9ZE2: Endogenous U2/branchpoint spliceosomal complex

Cryo-EM structure of the endogenous U2/branchpoint spliceosomal complex (core). Determined by electron microscopy at 3.26 Å resolution. Released 15 Apr 2026.

Method
Electron microscopy
Resolution
3.26 Å
Organism
Homo sapiens
Chains
12
Atoms
24,516
Mol. weight
690.23 kDa
Ligands
ZN
Released
15 Apr 2026

Explore 9ZE2 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9ZE2 contains 134 α-helices and 138 β-strands across 10 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: 2 helices, 0 β-strands

ElementResiduesLengthSheet
α-helix704-7118
α-helix800-81415
Chain A2: 2 helices, 3 β-strands
ElementResiduesLengthSheet
β-strand46-48339
β-strand54-56339
β-strand61-63339
α-helix66-738
α-helix76-9217
Chain A3: 9 helices, 6 β-strands
ElementResiduesLengthSheet
α-helix383-3842
β-strand385140
β-strand391140
α-helix392-3932
α-helix394-3996
β-strand406-408341
β-strand414-416341
α-helix419-4246
α-helix425-4273
α-helix429-43810
α-helix444-4463
α-helix452-46918
β-strand479-480242
β-strand486-487242
α-helix489-4968
Chain B1: 66 helices, 1 β-strand
ElementResiduesLengthSheet
α-helix380-39516
α-helix401-4077
β-strand412-41435
α-helix416-4183
α-helix466-4683
α-helix469-4713
α-helix489-50517
α-helix509-52113
α-helix523-5264
α-helix528-53912
α-helix546-56217
α-helix568-5703
α-helix571-5788
α-helix579-5835
α-helix587-60317
α-helix607-6137
α-helix615-6184
α-helix622-63918
α-helix641-65313
α-helix658-67417
α-helix677-6793
α-helix683-6919
α-helix692-6943
α-helix699-71517
α-helix722-7243
α-helix725-73511
α-helix741-75313
α-helix754-7563
α-helix759-77517
α-helix776-7783
α-helix782-79716
α-helix803-8053
α-helix806-8105
α-helix811-8166
α-helix822-8243
α-helix827-84418
α-helix846-8538
α-helix855-8584
α-helix862-87817
α-helix886-90217
α-helix909-92113
α-helix922-9287
α-helix929-93810
α-helix945-95713
α-helix959-9646
α-helix968-97912
α-helix987-100418
α-helix1013-102311
α-helix1029-104517
α-helix1047-10493
α-helix1052-106413
α-helix1065-10673
α-helix1071-108818
α-helix1091-10999
α-helix1109-112214
α-helix1125-11273
α-helix1129-11357
α-helix1141-115818
α-helix1159-11646
α-helix1166-117813
α-helix1182-119817
α-helix1205-121511
α-helix1216-12205
α-helix1224-124118
α-helix1243-12508
α-helix1259-127517
α-helix1279-12835
Chain B2: 17 helices, 4 β-strands
ElementResiduesLengthSheet
α-helix452-4587
α-helix459-4613
α-helix462-4665
α-helix472-4743
α-helix477-4815
α-helix485-4928
α-helix499-5013
α-helix502-5043
α-helix526-5294
α-helix533-54614
α-helix552-5609
α-helix568-57710
α-helix595-6028
α-helix612-6165
α-helix625-6273
α-helix631-6399
β-strand65116
β-strand65516
α-helix656-6583
