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

Cryo-EM structures and dynamics of substrate-engaged human 26S proteasome. Determined by electron microscopy at 3.3 Å resolution. Released 21 Nov 2018.

Method: Electron microscopy
Resolution: 3.3 Å
Organisms: Homo sapiens, Saccharomyces cerevisiae RM11-1a
Chains: 48
Atoms: 105,062
Mol. weight: 1673.08 kDa
Ligands: ZN, ATP, MG, ADP
Released: 21 Nov 2018

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

Secondary structure: helices and β-sheets

6MSE contains 610 α-helices and 553 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, 5 β-strands

Element	Residues	Length	Sheet
α-helix	7-11	5
α-helix	18-28	11
α-helix	29-31	3
α-helix	36-46	11
α-helix	54-66	13
α-helix	74-87	14
α-helix	90-103	14
α-helix	111-120	10
α-helix	134-141	8
α-helix	150-162	13
α-helix	170-182	13
α-helix	190-205	16
α-helix	223-228	6
α-helix	232-242	11
α-helix	249-256	8
α-helix	265-273	9
α-helix	277-283	7
β-strand	291-292	2	37
α-helix	293-299	7
α-helix	304-306	3
α-helix	307-317	11
β-strand	320	1	38
β-strand	323-324	2	37
β-strand	330-332	3	37
β-strand	336	1	38
α-helix	343-370	28

Chain A: 16 helices, 16 β-strands

Element	Residues	Length	Sheet
α-helix	42-63	22
α-helix	87-90	4
β-strand	97-98	2	73
β-strand	100-103	4	74
β-strand	111-114	4	74
β-strand	120-123	4	74
β-strand	124-125	2	73
β-strand	140-142	3	73
β-strand	148-152	5	73
α-helix	159-162	4
β-strand	165	1	75
α-helix	174-177	4
α-helix	181-187	7
α-helix	188-192	5
α-helix	193-196	4
α-helix	200-204	5
β-strand	211-215	5	75
α-helix	222-231	10
β-strand	236-239	4	75
α-helix	242-244	3
β-strand	249	1	76
α-helix	252-266	15
β-strand	270-273	4	75
α-helix	294-306	13
β-strand	315-320	6	75
β-strand	339-342	4	75
α-helix	349-358	10
β-strand	364	1	77
α-helix	371-374	4
α-helix	382-398	17
β-strand	404	1	77
α-helix	406-421	16

Chain b: 8 helices, 6 β-strands

Element	Residues	Length	Sheet
β-strand	4-10	7	43
α-helix	14-16	3
α-helix	25-43	19
β-strand	48-53	6	43
β-strand	65	1	43
α-helix	69-74	6
α-helix	87-94	8
α-helix	96-98	3
β-strand	106-113	8	43
α-helix	121-132	12
β-strand	137-142	6	43
α-helix	154-158	5
β-strand	170-171	2	43
α-helix	180-183	4

Chain B: 19 helices, 16 β-strands

Element	Residues	Length	Sheet
α-helix	59-80	22
α-helix	94-100	7
β-strand	107-108	2	74
β-strand	111-115	5	81
β-strand	120-124	5	81
β-strand	131	1	73
β-strand	132-134	3	81
β-strand	135	1	82
α-helix	136	1
β-strand	149	1	81
β-strand	150-151	2	83
β-strand	152	1	74
β-strand	159	1	82
β-strand	161-162	2	83
α-helix	163-165	3
α-helix	169-174	6
α-helix	193-197	5
α-helix	198-202	5
α-helix	203-206	4
α-helix	211-214	4
α-helix	216-219	4
β-strand	221-225	5	84
α-helix	232-241	10
β-strand	246-251	6	84
α-helix	252-254	3
α-helix	262-277	16
β-strand	280-285	6	84
α-helix	302-316	15
β-strand	325-330	6	84
α-helix	339-342	4
β-strand	349-352	4	84
α-helix	361-368	8
α-helix	381-384	4
α-helix	392-408	17
α-helix	416-431	16

