9PD4: NER dual incision complex - DuIM

NER dual incision complex - DuIM. Determined by electron microscopy at 3.4 Å resolution. Released 18 Feb 2026.

Method
Electron microscopy
Resolution
3.4 Å
Organisms
Homo sapiens, synthetic construct
Chains
16
Atoms
40,053
Mol. weight
850.52 kDa
Ligands
ZN, SF4
Released
18 Feb 2026

Explore 9PD4 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9PD4 contains 224 α-helices and 197 β-strands across 14 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: 28 helices, 29 β-strands

ElementResiduesLengthSheet
β-strand5311
β-strand59-6021
α-helix62-643
β-strand77-7822
β-strand84-8742
α-helix93-10311
β-strand105-10842
β-strand113-11752
α-helix120-1289
α-helix133-14210
α-helix150-16112
β-strand167-17153
β-strand174-17963
α-helix182-1898
α-helix193-1964
β-strand268-27143
α-helix274-2763
α-helix277-28610
α-helix290-2912
β-strand29213
β-strand294-29524
α-helix315-3173
α-helix318-32710
β-strand32814
β-strand333-33424
β-strand336-33945
α-helix347-35711
β-strand362-36545
α-helix368-38215
β-strand389-39245
β-strand405-40955
α-helix421-43010
β-strand43516
β-strand438-44145
α-helix443-4453
α-helix452-4576
β-strand46016
β-strand463-46755
α-helix477-4837
β-strand487-49045
α-helix493-4986
β-strand506-51277
α-helix513-5153
α-helix516-5249
α-helix538-55316
β-strand558-56257
α-helix565-57511
β-strand57917
α-helix587-59711
β-strand604-60747
β-strand622-62657
α-helix633-64210
β-strand658-66477
α-helix669-6768
α-helix678-6836
β-strand68817
α-helix706-71611
Chain B: 38 helices, 29 β-strands
ElementResiduesLengthSheet
β-strand2-548
β-strand8-1148
α-helix19-3416
β-strand3719
β-strand38-41410
α-helix48-6215
β-strand70-71211
β-strand74112
α-helix77-9822
β-strand105-108411
α-helix119-1213
α-helix127-13812
α-helix140-1467
α-helix156-1605
α-helix161-1655
α-helix169-1702
β-strand173-175311
α-helix177-18711
α-helix191-1988
β-strand204-206311
β-strand208112
α-helix210-2134
α-helix215-2217
β-strand230-231211
β-strand232-234310
α-helix239-2468
β-strand249-252413
α-helix253-28634
α-helix324-34421
β-strand352113
α-helix354-3563
α-helix357-3604
α-helix361-3655
α-helix369-3735
α-helix376-38611
α-helix395-40915
β-strand414-419613
β-strand432-437613
α-helix445-4506
β-strand455-458410
α-helix467-4704
β-strand47519
β-strand479-480210
β-strand490114
β-strand493-494215
β-strand495116
α-helix514-52815
β-strand535-539515
α-helix542-55413
α-helix558-5647
β-strand566-569415
α-helix574-58916
β-strand594-599615
α-helix603-6075
β-strand617-621515
α-helix631-64313
α-helix648-66417
α-helix665-6673
β-strand675114
β-strand676-679415
α-helix682-6854
α-helix687-6904
α-helix695-6984
β-strand708116
α-helix710-72415
α-helix727-7282
α-helix730-7345
Chain C: 8 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix399-41012
α-helix419-4213
α-helix424-43411
α-helix455-47723
α-helix485-50016
α-helix501-5055
α-helix506-51510
α-helix521-54727
Chain D: 22 helices, 18 β-strands
ElementResiduesLengthSheet
α-helix19-257
α-helix28-347
α-helix38-4710
α-helix50-589
β-strand66-67217
α-helix68-736
β-strand75118
α-helix77-793
α-helix80-9314
β-strand96-101617
β-strand105-110617
α-helix112-12312
α-helix145-16420
α-helix173-18210
β-strand185-186219
β-strand195-196219
α-helix198-2058
α-helix208-22114
α-helix230-24112
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-helix322-3298
β-strand333-33751
β-strand342-34541
α-helix348-3569
α-helix361-37010
β-strand37211
α-helix385-39511
α-helix398-4003
β-strand402-409822
α-helix415-42814
β-strand431-435522
β-strand440-443422
α-helix448-46114
Chain E: 15 helices, 12 β-strands
ElementResiduesLengthSheet
α-helix30-334
β-strand60-66723
α-helix69-724
α-helix80-9819
β-strand103-110823
β-strand113-121923
α-helix124-13411
α-helix137-1393
α-helix145-15713
β-strand166-171623
α-helix183-1919
β-strand195-200623
α-helix206-21510
β-strand219-221323
α-helix225-23612
α-helix238-2392
β-strand249-251324
α-helix273-2775
β-strand303-304225
β-strand311-312225
β-strand332-335426
α-helix336-3383
β-strand356-359426
β-strand366-367226
α-helix369-3746
α-helix375-3795
α-helix383-3864
Chain F: 12 helices, 11 β-strands
ElementResiduesLengthSheet
β-strand8-16927
α-helix19-2810
α-helix34-5118
β-strand56-62727
β-strand68-69227
α-helix103-12018
α-helix133-15018
β-strand157-165927
α-helix174-18613
β-strand190-195627
α-helix201-21010
β-strand214-216327
α-helix220-2223
α-helix223-2264
α-helix227-2315
α-helix235-2384
β-strand241118
α-helix243-2464
β-strand254128
β-strand261128
β-strand266-268324
β-strand274-275224
α-helix283-2875
Chain G: 2 helices, 3 β-strands
ElementResiduesLengthSheet
β-strand6-11622
α-helix14-2613
β-strand36-37222
β-strand42-45422
α-helix50-6314
Chain K: 10 helices, 11 β-strands
ElementResiduesLengthSheet
α-helix23-4422
α-helix91-966
β-strand103-104229
β-strand111-112229
α-helix116-1216
β-strand138-140330
α-helix141-1488
α-helix152-1565
α-helix1611
β-strand165-167330
α-helix176-1772
β-strand178-182530
α-helix186-1949
α-helix197-22933
β-strand246131
β-strand250132
α-helix253-2553
β-strand258132
β-strand259133
β-strand261131
β-strand268133

