9QEQ: Transcribing Pol II-DSIF-SPT6-U1 snRNP complex

Structure of the transcribing Pol II-DSIF-SPT6-U1 snRNP complex. Determined by electron microscopy at 3.5 Å resolution. Released 9 Jul 2025.

Method
Electron microscopy
Resolution
3.5 Å
Organisms
Sus scrofa domesticus, Homo sapiens, synthetic construct
Chains
28
Atoms
56,145
Mol. weight
1132.74 kDa
Ligands
MG, ZN
Released
9 Jul 2025

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

Secondary structure: helices and β-sheets

9QEQ contains 300 α-helices and 347 β-strands across 24 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: 80 helices, 74 β-strands

ElementResiduesLengthSheet
α-helix141
β-strand1511
β-strand20-2232
α-helix28-347
β-strand4313
β-strand47-4824
β-strand51-5224
β-strand5613
α-helix60-623
α-helix81-833
β-strand86-9495
α-helix97-993
α-helix100-10910
β-strand11016
β-strand11716
α-helix124-1329
α-helix137-15014
α-helix188-1892
β-strand190-19457
β-strand197-20157
β-strand213-21647
α-helix218-22710
α-helix230-2367
α-helix245-2484
β-strand250-25565
α-helix262-2632
β-strand26418
α-helix265-2662
α-helix2711
β-strand27218
α-helix273-2742
α-helix275-29521
α-helix300-31819
α-helix326-3283
β-strand32919
β-strand33519
α-helix339-3435
α-helix350-3534
β-strand357-359310
β-strand362-369811
β-strand377-381511
α-helix382-3876
β-strand389-393512
α-helix399-40810
β-strand416-420512
β-strand426-428312
α-helix435-4384
β-strand445-449512
α-helix4501
β-strand455-459511
α-helix466-4683
β-strand469-477911
β-strand482-484311
α-helix486-4883
α-helix489-4924
β-strand500-504511
α-helix509-51810
β-strand520111
β-strand526-527213
β-strand532-533213
α-helix540-5489
β-strand554-555214
α-helix557-56610
α-helix5741
β-strand575115
α-helix576-5772
β-strand579-581316
β-strand585116
β-strand586-587214
α-helix588-5958
β-strand602-605417
α-helix612-6143
β-strand627-630417
β-strand635-637317
α-helix642-6454
α-helix652-6598
α-helix662-68322
β-strand686118
α-helix689-6924
α-helix696-72227
α-helix727-7282
α-helix733-75927
α-helix765-7728
α-helix778-7858
β-strand789-790219
β-strand793120
β-strand796120
α-helix798-7992
β-strand802121
β-strand805121
α-helix817-8204
β-strand822-823219
α-helix833-86836
β-strand872-873222
β-strand879-880222
β-strand886-888322
α-helix891-8933
β-strand896123
β-strand902-905424
α-helix913-9208
α-helix927-9337
β-strand934125
α-helix936-9449
α-helix946-96924
β-strand976-979424
α-helix983-99412
β-strand1002125
α-helix1006-101914
α-helix1028-103710
α-helix1039-104810
α-helix1051-10533
α-helix1054-10585
α-helix1062-107918
β-strand1080123
α-helix1081-10822
α-helix1087-110115
α-helix1121-11299
β-strand1139-1141326
β-strand1143127
α-helix1146-11483
α-helix1151-116111
β-strand1162128
β-strand1164-1165229
α-helix1166-11694
β-strand1170-1177830
α-helix1187-119610
β-strand1210-1216730
α-helix1218-12236
α-helix1228-123912
α-helix1240-12423
β-strand1243-1247530
β-strand1256-1261630
α-helix1281-129515
β-strand1297-1299329
β-strand1301128
β-strand1307-1310426
β-strand1319-1322431
β-strand1328-1331431
β-strand1335-1338426
α-helix1343-13486
β-strand1352127
β-strand1358-1359226
α-helix1362-13698
α-helix1371-138717
α-helix1395-140612
β-strand1408-1409232
α-helix1416-14205
α-helix1426-14327
α-helix1435-144511
β-strand1448-144922
α-helix1454-14607
α-helix1467-14704
β-strand1471-1475533
α-helix1478-14803
Chain b: 5 helices, 6 β-strands
ElementResiduesLengthSheet
α-helix53-553
α-helix66-8823
β-strand104-108596
α-helix116-1238
β-strand129-136896
β-strand143-151996
α-helix154-16411
β-strand168-169297
β-strand172-173297
β-strand175-177396
α-helix190-1923
Chain B: 55 helices, 81 β-strands
ElementResiduesLengthSheet
α-helix19-3719
α-helix41-499
α-helix50-545
α-helix55-617
α-helix63-653
β-strand67-68236
β-strand83-84236
β-strand87-93737
β-strand98-99238
