9XYU: NER complex - C7CAD.ATP

NER complex - C7CAD.ATP. Determined by electron microscopy at 3.5 Å resolution. Released 18 Feb 2026.

Method
Electron microscopy
Resolution
3.5 Å
Organisms
Homo sapiens, synthetic construct
Chains
11
Atoms
23,406
Mol. weight
574.71 kDa
Ligands
ZN, SF4
Released
18 Feb 2026

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

Secondary structure: helices and β-sheets

9XYU contains 140 α-helices and 122 β-strands across 9 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: 26 helices, 33 β-strands

ElementResiduesLengthSheet
β-strand59-6021
β-strand76-7832
β-strand84-8742
α-helix93-10311
β-strand106-10832
β-strand113-11642
α-helix120-1289
α-helix133-14210
α-helix150-16011
β-strand166-17163
β-strand174-17963
α-helix182-1898
α-helix192-1976
β-strand20613
β-strand268-27143
α-helix277-28610
β-strand291-29223
β-strand294-29524
α-helix318-32710
β-strand32814
β-strand33015
β-strand333-33424
β-strand336-33946
α-helix346-35712
β-strand361-36556
α-helix368-38114
β-strand389-39246
β-strand405-40956
α-helix421-43212
β-strand43517
β-strand438-44146
α-helix443-4453
α-helix449-4579
β-strand46017
β-strand463-46756
α-helix477-4793
α-helix480-4834
β-strand487-49046
α-helix493-4986
β-strand505-50738
β-strand50819
β-strand511-512210
α-helix513-5153
α-helix516-5249
α-helix531-5355
α-helix538-55316
β-strand558-56258
α-helix565-57511
β-strand579-58028
α-helix586-59712
β-strand604-60748
β-strand622-62658
α-helix633-64311
β-strand657-66268
β-strand663-664210
α-helix668-6769
α-helix678-6825
β-strand68819
α-helix707-71711
Chain B: 36 helices, 24 β-strands
ElementResiduesLengthSheet
β-strand2-4311
β-strand9-11311
α-helix19-3416
β-strand38-41412
α-helix48-6215
β-strand69-74612
α-helix77-9822
β-strand106-108312
α-helix112-1154
α-helix127-13711
α-helix146-1483
α-helix156-1649
α-helix177-18711
α-helix191-1988
β-strand204-208512
α-helix209-2135
α-helix215-2217
β-strand229-234612
α-helix239-2479
β-strand249-252413
α-helix253-28331
α-helix322-34524
β-strand351-352213
α-helix356-36510
α-helix369-3735
α-helix375-38511
α-helix395-40915
β-strand414114
β-strand417-418213
β-strand421115
β-strand429115
β-strand432-435413
β-strand437114
α-helix440-4434
α-helix445-4506
β-strand453-458612
α-helix467-4704
β-strand479-480212
β-strand490-495616
α-helix508-5103
α-helix514-52815
β-strand535-539516
α-helix542-55413
α-helix558-5647
β-strand566-569416
α-helix574-58916
β-strand594-599616
α-helix603-6064
β-strand617-621516
α-helix631-64212
α-helix648-66518
β-strand675-680616
α-helix682-6854
α-helix687-6904
α-helix695-6984
