLLNQLKNASNDIVRHGGSLG
LGLAAMGTARQDVYDLLKTNLYQDDAVTGEAAGLALGLVMLGSKNAQAIEDMVGYAQETQ
HEKILRGLAVGIALVMYGRMEEADALIESLCRDKDPILRRSGMYTVAMAYCGSGNNKAIR
RLLHVAVSDVNDDVRRAAVESLGFILFRTPEQCPSVVSLLSESYNPHVRYGAAMALGICC
AGTGNKEAINLLEPMTNDPVNYVRQGALIASALIMIQQTEITCPKVNQFRQLYSKVINDK
HDDVMAKFGAILAQGILDAGGHNVTISLQSRTGHTHMPSVVGVLVFTQFWFWFPLSHFLS
LAYTPTCVIGLNKDLKMPKVQYKSNCKPSTFAYPAPLEVPKEKEKEKVSTAVLSITAKAK
KKEKEKEKKEEEKMEVDEAEKKEEKEKKKEPEPNFQLLDNPARVMPAQLKVLTMPETCRY
QPFKPLSIGGIIILKDTSEDIEELVEPVAAHGPKIEEEEQEPEPPEPFEYIDD

Sequence of entity 2 (V), FASTA

>6MSB_2 26S proteasome non-ATPase regulatory subunit 3 (chains V)
MKQEGSARRRGADKAKPPPGGGEQEPPPPPAPQDVEMKEEAATGGGSTGEADGKTAAAAA
EHSQRELDTVTLEDIKEHVKQLEKAVSGKEPRFVLRALRMLPSTSRRLNHYVLYKAVQGF
FTSNNATRDFLLPFLEEPMDTEADLQFRPRTGKAASTPLLPEVEAYLQLLVVIFMMNSKR
YKEAQKISDDLMQKISTQNRRALDLVAAKCYYYHARVYEFLDKLDVVRSFLHARLRTATL
RHDADGQATLLNLLLRNYLHYSLYDQAEKLVSKSVFPEQANNNEWARYLYYTGRIKAIQL
EYSEARRTMTNALRKAPQHTAVGFKQTVHKLLIVVELLLGEIPDRLQFRQPSLKRSLMPY
FLLTQAVRTGNLAKFNQVLDQFGEKFQADGTYTLIIRLRHNVIKTGVRMISLSYSRISLA
DIAQKLQLDSPEDAEFIVAKAIRDGVIEASINHEKGYVQSKEMIDIYSTREPQLAFHQRI
SFCLDIHNMSVKAMRFPPKSYNKDLESAEERREREQQDLEFAKEMAEDDDDSFP

Sequence of entity 3 (W), FASTA

>6MSB_3 26S proteasome non-ATPase regulatory subunit 12 (chains W)
MADGGSERADGRIVKMEVDYSATVDQRLPECAKLAKEGRLQEVIETLLSLEKQTRTASDM
VSTSRILVAVVKMCYEAKEWDLLNENIMLLSKRRSQLKQAVAKMVQQCCTYVEEITDLPI
KLRLIDTLRMVTEGKIYVEIERARLTKTLATIKEQNGDVKEAASILQELQVETYGSMEKK
ERVEFILEQMRLCLAVKDYIRTQIISKKINTKFFQEENTEKLKLKYYNLMIQLDQHEGSY
LSICKHYRAIYDTPCIQAESEKWQQALKSVVLYVILAPFDNEQSDLVHRISGDKKLEEIP
KYKDLLKLFTTMELMRWSTLVEDYGMELRKGSLESPATDVFGSTEEGEKRWKDLKNRVVE
HNIRIMAKYYTRITMKRMAQLLDLSVDESEAFLSNLVVNKTIFAKVDRLAGIINFQRPKD
PNNLLNDWSQKLNSLMSLVNKTTHLIAKEEMIHNLQ

Sequence of entity 4 (X), FASTA

>6MSB_4 26S proteasome non-ATPase regulatory subunit 11 (chains X)
MAAAAVVEFQRAQSLLSTDREASIDILHSIVKRDIQENDEEAVQVKEQSILELGSLLAKT
GQAAELGGLLKYVRPFLNSISKAKAARLVRSLLDLFLDMEAATGQEVELCLECIEWAKSE
KRTFLRQALEARLVSLYFDTKRYQEALHLGSQLLRELKKMDDKALLVEVQLLESKTYHAL
SNLPKARAALTSARTTANAIYCPPKLQATLDMQSGIIHAAEEKDWKTAYSYFYEAFEGYD
SIDSPKAITSLKYMLLCKIMLNTPEDVQALVSGKLALRYAGRQTEALKCVAQASKNRSLA
DFEKALTDYRAELRDDPIISTHLAKLYDNLLEQNLIRVIEPFSRVQIEHISSLIKLSKAD
VERKLSQMILDKKFHGILDQGEGVLIIFDEPPVDKTYEAALETIQNMSKVVDSLYNKAKK
LT