6 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-A cellsKprotein273Homo sapiensP23025
DNA (Cy5)LDNA93synthetic construct
DNAMDNA94synthetic construct
Replication protein A 70 kDa DNA-binding subunitNprotein616Homo sapiensP27694
Replication protein A 32 kDa subunitOprotein270Homo sapiensP15927

4 more molecules are not listed.

Sequence of entity 1 (A), FASTA
>9PD4_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
>9PD4_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
>9PD4_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
>9PD4_4 General transcription factor IIH subunit 4,  p52 (chains D)
MESTPSRGLNRVHLQCRNLQEFLGGLSPGVLDRLYGHPATCLAVFRELPSLAKNWVMRML
FLEQPLPQAAVALWVKKEFSKAQEESTGLLSGLRIWHTQLLPGGLQGLILNPIFRQNLRI
ALLGGGKAWSDDTSQLGPDKHARDVPSLDKYAEERWEVVLHFMVGSPSAAVSQDLAQLLS
QAGLMKSTEPGEPPCITSAGFQFLLLDTPAQLWYFMLQYLQTAQSRGMDLVEILSFLFQL
SFSTLGKDYSVEGMSDSLLNFLQHLREFGLVFQRKRKSRRYYPTRLAINLSSGVSGAGGT
VHQPGFIVVETNYRLYAYTESELQIALIALFSEMLYRFPNMVVAQVTRESVQQAIASGIT
AQQIIHFLRTRAHPVMLKQTPVLPPTITDQIRLWELERDRLRFTEGVLYNQFLSQVDFEL
LLAHARELGVLVFENSAKRLMVVTPAGHSDVKRFWKRQKHSS
Sequence of entity 5 (E), FASTA
>9PD4_5 General transcription factor IIH subunit 2 (chains E)
MDEEPERTKRWEGGYERTWEILKEDESGSLKATIEDILFKAKRKRVFEHHGQVRLGMMRH
LYVVVDGSRTMEDQDLKPNRLTCTLKLLEYFVEEYFDQNPISQIGIIVTKSKRAEKLTEL
SGNPRKHITSLKKAVDMTCHGEPSLYNSLSIAMQTLKHMPGHTSREVLIIFSSLTTCDPS
NIYDLIKTLKAAKIRVSVIGLSAEVRVCTVLARETGGTYHVILDESHYKELLTHHVSPPP
ASSSSECSLIRMGFPQHTIASLSDQDAKPSFSMAHLDGNTEPGLTLGGYFCPQCRAKYCE
LPVECKICGLTLVSAPHLARSYHHLFPLDAFQEIPLEEYNGERFCYGCQGELKDQHVYVC
AVCQNVFCVDCDVFVHDSLHCCPGCIHKIPAPSGV
Sequence of entity 6 (F), FASTA
>9PD4_6 General transcription factor IIH subunit 3 (chains F)
MVSDEDELNLLVIVVDANPIWWGKQALKESQFTLSKCIDAVMVLGNSHLFMNRSNKLAVI
ASHIQESRFLYPGKNGRLGDFFGDPGNPPEFNPSGSKDGKYELLTSANEVIVEEIKDLMT
KSDIKGQHTETLLAGSLAKALCYIHRMNKEVKDNQEMKSRILVIKAAEDSALQYMNFMNV
IFAAQKQNILIDACVLDSDSGLLQQACDITGGLYLKVPQMPSLLQYLLWVFLPDQDQRSQ
LILPPPVHVDYRAACFCHRNLIEIGYVCSVCLSIFCNFSPICTTCETAFKISLPPVLKAK
KKKLKVSA
Sequence of entity 7 (G), FASTA