β-strand67517
β-strand68117
Chain B3: 19 helices, 100 β-strands
ElementResiduesLengthSheet
β-strand3-978
β-strand16-2059
β-strand29-3359
β-strand37-4379
β-strand50-5789
β-strand62-68710
β-strand77-82610
β-strand86-93810
β-strand98-106910
β-strand120-123411
β-strand129-133511
β-strand136112
β-strand139-143511
β-strand144-146313
β-strand152-154313
β-strand158-159211
β-strand165-173912
β-strand181-188812
β-strand207-214812
β-strand219-227912
β-strand232-238714
α-helix2391
β-strand247-252614
β-strand255-259514
α-helix264-2663
β-strand267-270414
α-helix271-2722
β-strand273-274215
β-strand287-294816
β-strand299-305716
β-strand309-317916
β-strand322-3321116
β-strand336-342717
β-strand346-351617
β-strand356-361617
β-strand373-374216
β-strand386-387215
β-strand395-401717
β-strand407-413718
β-strand422-427618
α-helix430-4323
β-strand434-440718
β-strand442119
β-strand445-450620
β-strand458-462521
β-strand473-477521
β-strand482-486521
β-strand492-494321
α-helix504-5052
β-strand507-509322
β-strand516-520522
β-strand523-527522
β-strand533-536422
α-helix537-5382
β-strand544-548523
β-strand553-558623
α-helix559-5613
β-strand563-568623
β-strand574-576323
β-strand587-592624
α-helix593-5964
β-strand605-609524
β-strand613-618624
α-helix621-6233
β-strand626-632724
β-strand637-644825
β-strand663-669725
β-strand674-678525
β-strand686-689425
β-strand700-706726
β-strand709-715726
β-strand720-723426
β-strand730-733426
β-strand734119
β-strand741-745520
β-strand753-758620
β-strand761-766620
β-strand774-780718
β-strand785-790627
β-strand797-802627
α-helix809-82618
α-helix833-84412
α-helix849-8524
β-strand864-869627
β-strand876-881627
β-strand886-894928
β-strand903-911928
β-strand914129
β-strand919129
β-strand923-931928
β-strand936-9451028
β-strand951-955530
β-strand958-962530
β-strand967-973730
β-strand976-983830
β-strand990-995631
β-strand999-1004631
β-strand1007132
β-strand1009-1014631
β-strand1021-1025531
β-strand1026133
β-strand1030132
β-strand1033-1038634
β-strand1043-1048634
β-strand1052-1057634
α-helix1073-10764
β-strand1088133
β-strand1090-1096734
β-strand1101-110778
β-strand1116-112168
β-strand1126-113168
α-helix1135-115117
α-helix1160-11645
β-strand1171134
β-strand1173-117538
α-helix1176-11805
α-helix1181-11833
α-helix1187-119610
α-helix1201-121515
Chain B4: 4 helices, 8 β-strands
ElementResiduesLengthSheet
α-helix11-133
β-strand14-18535
α-helix26-349
β-strand39-41335
β-strand46136
α-helix471
β-strand53136
β-strand56-61635
α-helix64-7411
β-strand77-79337
β-strand82-84337
β-strand85-88435
Chain B5: 5 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix9-179
α-helix28-4518
α-helix47-5610
α-helix61-7111
α-helix79-802