β-strand105-106238
α-helix110-1167
β-strand121-1301037
β-strand132-133236
β-strand142-1531237
α-helix168-1736
β-strand185-187339
β-strand190-193439
α-helix194-1952
β-strand196-200540
α-helix2011
β-strand206-208341
β-strand215-223941
α-helix2321
β-strand233-240841
α-helix241-2422
β-strand256-259441
β-strand263142
β-strand267-268241
α-helix269-2768
α-helix281-2888
α-helix295-30713
α-helix314-32411
β-strand325142
α-helix332-34110
α-helix342-3465
α-helix358-37720
α-helix381-3844
α-helix388-3903
β-strand391-394440
α-helix396-42328
α-helix431-4344
α-helix437-45014
β-strand453143
β-strand463143
β-strand468-469239
α-helix4701
α-helix475-4828
β-strand484-486340
α-helix496-4994
α-helix503-5053
β-strand509144
β-strand523-525340
β-strand526144
β-strand531-534445
α-helix539-5479
β-strand551-552246
α-helix558-5614
β-strand565-569546
β-strand572-577646
α-helix580-59314
β-strand602-606546
β-strand611-615546
β-strand618-622545
β-strand623-627547
β-strand628-629248
β-strand632-633248
α-helix637-6448
α-helix646-6494
α-helix653-6597
β-strand662-665447
α-helix667-6726
β-strand675-676247
α-helix679-6824
β-strand695-696247
α-helix700-7034
α-helix706-7094
α-helix714-7163
β-strand717149
α-helix719-73113
β-strand732150
β-strand734-735251
α-helix738-7414
β-strand747-751551
β-strand755152
β-strand759-760253
α-helix764-7674
β-strand776-782754
β-strand783118
β-strand793-797554
α-helix798-8025
β-strand808-8171051
β-strand827-829355
β-strand853156
β-strand859-861351
β-strand865-867356
β-strand869-872455
β-strand889-891355
β-strand894-895256
α-helix896-8972
β-strand901-9111151
β-strand917-9271151
β-strand934-936354
β-strand942-949854
β-strand956157
β-strand965-967354
α-helix971-9755
β-strand977149
α-helix978-99316
β-strand997-998253
α-helix1008-101710
β-strand1025-1026254
β-strand1029157
α-helix10351
β-strand1036157
α-helix10371
β-strand1041-10501054
β-strand1051150
α-helix1055-10573
β-strand1060-1062311
β-strand1067158
β-strand1074158
α-helix10751
α-helix1078-10803
β-strand1084-1086310
α-helix1088-109710
α-helix1100-11045
α-helix1105-11095
β-strand1113-111971
β-strand112511
β-strand1127-1129359
β-strand1134-1136359
β-strand1137160
β-strand1142160
β-strand1146-115271
α-helix1153-116412
β-strand1170-117232
Chain c: 4 helices, 6 β-strands
ElementResiduesLengthSheet
β-strand11-15598
α-helix23-3412
α-helix35-373
β-strand40-46798
β-strand54-59698
α-helix62-7211
β-strand76199
β-strand81199
β-strand83-86498
α-helix92-987
Chain C: 12 helices, 12 β-strands
ElementResiduesLengthSheet
β-strand8-14761
β-strand18-24761
α-helix28-4013
β-strand44-551262
α-helix61-7010
β-strand73163
α-helix78-814
α-helix82-843
β-strand99-107962
α-helix1121
β-strand113-116464
α-helix117-1193
β-strand121-122262
β-strand129163
α-helix145-1484
β-strand149-153564
β-strand157-1681262
α-helix174-1763
β-strand179-185761
α-helix200-2023
α-helix204-2063
α-helix218-2203
β-strand229-235761
α-helix241-26929
Chain D: 8 helices, 1 β-strand
ElementResiduesLengthSheet
α-helix24-285
β-strand30-31233
α-helix34-5017
α-helix55-584
α-helix59-7012
α-helix77-8913
α-helix94-10310
α-helix108-1147
α-helix124-13714
Chain e: 2 helices, 5 β-strands
ElementResiduesLengthSheet
α-helix19-268
α-helix30-389
β-strand42-46534
β-strand51-56634
β-strand66-761134
β-strand90-1011234
β-strand106-111634
Chain E: 13 helices, 14 β-strands
ElementResiduesLengthSheet
α-helix3-2422
β-strand26165
α-helix30-334
α-helix37-437
β-strand47166
α-helix48-503
β-strand52166
α-helix55-584
β-strand60-63467
β-strand64165
β-strand71-75567
β-strand81-82268
α-helix84-9714
β-strand101-106667
β-strand109-110268
α-helix112-1209
β-strand126-131667
α-helix132-1354
α-helix139-1413
β-strand147-150432
α-helix151-1522
α-helix153-16311
α-helix167-1693
β-strand172-173232
α-helix178-1836
β-strand190-197832
β-strand201-209932

