9GEV: Human INO80 core-nucleosome complex state N-6

CryoEM structure of the human INO80 core-nucleosome complex state N-6. Determined by electron microscopy at 3.47 Å resolution. Released 20 Aug 2025.

Method
Electron microscopy
Resolution
3.47 Å
Organisms
Homo sapiens, synthetic construct
Chains
20
Atoms
42,875
Mol. weight
815.41 kDa
Ligands
ATP, ADP
Released
20 Aug 2025

Explore 9GEV in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9GEV contains 271 α-helices and 234 β-strands across 18 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: 27 helices, 26 β-strands

ElementResiduesLengthSheet
α-helix13-142
β-strand15134
β-strand26135
α-helix311
β-strand32135
α-helix331
β-strand35-36236
β-strand39-40236
α-helix43-5715
β-strand61137
β-strand65-69538
α-helix76-8712
β-strand93-97538
α-helix98-1014
α-helix108-11811
β-strand120-1402139
β-strand157-163739
β-strand166-172739
α-helix174-1829
β-strand189-194639
β-strand200-206739
α-helix207-2093
β-strand215140
β-strand220-221239
α-helix222-2243
β-strand229-2391139
α-helix240-2489
α-helix250-2512
α-helix260-2623
α-helix273-28715
β-strand293-296439
β-strand298-302538
α-helix304-3063
α-helix309-31810
β-strand326-331638
β-strand333141
α-helix3361
β-strand337142
α-helix3381
β-strand345142
α-helix347-3493
α-helix352-3554
β-strand359-362438
α-helix365-3673
α-helix368-38215
β-strand386143
α-helix388-40013
α-helix403-41917
β-strand425143
α-helix427-4348
β-strand438144
α-helix442-4498
α-helix451-4533
Chain B: 28 helices, 28 β-strands
ElementResiduesLengthSheet
β-strand3-5345
α-helix13-142
β-strand15115
β-strand26146
α-helix311
β-strand32146
α-helix331
β-strand35-36247
β-strand39-40247
α-helix43-5715
β-strand61148
β-strand65-69549
α-helix76-8611
β-strand93-97549
α-helix98-1014
α-helix108-11811
β-strand120-1432450
β-strand153-1631150
β-strand166-172750
α-helix174-1829
β-strand189-194650
β-strand199-205750
β-strand206151
α-helix207-2093
β-strand21516
β-strand221151
α-helix222-2243
β-strand229-2391150
α-helix240-2489
α-helix258-2614
α-helix273-28816
β-strand292-296550
β-strand298-302549
α-helix304-3063
α-helix309-31911
α-helix324-3252
β-strand326-331649
β-strand333152
α-helix3361
β-strand337153
α-helix3381
β-strand345153
α-helix347-3493
α-helix352-3554
β-strand359-362449
α-helix365-3673
α-helix368-38215
β-strand386154
α-helix388-40013
α-helix403-4086
α-helix410-41910
β-strand425154
α-helix427-43610
β-strand437-438216
α-helix440-44910
α-helix451-4533
Chain C: 31 helices, 27 β-strands
ElementResiduesLengthSheet
β-strand1511
β-strand2612