Sequence of entity 5 (Y), FASTA

>6MSB_5 26S proteasome non-ATPase regulatory subunit 6 (chains Y)
MPLENLEEEGLPKNPDLRIAQLRFLLSLPEHRGDAAVRDELMAAVRDNNMAPYYEALCKS
LDWQIDVDLLNKMKKANEDELKRLDEELEDAEKNLGESEIRDAMMAKAEYLCRIGDKEGA
LTAFRKTYDKTVALGHRLDIVFYLLRIGLFYMDNDLITRNTEKAKSLIEEGGDWDRRNRL
KVYQGLYCVAIRDFKQAAELFLDTVSTFTSYELMDYKTFVTYTVYVSMIALERPDLREKV
IKGAEILEVLHSLPAVRQYLFSLYECRYSVFFQSLAVVEQEMKKDWLFAPHYRYYVREMR
IHAYSQLLESYRSLTLGYMAEAFGVGVEFIDQELSRFIAAGRLHCKIDKVNEIVETNRPD
SKNWQYQETIKKGDLLLNRVQKLSRVINM

Sequence of entity 6 (Z), FASTA

>6MSB_6 26S proteasome non-ATPase regulatory subunit 7 (chains Z)
MPELAVQKVVVHPLVLLSVVDHFNRIGKVGNQKRVVGVLLGSWQKKVLDVSNSFAVPFDE
DDKDDSVWFLDHDYLENMYGMFKKVNARERIVGWYHTGPKLHKNDIAINELMKRYCPNSV
LVIIDVKPKDLGLPTEAYISVEEVHDDGTPTSKTFEHVTSEIGAEEAEEVGVEHLLRDIK
DTTVGTLSQRITNQVHGLKGLNSKLLDIRSYLEKVATGKLPINHQIIYQLQDVFNLLPDV
SLQEFVKAFYLKTNDQMVVVYLASLIRSVVALHNLINNKIANRDAEKKEGQEKEESKKDR
KEDKEKDKDKEKSDVKKEEKKEKK

Sequence of entity 7 (a), FASTA

>6MSB_7 26S proteasome non-ATPase regulatory subunit 13 (chains a)
MKDVPGFLQQSQNSGPGQPAVWHRLEELYTKKLWHQLTLQVLDFVQDPCFAQGDGLIKLY
ENFISEFEHRVNPLSLVEIILHVVRQMTDPNVALTFLEKTREKVKSSDEAVILCKTAIGA
LKLNIGDLQVTKETIEDVEEMLNNLPGVTSVHSRFYDLSSKYYQTIGNHASYYKDALRFL
GCVDIKDLPVSEQQERAFTLGLAGLLGEGVFNFGELLMHPVLESLRNTDRQWLIDTLYAF
NSGNVERFQTLKTAWGQQPDLAANEAQLLRKIQLLCLMEMTFTRPANHRQLTFEEIAKSA
KITVNEVELLVMKALSVGLVKGSIDEVDKRVHMTWVQPRVLDLQQIKGMKDRLEFWCTDV
KSMEMLVEHQAHDILT

Sequence of entity 8 (b), FASTA

>6MSB_8 26S proteasome non-ATPase regulatory subunit 4 (chains b)
MVLESTMVCVDNSEYMRNGDFLPTRLQAQQDAVNIVCHSKTRSNPENNVGLITLANDCEV
LTTLTPDTGRILSKLHTVQPKGKITFCTGIRVAHLALKHRQGKNHKMRIIAFVGSPVEDN
EKDLVKLAKRLKKEKVNVDIINFGEEEVNTEKLTAFVNTLNGKDGTGSHLVTVPPGPSLA
DALISSPILAGEGGAMLGLGASDFEFGVDPSADPELALALRVSMEEQRQRQEEEARRAAA
ASAAEAGIATTGTEDSDDALLKMTISQQEFGRTGLPDLSSMTEEEQIAYAMQMSLQGAEF
GQAESADIDASSAMDTSEPAKEEDDYDVMQDPEFLQSVLENLPGVDPNNEAIRNAMGSLA
SQATKDGKKDKKEEDKK

Sequence of entity 9 (c), FASTA

>6MSB_9 26S proteasome non-ATPase regulatory subunit 14 (chains c)
DRLLRLGGGMPGLGQGPPTDAPAVDTAEQVYISSLALLKMLKHGRAGVPMEVMGLMLGEF
VDDYTVRVIDVFAMPQSGTGVSVEAVDPVFQAKMLDMLKQTGRPEMVVGWYHSHPGFGCW
LSGVDINTQQSFEALSERAVAVVVDPIQSVKGKVVIDAFRLINANMMVLGHEPRQTTSNL
GHLNKPSIQALIHGLNRHYYSITINYRKNELEQKMLLNLHKKSWMEGLTLQDYSEHCKHN
ESVVKEMLELAKNYNKAVEEEDKMTPEQLAIKNVGKQDPKRHLEEHVDVLMTSNIVQCLA
AMLDTVVFK