α-helix704-7063
β-strand708116
α-helix710-72415
α-helix727-7282
α-helix730-7356
Chain C: 9 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix399-41214
α-helix424-43411
α-helix455-47824
α-helix4791
α-helix4811
α-helix484-50017
α-helix501-5055
α-helix506-51510
α-helix522-54625
Chain D: 24 helices, 18 β-strands
ElementResiduesLengthSheet
α-helix19-257
α-helix28-358
α-helix38-4710
α-helix50-589
β-strand66-67217
α-helix68-747
β-strand75118
α-helix77-793
α-helix80-9314
β-strand96-100517
β-strand106-110517
α-helix112-12211
α-helix145-16420
α-helix173-1819
β-strand185-186219
α-helix193-1942
β-strand195-196219
α-helix198-2058
α-helix208-22114
α-helix230-24112
α-helix243-2442
β-strand249-250220
α-helix256-26813
β-strand271-272221
β-strand280-281220
β-strand282-283221
α-helix285-2917
β-strand307-30931
β-strand314-31741
α-helix322-33110
β-strand333-33751
β-strand342-34541
α-helix348-3569
α-helix361-37010
β-strand37211
α-helix374-3774
α-helix385-39814
β-strand402-408722
α-helix415-42814
β-strand431-435522
β-strand440-443422
α-helix445-46117
Chain E: 19 helices, 20 β-strands
ElementResiduesLengthSheet
β-strand1415
α-helix15-162
α-helix18-214
α-helix30-4112
β-strand59-66823
α-helix69-724
α-helix80-9819
β-strand103-110823
β-strand113-121923
α-helix124-13512
α-helix145-15713
β-strand164-171823
α-helix182-19110
β-strand195-200623
α-helix206-21510
β-strand219-221323
α-helix225-23410
α-helix238-2403
β-strand250-253424
β-strand256-258325
β-strand270126
α-helix273-2775
α-helix281-2822
β-strand285126
β-strand288-290325
β-strand297-298225
β-strand303-304224
β-strand311-312224
α-helix315-3195
α-helix322-3254
α-helix327-3315
β-strand332-335427
β-strand344128
β-strand351128
β-strand356-359427
β-strand366-367227
α-helix369-3746
α-helix375-3795
α-helix383-3864
Chain F: 13 helices, 14 β-strands
ElementResiduesLengthSheet
β-strand7-8229
β-strand10-16730
α-helix19-2810
α-helix34-5118
β-strand56-62730
β-strand67-69330
α-helix102-12019
α-helix133-15018
β-strand156-157229
β-strand160-164530
α-helix171-1733
α-helix174-18613
β-strand190-195630
α-helix201-21010
β-strand214-216330
α-helix220-2223
α-helix223-2308
α-helix235-2406
β-strand241118
α-helix244-2485
β-strand249125
β-strand253-254231
α-helix2551
β-strand261-262231
β-strand264-267424
β-strand274-275224
α-helix284-2874
Chain G: 3 helices, 3 β-strands
ElementResiduesLengthSheet
β-strand5-11722
α-helix14-2613
β-strand36-37222
β-strand42-45422
α-helix47-493
α-helix50-6314
Chain H: 3 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix890-90112
α-helix904-91310
α-helix927-9337

