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

Cryo-EM structures and dynamics of substrate-engaged human 26S proteasome. Determined by electron microscopy at 3.2 Å resolution. Released 21 Nov 2018.

Method
Electron microscopy
Resolution
3.2 Å
Organisms
Homo sapiens, Saccharomyces cerevisiae RM11-1a
Chains
47
Atoms
105,222
Mol. weight
1665.24 kDa
Ligands
ADP, MG, ATP, ZN
Released
21 Nov 2018

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

Secondary structure: helices and β-sheets

6MSK contains 601 α-helices and 531 β-strands across 47 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: 23 helices, 3 β-strands

ElementResiduesLengthSheet
α-helix5-106
α-helix18-2811
α-helix36-4611
α-helix52-543
α-helix55-6612
α-helix73-8715
α-helix90-10314
α-helix111-12010
α-helix131-14111
α-helix150-16011
α-helix161-1633
α-helix170-18213
α-helix190-20617
α-helix220-2234
α-helix231-24212
α-helix249-2568
α-helix262-2643
α-helix265-27511
α-helix277-2837
β-strand291-292216
α-helix293-2997
α-helix304-3063
α-helix307-31711
β-strand323-324216
β-strand330-332316
α-helix344-37027
Chain A: 17 helices, 15 β-strands
ElementResiduesLengthSheet
α-helix27-337
α-helix42-6322
α-helix81-855
α-helix86-883
β-strand97-98268
β-strand100-103469
β-strand111-114469
β-strand121-123369
β-strand125168
β-strand140-142368
β-strand149-152468
α-helix154-1585
α-helix161-1644
β-strand165170
α-helix174-1763
α-helix183-1908
α-helix192-1965
β-strand211-215570
α-helix222-2298
β-strand236-239470
α-helix242-2443
α-helix252-26615
β-strand270-273470
α-helix293-30412
β-strand315-320670
β-strand339-342470
α-helix349-35810
β-strand364171
α-helix371-3755
α-helix382-39817
β-strand404171
α-helix406-41611
Chain b: 7 helices, 8 β-strands
ElementResiduesLengthSheet
β-strand4-10744
β-strand13145
α-helix14-163
α-helix25-4319
β-strand48-53644
β-strand62-65444
α-helix69-746
β-strand82145
α-helix87-9711
β-strand106-113844
α-helix121-13414
β-strand137-142644
α-helix151-1588
β-strand171144
α-helix181-1855
Chain B: 15 helices, 17 β-strands
ElementResiduesLengthSheet
α-helix33-353
α-helix59-8224
α-helix95-1028
β-strand107-114875
β-strand120-124575
β-strand131168
β-strand132-134375
β-strand135176
β-strand149-152475
β-strand159176
β-strand162175
β-strand175177
α-helix191-20010
α-helix202-2065
α-helix216-2194
β-strand221-225577
α-helix232-24110
β-strand247-251577
α-helix262-27615
β-strand281-285577
α-helix287-2893
α-helix301-31717
β-strand326-330577
α-helix339-3424
β-strand349-352477
α-helix361-3677
β-strand374178
α-helix381-3866
α-helix392-40817
β-strand414178
α-helix416-42712
β-strand439179
Chain c: 12 helices, 8 β-strands
ElementResiduesLengthSheet
β-strand30-33450
α-helix35-4713
β-strand54-61850
β-strand66-74950
α-helix76-783
α-helix84-863
α-helix89-10113
β-strand108-114750
α-helix124-13613
β-strand141-145550
β-strand157-160450
α-helix189-1924
β-strand202150
β-strand205-207350
α-helix214-2218
α-helix226-2294
α-helix238-25922
α-helix267-2726
α-helix279-2835
α-helix286-30823
Chain C: 18 helices, 16 β-strands
ElementResiduesLengthSheet
α-helix14-174
α-helix22-6645
α-helix69-702
β-strand71-76675
β-strand84183
β-strand87-88275
β-strand94-96375
β-strand98183
β-strand112-117675
β-strand122183
β-strand123-127575
α-helix155-1617
α-helix162-1665
α-helix167-1704
α-helix173-1786
α-helix181-1833
β-strand185-188484
α-helix198-2069
β-strand210-215684
β-strand223185
α-helix226-23712
β-strand244-249684
α-helix267-27913
β-strand289-294684
α-helix298-3003
β-strand313-315384
α-helix318-3214
α-helix322-33211
β-strand338186
α-helix344-3485
α-helix359-37113
β-strand378186
α-helix380-39011
α-helix400-4045
Chain d: 14 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix17-2913
α-helix46-6520
α-helix68-8114
α-helix95-10814
α-helix112-1187
α-helix119-1213
α-helix125-1284
α-helix131-14414
α-helix149-1535
α-helix164-18219
α-helix191-1933
α-helix194-1985
α-helix205-2117
α-helix239-25315
Chain D: 13 helices, 12 β-strands
ElementResiduesLengthSheet
α-helix40-7940
β-strand88-971090
β-strand100-105690
β-strand111-112275
β-strand113-114290
β-strand129-132490
β-strand139-142490
α-helix143-1453
β-strand153-154291
α-helix164-1663
α-helix171-18010
α-helix182-1865
α-helix189-1946
β-strand201-205591
α-helix213-2219
β-strand226-230591
α-helix242-25514
β-strand260-264591
α-helix283-29412
β-strand305-311791
β-strand329-332491
α-helix341-35010
α-helix372-38716
α-helix399-4068
α-helix413-4153