α-helix311
β-strand3212
α-helix331
β-strand35-3623
β-strand39-4023
α-helix43-5715
β-strand6114
β-strand65-6955
α-helix76-8712
β-strand93-9755
α-helix98-1014
α-helix108-11811
β-strand120-142236
α-helix1431
α-helix146-1472
β-strand152-163126
β-strand166-17276
α-helix174-1763
α-helix177-1804
β-strand189-19466
β-strand200-20566
β-strand20617
α-helix207-2093
β-strand21518
α-helix2201
β-strand22117
α-helix222-2243
β-strand229-239116
α-helix240-2489
α-helix256-2638
α-helix265-2662
α-helix273-28816
β-strand292-29656
β-strand298-30255
α-helix304-3063
α-helix309-32012
β-strand326-33165
β-strand33319
α-helix3361
β-strand337110
α-helix3381
β-strand345110
α-helix347-3493
α-helix352-3554
β-strand359-36245
α-helix365-3673
α-helix368-38215
β-strand386111
α-helix388-40013
α-helix403-4086
α-helix410-41910
β-strand425111
α-helix427-43610
β-strand438112
α-helix440-44910
α-helix451-4533
Chain D: 24 helices, 23 β-strands
ElementResiduesLengthSheet
β-strand22148
α-helix38-403
β-strand42-43255
β-strand46-47255
α-helix50-6516
β-strand68134
β-strand72-77644
α-helix83-9412
β-strand100-104544
α-helix105-1084
α-helix115-12511
β-strand127-1482240
β-strand158-164740
β-strand169-174640
α-helix1751
α-helix176-1827
β-strand191-196640
β-strand201-208840
α-helix209-2113
β-strand224-225240
α-helix226-2294
β-strand233-2431140
α-helix244-2507
α-helix258-2614
α-helix270-28516
β-strand290-293440
β-strand295-299544
α-helix301-3033
α-helix306-31510
β-strand323-328644
β-strand333-334256
α-helix3351
β-strand341-342256
α-helix343-3453
α-helix348-3514
β-strand355-359544
α-helix360-3634
α-helix364-37815
β-strand382157
α-helix384-39613
α-helix400-41617
β-strand421157
α-helix423-43210
β-strand434149
α-helix436-4416
α-helix444-4463
β-strand451152
Chain E: 21 helices, 27 β-strands
ElementResiduesLengthSheet
β-strand2214
β-strand33113
α-helix381
β-strand39113
α-helix401
β-strand42-43214
β-strand46-47214
α-helix50-6516
β-strand68115
β-strand72-76516
α-helix83-9311
β-strand100-104516
α-helix105-1084
α-helix115-12410
β-strand127-150246
β-strand158-16476
β-strand169-17466
α-helix176-1849
β-strand191-19666
β-strand20116
β-strand204-20746
β-strand208117
α-helix209-2124
β-strand225117
α-helix226-2294
β-strand233-243116
α-helix244-2529
α-helix259-2624
α-helix270-28516
β-strand290-29346
β-strand295-299516
α-helix301-3033
α-helix306-31510
β-strand323-328616
β-strand332-334318
α-helix3351
β-strand341-343318
α-helix348-3514
β-strand355-358416
α-helix364-37714
β-strand382119
α-helix384-39613
α-helix399-41517
β-strand421119