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

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
TFIIH basal transcription factor complex helicase XPB subunitAprotein782Homo sapiensP19447 (AlphaFold model)
General transcription and DNA repair factor IIH helicase subunit XPDBprotein768Homo sapiensP18074 (AlphaFold model)
General transcription factor IIH subunit 1Cprotein548Homo sapiensP32780 (AlphaFold model)
General transcription factor IIH subunit 4, p52Dprotein462Homo sapiensQ92759 (AlphaFold model)
General transcription factor IIH subunit 2Eprotein395Homo sapiensQ13888
General transcription factor IIH subunit 3Fprotein308Homo sapiensQ13889
General transcription factor IIH subunit 5Gprotein71Homo sapiensQ6ZYL4
DNA repair protein complementing XP-C cellsHprotein940Homo sapiensQ01831
DNA repair protein complementing XP-A cellsKprotein273Homo sapiensP23025
DNA (CY5)LDNA93synthetic construct
DNAMDNA94synthetic construct
Sequence of entity 1 (A), FASTA
>9XYU_1 TFIIH basal transcription factor complex helicase XPB subunit (chains A)
MGKRDRADRDKKKSRKRHYEDEEDDEEDAPGNDPQEAVPSAAGKQVDESGTKVDEYGAKD
YRLQMPLKDDHTSRPLWVAPDGHIFLEAFSPVYKYAQDFLVAIAEPVCRPTHVHEYKLTA
YSLYAAVSVGLQTSDITEYLRKLSKTGVPDGIMQFIKLCTVSYGKVKLVLKHNRYFVESC
HPDVIQHLLQDPVIRECRLRNSEGEATELITETFTSKSAISKTAESSGGPSTSRVTDPQG
KSDIPMDLFDFYEQMDKDEEEEEETQTVSFEVKQEMIEELQKRCIHLEYPLLAEYDFRND
SVNPDINIDLKPTAVLRPYQEKSLRKMFGNGRARSGVIVLPCGAGKSLVGVTAACTVRKR
CLVLGNSAVSVEQWKAQFKMWSTIDDSQICRFTSDAKDKPIGCSVAISTYSMLGHTTKRS
WEAERVMEWLKTQEWGLMILDEVHTIPAKMFRRVLTIVQAHCKLGLTATLVREDDKIVDL
NFLIGPKLYEANWMELQNNGYIAKVQCAEVWCPMSPEFYREYVAIKTKKRILLYTMNPNK
FRACQFLIKFHERRNDKIIVFADNVFALKEYAIRLNKPYIYGPTSQGERMQILQNFKHNP
KINTIFISKVGDTSFDLPEANVLIQISSHGGSRRQEAQRLGRVLRAKKGMVAEEYNAFFY
SLVSQDTQEMAYSTKRQRFLVDQGYSFKVITKLAGMEEEDLAFSTKEEQQQLLQKVLAAT
DLDAEEEVVAGEFGSRSSQASRRFGTMSSMSGADDTVYMEYHSSRSKAPSKHVHPLFKRF
RK
Sequence of entity 2 (B), FASTA
>9XYU_2 General transcription and DNA repair factor IIH helicase subunit XPD (chains B)
MKLNVDGLLVYFPYDYIYPEQFSYMRELKRTLDAKGHGVLEMPSGTGKTVSLLALIMAYQ
RAYPLEVTKLIYCSRTVPEIEKVIEELRKLLNFYEKQEGEKLPFLGLALSSRKNLCIHPE
VTPLRFGKDVDGKCHSLTASYVRAQYQHDTSLPHCRFYEEFDAHGREVPLPAGIYNLDDL
KALGRRQGWCPYFLARYSILHANVVVYSYHYLLDPKIADLVSKELARKAVVVFDEAHNID
NVCIDSMSVNLTRRTLDRCQGNLETLQKTVLRIKETDEQRLRDEYRRLVEGLREASAARE
TDAHLANPVLPDEVLQEAVPGSIRTAEHFLGFLRRLLEYVKWRLRVQHVVQESPPAFLSG
LAQRVCIQRKPLRFCAERLRSLLHTLEITDLADFSPLTLLANFATLVSTYAKGFTIIIEP