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

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
26S proteasome non-ATPase regulatory subunit 1Uprotein953Homo sapiensQ99460 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 3Vprotein533Homo sapiensO43242 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 12Wprotein456Homo sapiensO00232 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 11Xprotein422Homo sapiensO00231 (AlphaFold model)
26S proteasome non-ATPase regulatory subunit 6Yprotein389Homo sapiensQ15008
26S proteasome non-ATPase regulatory subunit 7Zprotein324Homo sapiensP51665
26S proteasome non-ATPase regulatory subunit 13aprotein376Homo sapiensQ9UNM6
26S proteasome non-ATPase regulatory subunit 4bprotein377Homo sapiensP55036
26S proteasome non-ATPase regulatory subunit 14cprotein309Homo sapiensO00487
26S proteasome non-ATPase regulatory subunit 8dprotein349Homo sapiensP48556
26S proteasome complex subunit SEM1eprotein70Homo sapiensP60896
26S proteasome non-ATPase regulatory subunit 2fprotein892Homo sapiensQ13200

21 more molecules are not listed.

Sequence of entity 1 (U), FASTA
>6MSK_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
>6MSK_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
>6MSK_3 26S proteasome non-ATPase regulatory subunit 12 (chains W)
MADGGSERADGRIVKMEVDYSATVDQRLPECAKLAKEGRLQEVIETLLSLEKQTRTASDM
VSTSRILVAVVKMCYEAKEWDLLNENIMLLSKRRSQLKQAVAKMVQQCCTYVEEITDLPI
KLRLIDTLRMVTEGKIYVEIERARLTKTLATIKEQNGDVKEAASILQELQVETYGSMEKK
ERVEFILEQMRLCLAVKDYIRTQIISKKINTKFFQEENTEKLKLKYYNLMIQLDQHEGSY
LSICKHYRAIYDTPCIQAESEKWQQALKSVVLYVILAPFDNEQSDLVHRISGDKKLEEIP
KYKDLLKLFTTMELMRWSTLVEDYGMELRKGSLESPATDVFGSTEEGEKRWKDLKNRVVE
HNIRIMAKYYTRITMKRMAQLLDLSVDESEAFLSNLVVNKTIFAKVDRLAGIINFQRPKD
PNNLLNDWSQKLNSLMSLVNKTTHLIAKEEMIHNLQ
Sequence of entity 4 (X), FASTA
>6MSK_4 26S proteasome non-ATPase regulatory subunit 11 (chains X)