α-helix423-43210
β-strand43415
α-helix436-44510
β-strand45119
Chain F: 25 helices, 26 β-strands
ElementResiduesLengthSheet
α-helix16-183
β-strand22137
β-strand33158
α-helix381
β-strand39158
α-helix401
β-strand42-43259
β-strand46-47259
α-helix50-6516
β-strand6811
β-strand72-76512
α-helix83-9412
β-strand100-104512
α-helix105-1084
α-helix115-12410
β-strand127-146208
β-strand159-16578
β-strand168-17478
α-helix176-1849
β-strand191-19668
β-strand202-20878
α-helix209-2113
β-strand217139
β-strand224-22528
α-helix226-2283
β-strand233-243118
α-helix244-2529
α-helix258-2625
α-helix270-28617
β-strand289-29358
β-strand295-299512
α-helix301-3033
α-helix306-31510
β-strand323-328612
β-strand333-334260
α-helix3351
β-strand341-342260
α-helix343-3453
α-helix348-3514
β-strand354-358512
α-helix361-3633
α-helix364-37815
β-strand382161
α-helix384-39613
α-helix399-41517
β-strand421161
α-helix423-43210
β-strand434138
α-helix436-44510
α-helix447-4493
β-strand451141
Chain G: 47 helices, 22 β-strands
ElementResiduesLengthSheet
α-helix520-53516
β-strand539-540262
α-helix543-5442
α-helix551-5544
α-helix556-5594
α-helix564-5674
β-strand570-574562
α-helix576-5783
α-helix579-58911
β-strand595162
β-strand598162
α-helix602-6098
α-helix610-6123
β-strand626-630562
α-helix631-6344
α-helix638-6425
β-strand648-652562
α-helix654-6574
α-helix663-6686
β-strand675-679562
α-helix688-69811
α-helix708-7114
α-helix730-73910
β-strand743162
α-helix748-7503
α-helix756-7572
β-strand758-765845
α-helix769-77911
α-helix809-8179
α-helix819-8213
α-helix824-8263
β-strand828163
β-strand831164
α-helix835-8373
β-strand838-840365
α-helix842-8476
α-helix850-8534
α-helix855-8639
α-helix869-8768
α-helix889-8924
α-helix898-9069
α-helix908-93225
α-helix944-9452
α-helix959-9646
α-helix968-9714
β-strand980-990116
α-helix992-9943
β-strand996-1005106
α-helix1006-10072
β-strand1017164
α-helix1019-10213
β-strand1024-1026365
α-helix1029-104012
α-helix1044-10518
α-helix1055-106410
α-helix1070-10734
α-helix1076-10783
β-strand1089163
α-helix1094-10996
α-helix1101-111616
β-strand1120-1123445
α-helix1127-113913
β-strand1144-1147445
α-helix1153-116311
β-strand1171-1174445
β-strand1190-1193445
α-helix1202-12087
β-strand1219-1226845
α-helix1231-124717
Chain H: 8 helices, 7 β-strands
ElementResiduesLengthSheet
α-helix142-15817
α-helix173-1753
α-helix178-18811
α-helix213-23321
β-strand272-27766
β-strand282-28656
β-strand297-29826
α-helix300-3034
α-helix306-3072
β-strand308166
β-strand317166
α-helix3181
β-strand320-32238
β-strand328-32928
α-helix332-3409