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

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
U2 snRNA2RNA37Homo sapiens
PHD finger-like domain-containing protein 5AHprotein110Homo sapiensQ7RTV0 (AlphaFold model)
pre-mRNARRNA42Homo sapiens
RNA-binding protein 5Aprotein824Homo sapiensP52756 (AlphaFold model)
Splicing factor 3B subunit 1B1protein1304Homo sapiensO75533 (AlphaFold model)
Splicing factor 3B subunit 2B2protein895Homo sapiensQ13435 (AlphaFold model)
Splicing factor 3B subunit 3B3protein1217Homo sapiensQ15393
Splicing factor 3B subunit 4B4protein424Homo sapiensQ15427
Splicing factor 3B subunit 5B5protein86Homo sapiensQ9BWJ5
Splicing factor 3B subunit 6B6protein125Homo sapiensQ9Y3B4
Splicing factor 3A subunit 2A2protein464Homo sapiensQ15428
Splicing factor 3A subunit 3A3protein501Homo sapiensQ12874
Sequence of entity 1 (2), FASTA
>9ZE2_1 U2 snRNA (chains 2)
AAGUGUAGUAUCUGUUCUCAUCAGUUUAAUAUCUGAU
Sequence of entity 2 (H), FASTA
>9ZE2_2 PHD finger-like domain-containing protein 5A (chains H)
MAKHHPDLIFCRKQAGVAIGRLCEKCDGKCVICDSYVRPCTLVRICDECNYGSYQGRCVI
CGGPGVSDAYYCKECTIQEKDRDGCPKIVNLGSSKTDLFYERKKYGFKKR
Sequence of entity 3 (R), FASTA
>9ZE2_3 pre-mRNA (chains R)
UUUUUUUUUUUUUUUCUCAUUUUUUUUUUUUUUUUUUUUUUU
Sequence of entity 4 (A), FASTA
>9ZE2_4 RNA-binding protein 5 (chains A)
MGSDKRVSRTERSGRYGSIIDRDDRDERESRSRRRDSDYKRSSDDRRGDRYDDYRDYDSP
ERERERRNSDRSEDGYHSDGDYGEHDYRHDISDERESKTIMLRGLPITITESDIREMMES
FEGPQPADVRLMKRKTGVSRGFAFVEFYHLQDATSWMEANQKKLVIQGKHIAMHYSNPRP
KFEDWLCNKCCLNNFRKRLKCFRCGADKFDSEQEVPPGTTESVQSVDYYCDTIILRNIAP
HTVVDSIMTALSPYASLAVNNIRLIKDKQTQQNRGFAFVQLSSAMDASQLLQILQSLHPP
LKIDGKTIGVDFAKSARKDLVLSDGNRVSAFSVASTAIAAAQWSSTQSQSGEGGSVDYSY
LQPGQDGYAQYAQYSQDYQQFYQQQAGGLESDASSASGTAVTTTSAAVVSQSPQLYNQTS
NPPGSPTEEAQPSTSTSTQAPAASPTGVVPGTKYAVPDTSTYQYDESSGYYYDPTTGLYY
DPNSQYYYNSLTQQYLYWDGEKETYVPAAESSSHQQSGLPPAKEGKEKKEKPKSKTAQQI
AKDMERWAKSLNKQKENFKNSFQPVNSLREEERRESAAADAGFALFEKKGALAERQQLIP
ELVRNGDEENPLKRGLVAAYSGDSDNEEELVERLESEEEKLADWKKMACLLCRRQFPNKD
ALVRHQQLSDLHKQNMDIYRRSRLSEQELEALELREREMKYRDRAAERREKYGIPEPPEP
KRKKQFDAGTVNYEQPTKDGIDHSNIGNKMLQAMGWREGSGLGRKCQGITAPIEAQVRLK