Chain c: 10 helices, 10 β-strands

Element	Residues	Length	Sheet
β-strand	30-32	3	49
α-helix	35-47	13
β-strand	54-60	7	49
β-strand	66-73	8	49
β-strand	77-79	3	50
β-strand	84-86	3	50
α-helix	90-100	11
β-strand	107-114	8	49
α-helix	130-136	7
α-helix	140	1
β-strand	141-146	6	49
β-strand	156-162	7	49
α-helix	187-188	2
α-helix	189-192	4
β-strand	200-202	3	49
β-strand	204-205	2	49
α-helix	211-218	8
α-helix	225-228	4
α-helix	234-261	28
α-helix	283-307	25

Chain C: 22 helices, 11 β-strands

Element	Residues	Length	Sheet
α-helix	22-25	4
α-helix	27-60	34
α-helix	62-65	4
β-strand	71-76	6	74
β-strand	80	1	74
β-strand	84-88	5	74
β-strand	94-98	5	74
β-strand	112-117	6	74
β-strand	122-126	5	74
α-helix	127-129	3
α-helix	133-136	4
α-helix	138-140	3
α-helix	145-147	3
α-helix	155-161	7
α-helix	162-166	5
α-helix	167-170	4
α-helix	173-178	6
α-helix	181-183	3
β-strand	185-189	5	89
α-helix	196-206	11
β-strand	210-213	4	89
α-helix	232-241	10
β-strand	244-248	5	89
α-helix	251-255	5
α-helix	259-261	3
α-helix	269-282	14
β-strand	290-295	6	89
α-helix	303-306	4
β-strand	314-316	3	89
α-helix	322-332	11
α-helix	344-349	6
α-helix	356-372	17
α-helix	380-393	14

Chain d: 13 helices, 2 β-strands

Element	Residues	Length	Sheet
α-helix	17-29	13
α-helix	42-45	4
α-helix	46-65	20
α-helix	68-82	15
α-helix	90-92	3
α-helix	95-107	13
α-helix	112-116	5
α-helix	131-142	12
α-helix	150-153	4
α-helix	164-183	20
α-helix	193-196	4
α-helix	205-212	8
β-strand	217	1	56
β-strand	222	1	56
α-helix	238-254	17

Chain D: 16 helices, 10 β-strands

Element	Residues	Length	Sheet
α-helix	40-79	40
α-helix	80-82	3
β-strand	87-97	11	74
β-strand	100-105	6	74
β-strand	110-115	6	74
α-helix	121-123	3
β-strand	129-132	4	74
β-strand	139-142	4	74
α-helix	164-166	3
α-helix	171-181	11
α-helix	183-185	3
α-helix	189-194	6
β-strand	201-205	5	93
α-helix	212-221	10
β-strand	226-230	5	93
α-helix	242-257	16
β-strand	260-265	6	93
α-helix	267-269	3
α-helix	282-297	16
β-strand	305-310	6	93
β-strand	327-332	6	93
α-helix	338-349	12
α-helix	361-363	3
α-helix	372-388	17
α-helix	396-403	8
α-helix	415-417	3

27 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	533	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	892	Homo sapiens	Q13200

22 more molecules are not listed.