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

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
RuvB-like 1A, B, Cprotein456Homo sapiensQ9Y265 (AlphaFold model)
RuvB-like 2D, E, Fprotein463Homo sapiensQ9Y230 (AlphaFold model)
INO80 complex subunit CIprotein192Homo sapiensQ6PI98 (AlphaFold model)
Nucleosomal DNA Strand 1KDNA152synthetic construct
Nucleosomal DNA Strand 2LDNA152synthetic construct
Histone H3.1M, Qprotein136Homo sapiensP68431 (AlphaFold model)
Histone H4N, Rprotein102Homo sapiensP62805
Histone H2A type 1-B/EO, Sprotein129Homo sapiensP04908
Histone H2B type 2-EP, Tprotein125Homo sapiensQ16778
Chromatin-remodeling ATPase INO80Gprotein1556Homo sapiensQ9ULG1
INO80 complex subunit BHprotein356Homo sapiensQ9C086
Actin-related protein 5Jprotein607Homo sapiensQ9H9F9
Sequence of entity 1 (A, B, C), FASTA
>9GEV_1 RuvB-like 1 (chains A, B, C)
MKIEEVKSTTKTQRIASHSHVKGLGLDESGLAKQAASGLVGQENAREACGVIVELIKSKK
MAGRAVLLAGPPGTGKTALALAIAQELGSKVPFCPMVGSEVYSTEIKKTEVLMENFRRAI
GLRIKETKEVYEGEVTELTPCETENPMGGYGKTISHVIIGLKTAKGTKQLKLDPSIFESL
QKERVEAGDVIYIEANSGAVKRQGRCDTYATEFDLEAEEYVPLPKGDVHKKKEIIQDVTL
HDLDVANARPQGGQDILSMMGQLMKPKKTEITDKLRGEINKVVNKYIDQGIAELVPGVLF
VDEVHMLDIECFTYLHRALESSIAPIVIFASNRGNCVIRGTEDITSPHGIPLDLLDRVMI
IRTMLYTPQEMKQIIKIRAQTEGINISEEALNHLGEIGTKTTLRYSVQLLTPANLLAKIN
GKDSIEKEHVEEISELFYDAKSSAKILADQQDKYMK
Sequence of entity 2 (D, E, F), FASTA
>9GEV_2 RuvB-like 2 (chains D, E, F)
MATVTATTKVPEIRDVTRIERIGAHSHIRGLGLDDALEPRQASQGMVGQLAARRAAGVVL
EMIREGKIAGRAVLIAGQPGTGKTAIAMGMAQALGPDTPFTAIAGSEIFSLEMSKTEALT
QAFRRSIGVRIKEETEIIEGEVVEIQIDRPATGTGSKVGKLTLKTTEMETIYDLGTKMIE
SLTKDKVQAGDVITIDKATGKISKLGRSFTRARDYDAMGSQTKFVQCPDGELQKRKEVVH
TVSLHEIDVINSRTQGFLALFSGDTGEIKSEVREQINAKVAEWREEGKAEIIPGVLFIDE
VHMLDIESFSFLNRALESDMAPVLIMATNRGITRIRGTSYQSPHGIPIDLLDRLLIVSTT
PYSEKDTKQILRIRCEEEDVEMSEDAYTVLTRIGLETSLRYAIQLITAASLVCRKRKGTE
VQVDDIKRVYSLFLDESRSTQYMKEYQDAFLFNELKGETMDTS
Sequence of entity 3 (I), FASTA
>9GEV_3 INO80 complex subunit C (chains I)
MAAQIPIVATTSTPGIVRNSKKRPASPSHNGSSGGGYGASKKKKASASSFAQGISMEAMS
ENKMVPSEFSTGPVEKAAKPLPFKDPNFVHSGHGGAVAGKKNRTWKYLKQILASERALPW
QLNDPNYFSIDAPPSFKPAKKYSDVSGLLANYTDPQSKLRFSTIEEFSYIRRLPSDVVTG
YLALRKATSIVP
Sequence of entity 4 (K), FASTA
>9GEV_4 Nucleosomal DNA Strand 1 (chains K)