FDDRTPTIANPILHFSCMDASLAIKPVFERFQSVIITSGTLSPLDIYPKILDFHPVTMAT
FTMTLARVCLCPMIIGRGNDQVAISSKFETREDIAVIRNYGNLLLEMSAVVPDGIVAFFT
SYQYMESTVASWYEQGILENIQRNKLLFIETQDGAETSVALEKYQEACENGRGAILLSVA
RGKVSEGIDFVHHYGRAVIMFGVPYVYTQSRILKARLEYLRDQFQIRENDFLTFDAMRHA
AQCVGRAIRGKTDYGLMVFADKRFARGDKRGKLPRWIQEHLTDANLNLTVDEGVQVAKYF
LRQMAQPFHREDQLGLSLLSLEQLESEETLKRIEQIAQQLDYKDDDDK
Sequence of entity 3 (C), FASTA
>9XYU_3 General transcription factor IIH subunit 1 (chains C)
MATSSEEVLLIVKKVRQKKQDGALYLMAERIAWAPEGKDRFTISHMYADIKCQKISPEGK
AKIQLQLVLHAGDTTNFHFSNESTAVKERDAVKDLLQQLLPKFKRKANKELEEKNRMLQE
DPVLFQLYKDLVVSQVISAEEFWANRLNVNATDSSSTSNHKQDVGISAAFLADVRPQTDG
CNGLRYNLTSDIIESIFRTYPAVKMKYAENVPHNMTEKEFWTRFFQSHYFHRDRLNTGSK
DLFAECAKIDEKGLKTMVSLGVKNPLLDLTALEDKPLDEGYGISSVPSASNSKSIKENSN
AAIIKRFNHHSAMVLAAGLRKQEAQNEQTSEPSNMDGNSGDADCFQPAVKRAKLQESIEY
EDLGKNNSVKTIALNLKKSDRYYHGPTPIQSLQYATSQDIINSFQSIRQEMEAYTPKLTQ
VLSSSAASSTITALSPGGALMQGGTQQAINQMVPNDIQSELKHLYVAVGELLRHFWSCFP
VNTPFLEEKVVKMKSNLERFQVTKLCPFQEKIRRQYLSTNLVSHIEEMLQTAYNKLHTWQ
SRRLMKKT
Sequence of entity 4 (D), FASTA
>9XYU_4 General transcription factor IIH subunit 4,  p52 (chains D)
MESTPSRGLNRVHLQCRNLQEFLGGLSPGVLDRLYGHPATCLAVFRELPSLAKNWVMRML
FLEQPLPQAAVALWVKKEFSKAQEESTGLLSGLRIWHTQLLPGGLQGLILNPIFRQNLRI
ALLGGGKAWSDDTSQLGPDKHARDVPSLDKYAEERWEVVLHFMVGSPSAAVSQDLAQLLS
QAGLMKSTEPGEPPCITSAGFQFLLLDTPAQLWYFMLQYLQTAQSRGMDLVEILSFLFQL
SFSTLGKDYSVEGMSDSLLNFLQHLREFGLVFQRKRKSRRYYPTRLAINLSSGVSGAGGT
VHQPGFIVVETNYRLYAYTESELQIALIALFSEMLYRFPNMVVAQVTRESVQQAIASGIT
AQQIIHFLRTRAHPVMLKQTPVLPPTITDQIRLWELERDRLRFTEGVLYNQFLSQVDFEL
LLAHARELGVLVFENSAKRLMVVTPAGHSDVKRFWKRQKHSS
Sequence of entity 5 (E), FASTA
>9XYU_5 General transcription factor IIH subunit 2 (chains E)
MDEEPERTKRWEGGYERTWEILKEDESGSLKATIEDILFKAKRKRVFEHHGQVRLGMMRH
LYVVVDGSRTMEDQDLKPNRLTCTLKLLEYFVEEYFDQNPISQIGIIVTKSKRAEKLTEL
SGNPRKHITSLKKAVDMTCHGEPSLYNSLSIAMQTLKHMPGHTSREVLIIFSSLTTCDPS
NIYDLIKTLKAAKIRVSVIGLSAEVRVCTVLARETGGTYHVILDESHYKELLTHHVSPPP
ASSSSECSLIRMGFPQHTIASLSDQDAKPSFSMAHLDGNTEPGLTLGGYFCPQCRAKYCE
LPVECKICGLTLVSAPHLARSYHHLFPLDAFQEIPLEEYNGERFCYGCQGELKDQHVYVC
AVCQNVFCVDCDVFVHDSLHCCPGCIHKIPAPSGV
Sequence of entity 6 (F), FASTA
>9XYU_6 General transcription factor IIH subunit 3 (chains F)
MVSDEDELNLLVIVVDANPIWWGKQALKESQFTLSKCIDAVMVLGNSHLFMNRSNKLAVI
ASHIQESRFLYPGKNGRLGDFFGDPGNPPEFNPSGSKDGKYELLTSANEVIVEEIKDLMT
KSDIKGQHTETLLAGSLAKALCYIHRMNKEVKDNQEMKSRILVIKAAEDSALQYMNFMNV
IFAAQKQNILIDACVLDSDSGLLQQACDITGGLYLKVPQMPSLLQYLLWVFLPDQDQRSQ
LILPPPVHVDYRAACFCHRNLIEIGYVCSVCLSIFCNFSPICTTCETAFKISLPPVLKAK