MAAAAVVEFQRAQSLLSTDREASIDILHSIVKRDIQENDEEAVQVKEQSILELGSLLAKT
GQAAELGGLLKYVRPFLNSISKAKAARLVRSLLDLFLDMEAATGQEVELCLECIEWAKSE
KRTFLRQALEARLVSLYFDTKRYQEALHLGSQLLRELKKMDDKALLVEVQLLESKTYHAL
SNLPKARAALTSARTTANAIYCPPKLQATLDMQSGIIHAAEEKDWKTAYSYFYEAFEGYD
SIDSPKAITSLKYMLLCKIMLNTPEDVQALVSGKLALRYAGRQTEALKCVAQASKNRSLA
DFEKALTDYRAELRDDPIISTHLAKLYDNLLEQNLIRVIEPFSRVQIEHISSLIKLSKAD
VERKLSQMILDKKFHGILDQGEGVLIIFDEPPVDKTYEAALETIQNMSKVVDSLYNKAKK
LT
Sequence of entity 5 (Y), FASTA
>6MSK_5 26S proteasome non-ATPase regulatory subunit 6 (chains Y)
MPLENLEEEGLPKNPDLRIAQLRFLLSLPEHRGDAAVRDELMAAVRDNNMAPYYEALCKS
LDWQIDVDLLNKMKKANEDELKRLDEELEDAEKNLGESEIRDAMMAKAEYLCRIGDKEGA
LTAFRKTYDKTVALGHRLDIVFYLLRIGLFYMDNDLITRNTEKAKSLIEEGGDWDRRNRL
KVYQGLYCVAIRDFKQAAELFLDTVSTFTSYELMDYKTFVTYTVYVSMIALERPDLREKV
IKGAEILEVLHSLPAVRQYLFSLYECRYSVFFQSLAVVEQEMKKDWLFAPHYRYYVREMR
IHAYSQLLESYRSLTLGYMAEAFGVGVEFIDQELSRFIAAGRLHCKIDKVNEIVETNRPD
SKNWQYQETIKKGDLLLNRVQKLSRVINM
Sequence of entity 6 (Z), FASTA
>6MSK_6 26S proteasome non-ATPase regulatory subunit 7 (chains Z)
MPELAVQKVVVHPLVLLSVVDHFNRIGKVGNQKRVVGVLLGSWQKKVLDVSNSFAVPFDE
DDKDDSVWFLDHDYLENMYGMFKKVNARERIVGWYHTGPKLHKNDIAINELMKRYCPNSV
LVIIDVKPKDLGLPTEAYISVEEVHDDGTPTSKTFEHVTSEIGAEEAEEVGVEHLLRDIK
DTTVGTLSQRITNQVHGLKGLNSKLLDIRSYLEKVATGKLPINHQIIYQLQDVFNLLPDV
SLQEFVKAFYLKTNDQMVVVYLASLIRSVVALHNLINNKIANRDAEKKEGQEKEESKKDR
KEDKEKDKDKEKSDVKKEEKKEKK
Sequence of entity 7 (a), FASTA
>6MSK_7 26S proteasome non-ATPase regulatory subunit 13 (chains a)
MKDVPGFLQQSQNSGPGQPAVWHRLEELYTKKLWHQLTLQVLDFVQDPCFAQGDGLIKLY
ENFISEFEHRVNPLSLVEIILHVVRQMTDPNVALTFLEKTREKVKSSDEAVILCKTAIGA
LKLNIGDLQVTKETIEDVEEMLNNLPGVTSVHSRFYDLSSKYYQTIGNHASYYKDALRFL
GCVDIKDLPVSEQQERAFTLGLAGLLGEGVFNFGELLMHPVLESLRNTDRQWLIDTLYAF
NSGNVERFQTLKTAWGQQPDLAANEAQLLRKIQLLCLMEMTFTRPANHRQLTFEEIAKSA
KITVNEVELLVMKALSVGLVKGSIDEVDKRVHMTWVQPRVLDLQQIKGMKDRLEFWCTDV
KSMEMLVEHQAHDILT