CAATATCCCGAGTACATGCACAGGATGTATATATCTGACACGTGCCTGGAGACTAGGGAG
TAATCCCCTTGGCGGTTAAAACGCGGGGGACAGCGCGTACGTGCGTTTAAGCGGTGCTAG
AGCTGTCTACGACCAATTGAGCGGCCTCGGCA
Sequence of entity 5 (L), FASTA
>9GEV_5 Nucleosomal DNA Strand 2 (chains L)
TGCCGAGGCCGCTCAATTGGTCGTAGACAGCTCTAGCACCGCTTAAACGCACGTACGCGC
TGTCCCCCGCGTTTTAACCGCCAAGGGGATTACTCCCTAGTCTCCAGGCACGTGTCAGAT
ATATACATCCTGTGCATGTACTCGGGATATTG
Sequence of entity 6 (M, Q), FASTA
>9GEV_6 Histone H3.1 (chains M, Q)
MARTKQTARKSTGGKAPRKQLATKAARKSAPATGGVKKPHRYRPGTVALREIRRYQKSTE
LLIRKLPFQRLVREIAQDFKTDLRFQSSAVMALQEACEAYLVGLFEDTNLCAIHAKRVTI
MPKDIQLARRIRGERA
Sequence of entity 7 (N, R), FASTA
>9GEV_7 Histone H4 (chains N, R)
SGRGKGGKGLGKGGAKRHRKVLRDNIQGITKPAIRRLARRGGVKRISGLIYEETRGVLKV
FLENVIRDAVTYTEHAKRKTVTAMDVVYALKRQGRTLYGFGG
Sequence of entity 8 (O, S), FASTA
>9GEV_8 Histone H2A type 1-B/E (chains O, S)
SGRGKQGGKARAKAKTRSSRAGLQFPVGRVHRLLRKGNYSERVGAGAPVYLAAVLEYLTA
EILELAGNAARDNKKTRIIPRHLQLAIRNDEELNKLLGRVTIAQGGVLPNIQAVLLPKKT
ESHHKAKGK
Sequence of entity 9 (P, T), FASTA
>9GEV_9 Histone H2B type 2-E (chains P, T)
PEPAKSAPAPKKGSKKAVTKAQKKDGKKRKRSRKESYSIYVYKVLKQVHPDTGISSKAMG
IMNSFVNDIFERIAGEASRLAHYNKRSTITSREIQTAVRLLLPGELAKHAVSEGTKAVTK
YTSSK
Sequence of entity 10 (G), FASTA
>9GEV_10 Chromatin-remodeling ATPase INO80 (chains G)
MASELGARDDGGCTELAKPLYLQYLERALRLDHFLRQTSAIFNRNISSDDSEDGLDDSNP
LLPQSGDPLIQVKEEPPNSLLGETSGAGSSGMLNTYSLNGVLQSESKCDKGNLYNFSKLK
KSRKWLKSILLSDESSEADSQSEDDDEEELNLSREELHNMLRLHKYKKLHQNKYSKDKEL
QQYQYYSAGLLSTYDPFYEQQRHLLGPKKKKFKEEKKLKAKLKKVKKKRRRDEELSSEES
PRRHHHQTKVFAKFSHDAPPPGTKKKHLSIEQLNARRRKVWLSIVKKELPKANKQKASAR
NLFLTNSRKLAHQCMKEVRRAALQAQKNCKETLPRARRLTKEMLLYWKKYEKVEKEHRKR
AEKEALEQRKLDEEMREAKRQQRKLNFLITQTELYAHFMSRKRDMGHDGIQEEILRKLED
SSTQRQIDIGGGVVVNITQEDYDSNHFKAQALKNAENAYHIHQARTRSFDEDAKESRAAA
LRAANKSGTGFGESYSLANPSIRAGEDIPQPTIFNGKLKGYQLKGMNWLANLYEQGINGI
LADEMGLGKTVQSIALLAHLAERENIWGPFLIISPASTLNNWHQEFTRFVPKFKVLPYWG
NPHDRKVIRRFWSQKTLYTQDAPFHVVITSYQLVVQDVKYFQRVKWQYMVLDEAQALKSS
SSVRWKILLQFQCRNRLLLTGTPIQNTMAELWALLHFIMPTLFDSHEEFNEWFSKDIESH
AENKSAIDENQLSRLHMILKPFMLRRIKKDVENELSDKIEILMYCQLTSRQKLLYQALKN
KISIEDLLQSSMGSTQQAQNTTSSLMNLVMQFRKVCNHPELFERQETWSPFHISLKPYHI
SKFIYRHGQIRVFNHSRDRWLRVLSPFAPDYIQRSLFHRKGINEESCFSFLRFIDISPAE
MANLMLQGLLARWLALFLSLKASYRLHQLRSWGAPEGESHQRYLRNKDFLLGVNFPLSFP