Sequence of entity 10 (d), FASTA

>6MSB_10 26S proteasome non-ATPase regulatory subunit 8 (chains d)
FIKGRAPRAPPRERRRATRGGLRQVVAPPRALGSTSRPHFRRASVCRRRCRKSGGLLAAS
RKMAAAAVNGAAGFSSSGPAATSGAVLQAATGMYEQLKGEWNRKSPNLSKCGEELGRLKL
VLLELNFLPTTGTKLTKQQLILARDILEIGAQWSILRKDIPSFERYMAQLKCYYFDYKEQ
LPESAYMHQLLGLNLLFLLSQNRVAEFHTELERLPAKDIQTNVYIKHPVSLEQYLMEGSY
NKVFLAKGNIPAESYTFFIDILLDTIRDEIAGCIEKAYEKILFTEATRILFFNTPKKMTD
YAKKRGWVLGPNNYYSFASQQQKPEDTTIPSTELAKQVIEYARQLEMIV

Sequence of entity 11 (e), FASTA

>6MSB_11 26S proteasome complex subunit SEM1 (chains e)
MSEKKQPVDLGLLEEDDEFEEFPAEDWAGLDEDEDAHVWEDNWDDDNVEDDFSNQLRAEL
EKHGYKMETS

Sequence of entity 12 (f), FASTA

>6MSB_12 26S proteasome non-ATPase regulatory subunit 2 (chains f)
MEEGGRDKAPVQPQQSPAAAPGGTDEKPSGKERRDAGDKDKEQELSEEDKQLQDELEMLV
ERLGEKDTSLYRPALEELRRQIRSSTTSMTSVPKPLKFLRPHYGKLKEIYENMAPGENKR
FAADIISVLAMTMSGERECLKYRLVGSQEELASWGHEYVRHLAGEVAKEWQELDDAEKVQ
REPLLTLVKEIVPYNMAHNAEHEACDLLMEIEQVDMLEKDIDENAYAKVCLYLTSCVNYV
PEPENSALLRCALGVFRKFSRFPEALRLALMLNDMELVEDIFTSCKDVVVQKQMAFMLGR
HGVFLELSEDVEEYEDLTEIMSNVQLNSNFLALARELDIMEPKVPDDIYKTHLENNRFGG
SGSQVDSARMNLASSFVNGFVNAAFGQDKLLTDDGNKWLYKNKDHGMLSAAASLGMILLW
DVDGGLTQIDKYLYSSEDYIKSGALLACGIVNSGVRNECDPALALLSDYVLHNSNTMRLG
SIFGLGLAYAGSNREDVLTLLLPVMGDSKSSMEVAGVTALACGMIAVGSCNGDVTSTILQ
TIMEKSETELKDTYARWLPLGLGLNHLGKGEAIEAILAALEVVSEPFRSFANTLVDVCAY
AGSGNVLKVQQLLHICSEHFDSKEKEEDKDKKEKKDKDKKEAPADMGAHQGVAVLGIALI
AMGEEIGAEMALRTFGHLLRYGEPTLRRAVPLALALISVSNPRLNILDTLSKFSHDADPE
VSYNSIFAMGMVGSGTNNARLAAMLRQLAQYHAKDPNNLFMVRLAQGLTHLGKGTLTLCP
YHSDRQLMSQVAVAGLLTVLVSFLDVRNIILGKSHYVLYGLVAAMQPRMLVTFDEELRPL
PVSVRVGQAVDVVGQAGKPKTITGFQTHTTPVLLAHGERAELATEEFLPVTPILEGFVIL
RKNPNYDL

Ligands and cofactors

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

Primary citation

Cryo-EM structures and dynamics of substrate-engaged human 26S proteasome. Dong, Y., Zhang, S., Wu, Z. et al. Nature (2019) 565:49-55. DOI 10.1038/s41586-018-0736-4 · PubMed

Other PDB entries of the same protein (UniProt Q99460 (AlphaFold model), which also has an AlphaFold model), best resolution first:

5V1Y 1.42 Å, Crystal structure of the ternary RPN13 PRU-RPN2 (940-953)-ubiquitin complex
6OI4 1.76 Å, RPN13 (19-132)-RPN2 (940-952) pY950-Ub complex
5V1Z 2.0 Å, Crystal structure of the RPN13 PRU-RPN2 (932-953)-ubiquitin complex
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…
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

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