Sequence of entity 8 (b), FASTA
>6MSK_8 26S proteasome non-ATPase regulatory subunit 4 (chains b)
MVLESTMVCVDNSEYMRNGDFLPTRLQAQQDAVNIVCHSKTRSNPENNVGLITLANDCEV
LTTLTPDTGRILSKLHTVQPKGKITFCTGIRVAHLALKHRQGKNHKMRIIAFVGSPVEDN
EKDLVKLAKRLKKEKVNVDIINFGEEEVNTEKLTAFVNTLNGKDGTGSHLVTVPPGPSLA
DALISSPILAGEGGAMLGLGASDFEFGVDPSADPELALALRVSMEEQRQRQEEEARRAAA
ASAAEAGIATTGTEDSDDALLKMTISQQEFGRTGLPDLSSMTEEEQIAYAMQMSLQGAEF
GQAESADIDASSAMDTSEPAKEEDDYDVMQDPEFLQSVLENLPGVDPNNEAIRNAMGSLA
SQATKDGKKDKKEEDKK
Sequence of entity 9 (c), FASTA
>6MSK_9 26S proteasome non-ATPase regulatory subunit 14 (chains c)
DRLLRLGGGMPGLGQGPPTDAPAVDTAEQVYISSLALLKMLKHGRAGVPMEVMGLMLGEF
VDDYTVRVIDVFAMPQSGTGVSVEAVDPVFQAKMLDMLKQTGRPEMVVGWYHSHPGFGCW
LSGVDINTQQSFEALSERAVAVVVDPIQSVKGKVVIDAFRLINANMMVLGHEPRQTTSNL
GHLNKPSIQALIHGLNRHYYSITINYRKNELEQKMLLNLHKKSWMEGLTLQDYSEHCKHN
ESVVKEMLELAKNYNKAVEEEDKMTPEQLAIKNVGKQDPKRHLEEHVDVLMTSNIVQCLA
AMLDTVVFK
Sequence of entity 10 (d), FASTA
>6MSK_10 26S proteasome non-ATPase regulatory subunit 8 (chains d)
FIKGRAPRAPPRERRRATRGGLRQVVAPPRALGSTSRPHFRRASVCRRRCRKSGGLLAAS
RKMAAAAVNGAAGFSSSGPAATSGAVLQAATGMYEQLKGEWNRKSPNLSKCGEELGRLKL
VLLELNFLPTTGTKLTKQQLILARDILEIGAQWSILRKDIPSFERYMAQLKCYYFDYKEQ
LPESAYMHQLLGLNLLFLLSQNRVAEFHTELERLPAKDIQTNVYIKHPVSLEQYLMEGSY
NKVFLAKGNIPAESYTFFIDILLDTIRDEIAGCIEKAYEKILFTEATRILFFNTPKKMTD
YAKKRGWVLGPNNYYSFASQQQKPEDTTIPSTELAKQVIEYARQLEMIV
Sequence of entity 11 (e), FASTA
>6MSK_11 26S proteasome complex subunit SEM1 (chains e)
MSEKKQPVDLGLLEEDDEFEEFPAEDWAGLDEDEDAHVWEDNWDDDNVEDDFSNQLRAEL
EKHGYKMETS
Sequence of entity 12 (f), FASTA
>6MSK_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

IDNameFormulaCopies
ADPAdenosine-5'-diphosphateC10 H15 N5 O10 P21
MGMagnesium ionMg5
ATPAdenosine-5'-triphosphateC10 H16 N5 O13 P34
ZNZinc ionZn1

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:

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