NLCSCPLLKSLVFSSHCKAVSGYSDQVVHQRRSATSSLRRCLLTELPSFLCVASPRVTAV
PLDSYCNDRSAEYERRVLKEGGSLAAKQCLLNGAPELAADWLNRRSQFFPEPAGGLWSIR
PQNGWSFIRIPGKESLITDSGKLYALDVLLTRLKSQGHRVLIYSQMTRMIDLLEEYMVYR
KHTYMRLDGSSKISERRDMVADFQNRNDIFVFLLSTRAGGLGINLTAADTVIFYDSDWNP
TVDQQAMDRAHRLGQTKQVTVYRLICKGTIEERILQRAKEKSEIQRMVISGGNFKPDTLK
PKEVVSLLLDDEELEKKLRLRQEEKRQQEETNRVKERKRKREKYAEKKKKEDELDGKRRK
EGVNLVIPFVPSADNSNLSADGDDSFISVDSAMPSPFSEISISSELHTGSIPLDESSSDM
LVIVDDPASSAPQSRATNSPASITGSVSDTVNGISIQEMPAAGRGHSARSRGRPKGSGST
AKGAGKGRSRKSTAGSAAAMAGAKAGAAAASAAAYAAYGYNVSKGISASSPLQTSLVRPA
GLADFGPSSASSPLSSPLSKGNNVPGNPKNLHMTSSLAPDSLVRKQGKGTNPSGGR
Sequence of entity 11 (H), FASTA
>9GEV_11 INO80 complex subunit B (chains H)
MSKLWRRGSTSGAMEAPEPGEALELSLAGAHGHGVHKKKHKKHKKKHKKKHHQEEDAGPT
QPSPAKPQLKLKIKLGGQVLGTKSVPTFTVIPEGPRSPSPLMVVDNEEEPMEGVPLEQYR
AWLDEDSNLSPSPLRDLSGGLGGQEEEEEQRWLDALEKGELDDNGDLKKEINERLLTARQ
RALLQKARSQPSPMLPLPVAEGCPPPALTEEMLLKREERARKRRLQAARRAEEHKNQTIE
RLTKTAATSGRGGRGGARGERRGGRAAAPAPMVRYCSGAQGSTLSFPPGVPAPTAVSQRP
SPSGPPPRCSVPGCPHPRRYACSRTGQALCSLQCYRINLQMRLGGPEGPGSPLLAT
Sequence of entity 12 (J), FASTA
>9GEV_12 Actin-related protein 5 (chains J)
MAANVFPFRDARAAPDPVLEAGPVAHGPLPVPLVLDNGSFQVRAGWACPGQDPGPEPRLQ
FRAVCARGRGGARGASGPQVGNALGSLEPLRWMLRSPFDRNVPVNLELQELLLDYSFQHL
GVSSQGCVDHPIVLTEAVCNPLYSRQMMSELLFECYGIPKVAYGIDSLFSFYHNKPKNSM
CSGLIISSGYQCTHVLPILEGRLDAKNCKRINLGGSQAAGYLQRLLQLKYPGHLAAITLS
RMEEILHEHSYIAEDYVEELHKWRCPDYYENNVHKMQLPFSSKLLGSTLTSEEKQERRQQ
QLRRLQELNARRREEKLQLDQERLDRLLYVQELLEDGQMDQFHKALIELNMDSPEELQSY
IQKLSIAVEQAKQKILQAEVNLEVDVVDSKPETPDLEQLEPSLEDVESMNDFDPLFSEET
PGVEKPVTTVQPVFNLAAYHQLFVGTERIRAPEIIFQPSLIGEEQAGIAETLQYILDRYP
KDIQEMLVQNVFLTGGNTMYPGMKARMEKELLEMRPFRSSFQVQLASNPVLDAWYGARDW
ALNHLDDNEVWITRKEYEEKGGEYLKEHCASNIYVPIRLPKQASRSSDAQASSKGSAAGG
GGAGEQA

Ligands and cofactors

IDNameFormulaCopies
ATPAdenosine-5'-triphosphateC10 H16 N5 O13 P31
ADPAdenosine-5'-diphosphateC10 H15 N5 O10 P26

Primary citation

Recognition and remodelling of nucleosomes and hexasomes by the human INO80 complex. Aggarwal, P., Sharma, M., Woike, S. et al. Nucleic Acids Res (2026) 54. DOI 10.1093/nar/gkag138 · PubMed

Other PDB entries of the same protein (UniProt Q9Y265 (AlphaFold model), which also has an AlphaFold model), best resolution first:

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