Sequence of entity 1 (U), FASTA

>6MSE_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

>6MSE_2 26S proteasome non-ATPase regulatory subunit 3 (chains V)
KQEGSARRRGADKAKPPPGGGEQEPPPPPAPQDVEMKEEAATGGGSTGEADGKTAAAAAE
HSQRELDTVTLEDIKEHVKQLEKAVSGKEPRFVLRALRMLPSTSRRLNHYVLYKAVQGFF
TSNNATRDFLLPFLEEPMDTEADLQFRPRTGKAASTPLLPEVEAYLQLLVVIFMMNSKRY
KEAQKISDDLMQKISTQNRRALDLVAAKCYYYHARVYEFLDKLDVVRSFLHARLRTATLR
HDADGQATLLNLLLRNYLHYSLYDQAEKLVSKSVFPEQANNNEWARYLYYTGRIKAIQLE
YSEARRTMTNALRKAPQHTAVGFKQTVHKLLIVVELLLGEIPDRLQFRQPSLKRSLMPYF
LLTQAVRTGNLAKFNQVLDQFGEKFQADGTYTLIIRLRHNVIKTGVRMISLSYSRISLAD
IAQKLQLDSPEDAEFIVAKAIRDGVIEASINHEKGYVQSKEMIDIYSTREPQLAFHQRIS
FCLDIHNMSVKAMRFPPKSYNKDLESAEERREREQQDLEFAKEMAEDDDDSFP

Sequence of entity 3 (W), FASTA

>6MSE_3 26S proteasome non-ATPase regulatory subunit 12 (chains W)
MADGGSERADGRIVKMEVDYSATVDQRLPECAKLAKEGRLQEVIETLLSLEKQTRTASDM
VSTSRILVAVVKMCYEAKEWDLLNENIMLLSKRRSQLKQAVAKMVQQCCTYVEEITDLPI
KLRLIDTLRMVTEGKIYVEIERARLTKTLATIKEQNGDVKEAASILQELQVETYGSMEKK
ERVEFILEQMRLCLAVKDYIRTQIISKKINTKFFQEENTEKLKLKYYNLMIQLDQHEGSY
LSICKHYRAIYDTPCIQAESEKWQQALKSVVLYVILAPFDNEQSDLVHRISGDKKLEEIP
KYKDLLKLFTTMELMRWSTLVEDYGMELRKGSLESPATDVFGSTEEGEKRWKDLKNRVVE
HNIRIMAKYYTRITMKRMAQLLDLSVDESEAFLSNLVVNKTIFAKVDRLAGIINFQRPKD
PNNLLNDWSQKLNSLMSLVNKTTHLIAKEEMIHNLQ

Sequence of entity 4 (X), FASTA

>6MSE_4 26S proteasome non-ATPase regulatory subunit 11 (chains X)
MAAAAVVEFQRAQSLLSTDREASIDILHSIVKRDIQENDEEAVQVKEQSILELGSLLAKT
GQAAELGGLLKYVRPFLNSISKAKAARLVRSLLDLFLDMEAATGQEVELCLECIEWAKSE
KRTFLRQALEARLVSLYFDTKRYQEALHLGSQLLRELKKMDDKALLVEVQLLESKTYHAL
SNLPKARAALTSARTTANAIYCPPKLQATLDMQSGIIHAAEEKDWKTAYSYFYEAFEGYD
SIDSPKAITSLKYMLLCKIMLNTPEDVQALVSGKLALRYAGRQTEALKCVAQASKNRSLA
DFEKALTDYRAELRDDPIISTHLAKLYDNLLEQNLIRVIEPFSRVQIEHISSLIKLSKAD
VERKLSQMILDKKFHGILDQGEGVLIIFDEPPVDKTYEAALETIQNMSKVVDSLYNKAKK
LT

Sequence of entity 5 (Y), FASTA

>6MSE_5 26S proteasome non-ATPase regulatory subunit 6 (chains Y)
MPLENLEEEGLPKNPDLRIAQLRFLLSLPEHRGDAAVRDELMAAVRDNNMAPYYEALCKS
LDWQIDVDLLNKMKKANEDELKRLDEELEDAEKNLGESEIRDAMMAKAEYLCRIGDKEGA
LTAFRKTYDKTVALGHRLDIVFYLLRIGLFYMDNDLITRNTEKAKSLIEEGGDWDRRNRL
KVYQGLYCVAIRDFKQAAELFLDTVSTFTSYELMDYKTFVTYTVYVSMIALERPDLREKV
IKGAEILEVLHSLPAVRQYLFSLYECRYSVFFQSLAVVEQEMKKDWLFAPHYRYYVREMR
IHAYSQLLESYRSLTLGYMAEAFGVGVEFIDQELSRFIAAGRLHCKIDKVNEIVETNRPD
SKNWQYQETIKKGDLLLNRVQKLSRVINM