>9PD4_7 General transcription factor IIH subunit 5 (chains G)
MVNVLKGVLIECDPAMKQFLLYLDESNALGKKFIIQDIDDTHVFVIAELVNVLQERVGEL
MDQNAFSLTQK
Sequence of entity 8 (K), FASTA
>9PD4_8 DNA repair protein complementing XP-A cells (chains K)
MAAADGALPEAAALEQPAELPASVRASIERKRQRALMLRQARLAARPYSATAAAATGGMA
NVKAAPKIIDTGGGFILEEEEEEEQKIGKVVHQPGPVMEFDYVICEECGKEFMDSYLMNH
FDLPTCDNCRDADDKHKLITKTEAKQEYLLKDCDLEKREPPLKFIVKKNPHHSQWGDMKL
YLKLQIVKRSLEVWGSQEALEEAKEVRQENREKMKQKKFDKKVKELRRAVRSSVWKRETI
VHQHEYGPEENLEDDMYRKTCTMCGHELTYEKM
Sequence of entity 9 (L), FASTA
>9PD4_9 DNA (Cy5) (chains L)
GCTGCGGTCCGTTGACGACATACAGCTCAACGATGCCACTTCAGCATTGCACTCATGTTC
TGXCACTAGTACTACGACTACGCCTCGGCTCGA
Sequence of entity 10 (M), FASTA
>9PD4_10 DNA (chains M)
TCGAGCCGAGGCGTAGTCGTAGTACTAGTGTTCAGAATATGAGTACAATACTGAAGTGGC
ATCGTTGAGCTGTATGTCGTCAACGGACCGCAGC
Sequence of entity 11 (N), FASTA
>9PD4_11 Replication protein A 70 kDa DNA-binding subunit (chains N)
MVGQLSEGAIAAIMQKGDTNIKPILQVINIRPITTGNSPPRYRLLMSDGLNTLSSFMLAT
QLNPLVEEEQLSSNCVCQIHRFIVNTLKDGRRVVILMELEVLKSAEAVGVKIGNPVPYNE
GLGQPQVAPPAPAASPAASSRPQPQNGSSGMGSTVSKAYGASKTFGKAAGPSLSHTSGGT
QSKVVPIASLTPYQSKWTICARVTNKSQIRTWSNSRGEGKLFSLELVDESGEIRATAFNE
QVDKFFPLIEVNKVYYFSKGTLKIANKQFTAVKNDYEMTFNNETSVMPCEDDHHLPTVQF
DFTGIDDLENKSKDSLVDIIGICKSYEDATKITVRSNNREVAKRNIYLMDTSGKVVTATL
WGEDADKFDGSRQPVLAIKGARVSDFGGRSLSVLSSSTIIANPDIPEAYKLRGWFDAEGQ
ALDGVSISDLKSGGVGGSNTNWKTLYEVKSENLGQGDKPDYFSSVATVVYLRKENCMYQA
CPTQDCNKKVIDQQNGLYRCEKCDTEFPNFKYRMILSVNIADFQENQWVTCFQESAEAIL
GQNAAYLGELKDKNEQAFEEVFQNANFRSFIFRVRVKVETYNDESRIKATVMDVKPVDYR
EYGRRLVMSIRRSALM
Sequence of entity 12 (O), FASTA
>9PD4_12 Replication protein A 32 kDa subunit (chains O)
MWNSGFESYGSSSYGGAGGYTQSPGGFGSPAPSQAEKKSRARAQHIVPCTISQLLSATLV
DEVFRIGNVEISQVTIVGIIRHAEKAPTNIVYKIDDMTAAPMDVRQWVDTDDTSSENTVV
PPETYVKVAGHLRSFQNKKSLVAFKIMPLEDMNEFTTHILEVINAHMVLSKANSQPSAGR
APISNPGMSEAGNFGGNSFMPANGLTVAQNQVLNLIKACPRPEGLNFQDLKNQLKHMSVS
SIKQAVDFLSNEGHIYSTVDDDHFKSTDAE

Ligands and cofactors

IDNameFormulaCopies
ZNZinc ionZn7
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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