GAGLGAKGSAYGLSGADSYKDAVRKAMFARFTEMEMDYKDDDDK
Sequence of entity 5 (B1), FASTA
>9ZE2_5 Splicing factor 3B subunit 1 (chains B1)
MAKIAKTHEDIEAQIREIQGKKAALDEAQGVGLDSTGYYDQEIYGGSDSRFAGYVTSIAA
TELEDDDDDYSSSTSLLGQKKPGYHAPVALLNDIPQSTEQYDPFAEHRPPKIADREDEYK
KHRRTMIISPERLDPFADGGKTPDPKMNARTYMDVMREQHLTKEEREIRQQLAEKAKAGE
LKVVNGAAASQPPSKRKRRWDQTADQTPGATPKKLSSWDQAETPGHTPSLRWDETPGRAK
GSETPGATPGSKIWDPTPSHTPAGAATPGRGDTPGHATPGHGGATSSARKNRWDETPKTE
RDTPGHGSGWAETPRTDRGGDSIGETPTPGASKRKSRWDETPASQMGGSTPVLTPGKTPI
GTPAMNMATPTPGHIMSMTPEQLQAWRWEREIDERNRPLSDEELDAMFPEGYKVLPPPAG
YVPIRTPARKLTATPTPLGGMTGFHMQTEDRTMKSVNDQPSGNLPFLKPDDIQYFDKLLV
DVDESTLSPEEQKERKIMKLLLKIKNGTPPMRKAALRQITDKAREFGAGPLFNQILPLLM
SPTLEDQERHLLVKVIDRILYKLDDLVRPYVHKILVVIEPLLIDEDYYARVEGREIISNL
AKAAGLATMISTMRPDIDNMDEYVRNTTARAFAVVASALGIPSLLPFLKAVCKSKKSWQA
RHTGIKIVQQIAILMGCAILPHLRSLVEIIEHGLVDEQQKVRTISALAIAALAEAATPYG
IESFDSVLKPLWKGIRQHRGKGLAAFLKAIGYLIPLMDAEYANYYTREVMLILIREFQSP
DEEMKKIVLKVVKQCCGTDGVEANYIKTEILPPFFKHFWQHRMALDRRNYRQLVDTTVEL
ANKVGAAEIISRIVDDLKDEAEQYRKMVMETIEKIMGNLGAADIDHKLEEQLIDGILYAF
QEQTTEDSVMLNGFGTVVNALGKRVKPYLPQICGTVLWRLNNKSAKVRQQAADLISRTAV
VMKTCQEEKLMGHLGVVLYEYLGEEYPEVLGSILGALKAIVNVIGMHKMTPPIKDLLPRL
TPILKNRHEKVQENCIDLVGRIADRGAEYVSAREWMRICFELLELLKAHKKAIRRATVNT
FGYIAKAIGPHDVLATLLNNLKVQERQNRVCTTVAIAIVAETCSPFTVLPALMNEYRVPE
LNVQNGVLKSLSFLFEYIGEMGKDYIYAVTPLLEDALMDRDLVHRQTASAVVQHMSLGVY
GFGCEDSLNHLLNYVWPNVFETSPHVIQAVMGALEGLRVAIGPCRMLQYCLQGLFHPARK
VRDVYWKIYNSIYIGSQDALIAHYPRIYNDDKNTYIRYELDYIL
Sequence of entity 6 (B2), FASTA
>9ZE2_6 Splicing factor 3B subunit 2 (chains B2)
MATEHPEPPKAELQLPPPPPPGHYGAWAAQELQAKLAEIGAPIQGNREELVERLQSYTRQ
TGIVLNRPVLRGEDGDKAAPPPMSAQLPGIPMPPPPLGLPPLQPPPPPPPPPPGLGLGFP
MAHPPNLGPPPPLRVGEPVALSEEERLKLAQQQAALLMQQEERAKQQGDHSLKEHELLEQ
QKRAAVLLEQERQQEIAKMGTPVPRPPQDMGQIGVRTPLGPRVAAPVGPVGPTPTVLPMG
APVPRPRGPPPPPGDENREMDDPSVGPKIPQALEKILQLKESRQEEMNSQQEEEEMETDA
RSSLGQSASETEEDTVSVSKKEKNRKRRNRKKKKKPQRVRGVSSESSGDREKDSTRSRGS
DSPAADVEIEYVTEEPEIYEPNFIFFKRIFEAFKLTDDVKKEKEKEPEKLDKLENSAAPK
KKGFEEEHKDSDDDSSDDEQEKKPEAPKLSKKKLRRMNRFTVAELKQLVARPDVVEMHDV
TAQDPKLLVHLKATRNSVPVPRHWCFKRKYLQGKRGIEKPPFELPDFIKRTGIQEMREAL