Sequence of entity 6 (Z), FASTA

>6MSE_6 26S proteasome non-ATPase regulatory subunit 7 (chains Z)
MPELAVQKVVVHPLVLLSVVDHFNRIGKVGNQKRVVGVLLGSWQKKVLDVSNSFAVPFDE
DDKDDSVWFLDHDYLENMYGMFKKVNARERIVGWYHTGPKLHKNDIAINELMKRYCPNSV
LVIIDVKPKDLGLPTEAYISVEEVHDDGTPTSKTFEHVTSEIGAEEAEEVGVEHLLRDIK
DTTVGTLSQRITNQVHGLKGLNSKLLDIRSYLEKVATGKLPINHQIIYQLQDVFNLLPDV
SLQEFVKAFYLKTNDQMVVVYLASLIRSVVALHNLINNKIANRDAEKKEGQEKEESKKDR
KEDKEKDKDKEKSDVKKEEKKEKK

Sequence of entity 7 (a), FASTA

>6MSE_7 26S proteasome non-ATPase regulatory subunit 13 (chains a)
MKDVPGFLQQSQNSGPGQPAVWHRLEELYTKKLWHQLTLQVLDFVQDPCFAQGDGLIKLY
ENFISEFEHRVNPLSLVEIILHVVRQMTDPNVALTFLEKTREKVKSSDEAVILCKTAIGA
LKLNIGDLQVTKETIEDVEEMLNNLPGVTSVHSRFYDLSSKYYQTIGNHASYYKDALRFL
GCVDIKDLPVSEQQERAFTLGLAGLLGEGVFNFGELLMHPVLESLRNTDRQWLIDTLYAF
NSGNVERFQTLKTAWGQQPDLAANEAQLLRKIQLLCLMEMTFTRPANHRQLTFEEIAKSA
KITVNEVELLVMKALSVGLVKGSIDEVDKRVHMTWVQPRVLDLQQIKGMKDRLEFWCTDV
KSMEMLVEHQAHDILT

Sequence of entity 8 (b), FASTA

>6MSE_8 26S proteasome non-ATPase regulatory subunit 4 (chains b)
MVLESTMVCVDNSEYMRNGDFLPTRLQAQQDAVNIVCHSKTRSNPENNVGLITLANDCEV
LTTLTPDTGRILSKLHTVQPKGKITFCTGIRVAHLALKHRQGKNHKMRIIAFVGSPVEDN
EKDLVKLAKRLKKEKVNVDIINFGEEEVNTEKLTAFVNTLNGKDGTGSHLVTVPPGPSLA
DALISSPILAGEGGAMLGLGASDFEFGVDPSADPELALALRVSMEEQRQRQEEEARRAAA
ASAAEAGIATTGTEDSDDALLKMTISQQEFGRTGLPDLSSMTEEEQIAYAMQMSLQGAEF
GQAESADIDASSAMDTSEPAKEEDDYDVMQDPEFLQSVLENLPGVDPNNEAIRNAMGSLA
SQATKDGKKDKKEEDKK

Sequence of entity 9 (c), FASTA

>6MSE_9 26S proteasome non-ATPase regulatory subunit 14 (chains c)
DRLLRLGGGMPGLGQGPPTDAPAVDTAEQVYISSLALLKMLKHGRAGVPMEVMGLMLGEF
VDDYTVRVIDVFAMPQSGTGVSVEAVDPVFQAKMLDMLKQTGRPEMVVGWYHSHPGFGCW
LSGVDINTQQSFEALSERAVAVVVDPIQSVKGKVVIDAFRLINANMMVLGHEPRQTTSNL
GHLNKPSIQALIHGLNRHYYSITINYRKNELEQKMLLNLHKKSWMEGLTLQDYSEHCKHN
ESVVKEMLELAKNYNKAVEEEDKMTPEQLAIKNVGKQDPKRHLEEHVDVLMTSNIVQCLA
AMLDTVVFK