KKKLKVSA
Sequence of entity 7 (G), FASTA
>9XYU_7 General transcription factor IIH subunit 5 (chains G)
MVNVLKGVLIECDPAMKQFLLYLDESNALGKKFIIQDIDDTHVFVIAELVNVLQERVGEL
MDQNAFSLTQK
Sequence of entity 8 (H), FASTA
>9XYU_8 DNA repair protein complementing XP-C cells (chains H)
MARKRAAGGEPRGRELRSQKSKAKSKARREEEEEDAFEDEKPPKKSLLSKVSQGKRKRGC
SHPGGSADGPAKKKVAKVTVKSENLKVIKDEALSDGDDLRDFPSDLKKAHHLKRGATMNE
DSNEEEEESENDWEEVEELSEPVLGDVRESTAFSRSLLPVKPVEIEIETPEQAKTRERSE
KIKLEFETYLRRAMKRFNKGVHEDTHKVHLLCLLANGFYRNNICSQPDLHAIGLSIIPAR
FTRVLPRDVDTYYLSNLVKWFIGTFTVNAELSASEQDNLQTTLERRFAIYSARDDEELVH
IFLLILRALQLLTRLVLSLQPIPLKSATAKGKKPSKERLTADPGGSSETSSQVLENHTKP
KTSKGTKQEETFAKGTCRPSAKGKRNKGGRKKRSKPSSSEEDEGPGDKQEKATQRRPHGR
ERRVASRVSYKEESGSDEAGSGSDFELSSGEASDPSDEDSEPGPPKQRKAPAPQRTKAGS
KSASRTHRGSHRKDPSLPAASSSSSSSKRGKKMCSDGEKAEKRSIAGIDQWLEVFCEQEE
KWVCVDCVHGVVGQPLTCYKYATKPMTYVVGIDSDGWVRDVTQRYDPVWMTVTRKCRVDA
EWWAETLRPYQSPFMDREKKEDLEFQAKHMDQPLPTAIGLYKNHPLYALKRHLLKYEAIY
PETAAILGYCRGEAVYSRDCVHTLHSRDTWLKKARVVRLGEVPYKMVKGFSNRARKARLA
EPQLREENDLGLFGYWQTEEYQPPVAVDGKVPRNEFGNVYLFLPSMMPIGCVQLNLPNLH
RVARKLDIDCVQAITGFDFHGGYSHPVTDGYIVCEEFKDVLLTAWENEQAVIERKEKEKK
EKRALGNWKLLAKGLLIRERLKRRYGPKSEAAAPHTDAGGGLSSDEEEGTSSQAEAARIL
AASWPQNREDEEKQKLKGGPKKTKREKKAAASHLFPFEKL
Sequence of entity 9 (K), FASTA
>9XYU_9 DNA repair protein complementing XP-A cells (chains K)
MAAADGALPEAAALEQPAELPASVRASIERKRQRALMLRQARLAARPYSATAAAATGGMA
NVKAAPKIIDTGGGFILEEEEEEEQKIGKVVHQPGPVMEFDYVICEECGKEFMDSYLMNH
FDLPTCDNCRDADDKHKLITKTEAKQEYLLKDCDLEKREPPLKFIVKKNPHHSQWGDMKL
YLKLQIVKRSLEVWGSQEALEEAKEVRQENREKMKQKKFDKKVKELRRAVRSSVWKRETI
VHQHEYGPEENLEDDMYRKTCTMCGHELTYEKM
Sequence of entity 10 (L), FASTA
>9XYU_10 DNA (CY5) (chains L)
GCTGCGGTCCGTTGACGACATACAGCTCAACGATGCCACTTCAGCATTGCACTCATGTTC
TGCCACTAGTACTACGACTACGCCTCGGCTCGA
Sequence of entity 11 (M), FASTA
>9XYU_11 DNA (chains M)
TCGAGCCGAGGCGTAGTCGTAGTACTAGTGTTCAGAATATGAGTACAATACTGAAGTGGC
ATCGTTGAGCTGTATGTCGTCAACGGACCGCAGC

Ligands and cofactors

IDNameFormulaCopies
ZNZinc ionZn6
SF4Iron/sulfur clusterFe4 S41

Primary citation

Pre-incision structures reveal principles of DNA nucleotide excision repair. Li, E.C.L., Kim, J., Brussee, S.J. et al. Nature (2026) 652:1060-1067. DOI 10.1038/s41586-026-10122-5 · PubMed

Other PDB entries of the same protein (UniProt P19447 (AlphaFold model), which also has an AlphaFold model), best resolution first:

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