QEKEEQKTMKSKMREKVRPKMGKIDIDYQKLHDAFFKWQTKPKLTIHGDLYYEGKEFETR
LKEKKPGDLSDELRISLGMPVGPNAHKVPPPWLIAMQRYGPPPSYPNLKIPGLNSPIPES
CSFGYHAGGWGKPPVDETGKPLYGDVFGTNAAEFQTKTEEEEIDRTPWGELEPSDEESSE
EEEEEESDEDKPDETGFITPADSGLITPGGFSSVPAGMETPELIELRKKKIEEAMDGSET
PQLFTVLPEKRTATVGGAMMGSTHIYDMSTVMSRKGPAPELQGVEVALAPEELELDPMAM
TQKYEEHVREQQAQVEKEDFSDMVAEHAAKQKQKKRKAQPQDSRGGSKKYKEFKF
Sequence of entity 7 (B3), FASTA
>9ZE2_7 Splicing factor 3B subunit 3 (chains B3)
MFLYNLTLQRATGISFAIHGNFSGTKQQEIVVSRGKILELLRPDPNTGKVHTLLTVEVFG
VIRSLMAFRLTGGTKDYIVVGSDSGRIVILEYQPSKNMFEKIHQETFGKSGCRRIVPGQF
LAVDPKGRAVMISAIEKQKLVYILNRDAAARLTISSPLEAHKANTLVYHVVGVDVGFENP
MFACLEMDYEEADNDPTGEAAANTQQTLTFYELDLGLNHVVRKYSEPLEEHGNFLITVPG
GSDGPSGVLICSENYITYKNFGDQPDIRCPIPRRRNDLDDPERGMIFVCSATHKTKSMFF
FLAQTEQGDIFKITLETDEDMVTEIRLKYFDTVPVAAAMCVLKTGFLFVASEFGNHYLYQ
IAHLGDDDEEPEFSSAMPLEEGDTFFFQPRPLKNLVLVDELDSLSPILFCQIADLANEDT
PQLYVACGRGPRSSLRVLRHGLEVSEMAVSELPGNPNAVWTVRRHIEDEFDAYIIVSFVN
ATLVLSIGETVEEVTDSGFLGTTPTLSCSLLGDDALVQVYPDGIRHIRADKRVNEWKTPG
KKTIVKCAVNQRQVVIALTGGELVYFEMDPSGQLNEYTERKEMSADVVCMSLANVPPGEQ
RSRFLAVGLVDNTVRIISLDPSDCLQPLSMQALPAQPESLCIVEMGGTEKQDELGERGSI
GFLYLNIGLQNGVLLRTVLDPVTGDLSDTRTRYLGSRPVKLFRVRMQGQEAVLAMSSRSW
LSYSYQSRFHLTPLSYETLEFASGFASEQCPEGIVAISTNTLRILALEKLGAVFNQVAFP
LQYTPRKFVIHPESNNLIIIETDHNAYTEATKAQRKQQMAEEMVEAAGEDERELAAEMAA
AFLNENLPESIFGAPKAGNGQWASVIRVMNPIQGNTLDLVQLEQNEAAFSVAVCRFSNTG
EDWYVLVGVAKDLILNPRSVAGGFVYTYKLVNNGEKLEFLHKTPVEEVPAAIAPFQGRVL
IGVGKLLRVYDLGKKKLLRKCENKHIANYISGIQTIGHRVIVSDVQESFIWVRYKRNENQ
LIIFADDTYPRWVTTASLLDYDTVAGADKFGNICVVRLPPNTNDEVDEDPTGNKALWDRG
LLNGASQKAEVIMNYHVGETVLSLQKTTLIPGGSESLVYTTLSGGIGILVPFTSHEDHDF
FQHVEMHLRSEHPPLCGRDHLSFRSYYFPVKNVIDGDLCEQFNSMEPNKQKNVSEELDRT
PPEVSKKLEDIRTRYAF
Sequence of entity 8 (B4), FASTA
>9ZE2_8 Splicing factor 3B subunit 4 (chains B4)
MAAGPISERNQDATVYVGGLDEKVSEPLLWELFLQAGPVVNTHMPKDRVTGQHQGYGFVE
FLSEEDADYAIKIMNMIKLYGKPIRVNKASAHNKNLDVGANIFIGNLDPEIDEKLLYDTF
SAFGVILQTPKIMRDPDTGNSKGYAFINFASFDASDAAIEAMNGQYLCNRPITVSYAFKK
DSKGERHGSAAERLLAAQNPLSQADRPHQLFADAPPPPSAPNPVVSSLGSGLPPPGMPPP
GSFPPPVPPPGALPPGIPPAMPPPPMPPGAAGHGPPSAGTPGAGHPGHGHSHPHPFPPGG