Sequence of entity 10 (d), FASTA

>6MSE_10 26S proteasome non-ATPase regulatory subunit 8 (chains d)
FIKGRAPRAPPRERRRATRGGLRQVVAPPRALGSTSRPHFRRASVCRRRCRKSGGLLAAS
RKMAAAAVNGAAGFSSSGPAATSGAVLQAATGMYEQLKGEWNRKSPNLSKCGEELGRLKL
VLLELNFLPTTGTKLTKQQLILARDILEIGAQWSILRKDIPSFERYMAQLKCYYFDYKEQ
LPESAYMHQLLGLNLLFLLSQNRVAEFHTELERLPAKDIQTNVYIKHPVSLEQYLMEGSY
NKVFLAKGNIPAESYTFFIDILLDTIRDEIAGCIEKAYEKILFTEATRILFFNTPKKMTD
YAKKRGWVLGPNNYYSFASQQQKPEDTTIPSTELAKQVIEYARQLEMIV

Sequence of entity 11 (e), FASTA

>6MSE_11 26S proteasome complex subunit SEM1 (chains e)
MSEKKQPVDLGLLEEDDEFEEFPAEDWAGLDEDEDAHVWEDNWDDDNVEDDFSNQLRAEL
EKHGYKMETS

Sequence of entity 12 (f), FASTA

>6MSE_12 26S proteasome non-ATPase regulatory subunit 2 (chains f)
MEEGGRDKAPVQPQQSPAAAPGGTDEKPSGKERRDAGDKDKEQELSEEDKQLQDELEMLV
ERLGEKDTSLYRPALEELRRQIRSSTTSMTSVPKPLKFLRPHYGKLKEIYENMAPGENKR
FAADIISVLAMTMSGERECLKYRLVGSQEELASWGHEYVRHLAGEVAKEWQELDDAEKVQ
REPLLTLVKEIVPYNMAHNAEHEACDLLMEIEQVDMLEKDIDENAYAKVCLYLTSCVNYV
PEPENSALLRCALGVFRKFSRFPEALRLALMLNDMELVEDIFTSCKDVVVQKQMAFMLGR
HGVFLELSEDVEEYEDLTEIMSNVQLNSNFLALARELDIMEPKVPDDIYKTHLENNRFGG
SGSQVDSARMNLASSFVNGFVNAAFGQDKLLTDDGNKWLYKNKDHGMLSAAASLGMILLW
DVDGGLTQIDKYLYSSEDYIKSGALLACGIVNSGVRNECDPALALLSDYVLHNSNTMRLG
SIFGLGLAYAGSNREDVLTLLLPVMGDSKSSMEVAGVTALACGMIAVGSCNGDVTSTILQ
TIMEKSETELKDTYARWLPLGLGLNHLGKGEAIEAILAALEVVSEPFRSFANTLVDVCAY
AGSGNVLKVQQLLHICSEHFDSKEKEEDKDKKEKKDKDKKEAPADMGAHQGVAVLGIALI
AMGEEIGAEMALRTFGHLLRYGEPTLRRAVPLALALISVSNPRLNILDTLSKFSHDADPE
VSYNSIFAMGMVGSGTNNARLAAMLRQLAQYHAKDPNNLFMVRLAQGLTHLGKGTLTLCP
YHSDRQLMSQVAVAGLLTVLVSFLDVRNIILGKSHYVLYGLVAAMQPRMLVTFDEELRPL
PVSVRVGQAVDVVGQAGKPKTITGFQTHTTPVLLAHGERAELATEEFLPVTP

Ligands and cofactors

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

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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