MPHPGMSQMQLAHHGPHGLGHPHAGPPGSGGQPPPRPPPGMPHPGPPPMGMPPRGPPFGS
PMGHPGPMPPHGMRGPPPLMPPHGYTGPPRPPPYGYQRGPLPPPRPTPRPPVPPRGPLRG
PLPQ
Sequence of entity 9 (B5), FASTA
>9ZE2_9 Splicing factor 3B subunit 5 (chains B5)
MTDRYTIHSQLEHLQSKYIGTGHADTTKWEWLVNQHRDSYCSYMGHFDLLNYFAIAENES
KARVRFNLMEKMLQPCGPPADKPEEN
Sequence of entity 10 (B6), FASTA
>9ZE2_10 Splicing factor 3B subunit 6 (chains B6)
MAMQAAKRANIRLPPEVNRILYIRNLPYKITAEEMYDIFGKYGPIRQIRVGNTPETRGTA
YVVYEDIFDAKNACDHLSGFNVCNRYLVVLYYNANRAFQKMDTKKKEEQLKLLKEKYGIN
TDPPK
Sequence of entity 11 (A2), FASTA
>9ZE2_11 Splicing factor 3A subunit 2 (chains A2)
MDFQHRPGGKTGSGGVASSSESNRDRRERLRQLALETIDINKDPYFMKNHLGSYECKLCL
TLHNNEGSYLAHTQGKKHQTNLARRAAKEAKEAPAQPAPEKVKVEVKKFVKIGRPGYKVT
KQRDSEMGQQSLLFQIDYPEIAEGIMPRHRFMSAYEQRIEPPDRRWQYLLMAAEPYETIA
FKVPSREIDKAEGKFWTHWNRETKQFFLQFHFKMEKPPAPPSLPAGPPGVKRPPPPLMNG
LPPRPPLPESLPPPPPGGLPLPPMPPTGPAPSGPPGPPQLPPPAPGVHPPAPVVHPPASG
VHPPAPGVHPPAPGVHPPAPGVHPPTSGVHPPAPGVHPPAPGVHPPAPGVHPPAPGVHPP
APGVHPPPSAGVHPQAPGVHPAAPAVHPQAPGVHPPAPGMHPQAPGVHPQPPGVHPSAPG
VHPQPPGVHPSNPGVHPPTPMPPMLRPPLPSEGPGNIPPPPPTN
Sequence of entity 12 (A3), FASTA
>9ZE2_12 Splicing factor 3A subunit 3 (chains A3)
METILEQQRRYHEEKERLMDVMAKEMLTKKSTLRDQINSDHRTRAMQDRYMEVSGNLRDL
YDDKDGLRKEELNAISGPNEFAEFYNRLKQIKEFHRKHPNEICVPMSVEFEELLKARENP
SEEAQNLVEFTDEEGYGRYLDLHDCYLKYINLKASEKLDYITYLSIFDQLFDIPKERKNA
EYKRYLEMLLEYLQDYTDRVKPLQDQNELFGKIQAEFEKKWENGTFPGWPKETSSALTHA
GAHLDLSAFSSWEELASLGLDRLKSALLALGLKCGGTLEERAQRLFSTKGKSLESLDTSL
FAKNPKSKGTKRDTERNKDIAFLEAQIYEYVEILGEQRHLTHENVQRKQARTGEEREEEE
EEQISESESEDEENEIIYNPKNLPLGWDGKPIPYWLYKLHGLNINYNCEICGNYTYRGPK
AFQRHFAEWRHAHGMRCLGIPNTAHFANVTQIEDAVSLWAKLKLQKASERWQPDTEEEYE
DSSGNVVNKKTYEDLKRQGLL

Ligands and cofactors

IDNameFormulaCopies
ZNZinc ionZn3

Primary citation

The tumour suppressor RBM5 activates the helicase DHX15 to regulate splicing. Liu, S., Su, T., Huang, J. et al. bioRxiv (2026). DOI 10.64898/2026.03.26.714623 · PubMed

Other PDB entries of the same protein (UniProt Q7RTV0 (AlphaFold model), which also has an AlphaFold model), best resolution first:

Browse structure collections

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