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

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
DNA-directed RNA polymerase subunitAprotein1970Sus scrofa domesticusA0A8D1DPV6
DNA-directed RNA polymerase subunit betaBprotein1174Sus scrofa domesticusA0A0B8RVL1 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB3Cprotein275Sus scrofa domesticusI3LCH3 (AlphaFold model)
RNA polymerase II subunit DDprotein142Sus scrofa domesticusA0A2Y9PTG7 (AlphaFold model)
DNA-directed RNA polymerase II subunit EEprotein210Sus scrofa domesticusI3LSI7
DNA-directed RNA polymerases I, II, and III subunit RPABC2Fprotein127Sus scrofa domesticusA0A4X1VEK9
DNA-directed RNA polymerase subunitGprotein172Sus scrofa domesticusA0A2Y9PKI8
DNA-directed RNA polymerases I, II, and III subunit RPABC3Hprotein150Sus scrofa domesticusA0A6I8N3L3
DNA-directed RNA polymerase II subunit RPB9Iprotein125Sus scrofa domesticusP60899
DNA-directed RNA polymerases I, II, and III subunit RPABC5Jprotein67Sus scrofa domesticusA0A8W4F9W9
DNA-directed RNA polymerase II subunit RPB11-aKprotein117Sus scrofa domesticusF7CZX7
RNA polymerase II, I and III subunit KLprotein58Sus scrofa domesticusA0A4X1TRS6

16 more molecules are not listed.

Sequence of entity 1 (A), FASTA
>9QEQ_1 DNA-directed RNA polymerase subunit (chains A)
MHGGGPPSGDSACPLRTIKRVQFGVLSPDELKRMSVTEGGIKYPETTEGGRPKLGGLMDP
RQGVIERTGRCQTCAGNMTECPGHFGHIELAKPVFHVGFLVKTMKVLRCVCFFCSKLLVD
SNNPKIKDILAKSKGQPKKRLTHVYDLCKGKNICEGGEEMDNKFGVEQPEGDEDLTKEKG
HGGCGRYQPRIRRSGLELYAEWKHVNEDSQEKKILLSPERVHEIFKRISDEECFVLGMEP
RYARPEWMIVTVLPVPPLSVRPAVVMQGSARNQDDLTHKLADIVKINNQLRRNEQNGAAA
HVIAEDVKLLQFHVATMVDNELPGLPRAMQKSGRPLKSLKQRLKGKEGRVRGNLMGKRVD
FSARTVITPDPNLSIDQVGVPRSIAANMTFAEIVTPFNIDRLQELVRRGNSQYPGAKYII
RDNGDRIDLRFHPKPSDLHLQTGYKVERHMCDGDIVIFNRQPTLHKMSMMGHRVRILPWS
TFRLNLSVTTPYNADFDGDEMNLHLPQSLETRAEIQELAMVPRMIVTPQSNRPVMGIVQD
TLTAVRKFTKRDVFLERGEVMNLLMFLSTWDGKVPQPAILKPRPLWTGKQIFSLIIPGHI
NCIRTHSTHPDDEDSGPYKHISPGDTKVVVENGELIMGILCKKSLGTSAGSLVHISYLEM
GHDITRLFYSNIQTVINNWLLIEGHTIGIGDSIADSKTYQDIQNTIKKAKQDVIEVIEKA
HNNELEPTPGNTLRQTFENQVNRILNDARDKTGSSAQKSLSEYNNFKSMVVSGAKGSKIN
ISQVIAVVGQQNVEGKRIPFGFKHRTLPHFIKDDYGPESRGFVENSYLAGLTPTEFFFHA
MGGREGLIDTAVKTAETGYIQRRLIKSMESVMVKYDATVRNSINQVVQLRYGEDGLAGES
VEFQNLATLKPSNKAFEKKFRFDYTNERALRRTLQEDLVKDVLSNAHIQNELEREFERMR
EDREVLRVIFPTGDSKVVLPCNLLRMIWNAQKIFHINPRLPSDLHPIKVVEGVKELSKKL
VIVNGDDPLSRQAQENATLLFNIHLRSTLCSRRMAEEFRLSGEAFDWLLGEIESKFNQAI
AHPGEMVGALAAQSLGEPATQMTLNTFHYAGVSAKNVTLGVPRLKELINISKKPKTPSLT
VFLLGQSARDAERAKDILCRLEHTTLRKVTANTAIYYDPNPQSTVVAEDQEWVNVYYEMP
DFDVARISPWLLRVELDRKHMTDRKLTMEQIAEKINAGFGDDLNCIFNDDNAEKLVLRIR
IMNSDENKMQEEEEVVDKMDDDVFLRCIESNMLTDMTLQGIEQISKVYMHLPQTDNKKKI
IITEDGEFKALQEWILETDGVSLMRVLSEKDVDPVRTTSNDIVEIFTVLGIEAVRKALER
ELYHVISFDGSYVNYRHLALLCDTMTCRGHLMAITRHGVNRQDTGPLMKCSFEETVDVLM
EAAAHGESDPMKGVSENIMLGQLAPAGTGCFDLLLDAEKCKYGMEIPTNIPGLGAAGPTG
MFFGSAPSPMGGISPAMTPWNQGATPAYGAWSPSVGSGMTPGAAGFSPSAASDASGFSPG
YSPAWSPTPGSPGSPGPSSPYIPSPGGAMSPSYSPTSPAYEPRSPGGYTPQSPSYSPTSP
SYSPTSPSYSPTSPNYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSP
TSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPS
YSPTSPNYSPTSPNYTPTSPSYSPTSPSYSPTSPNYTPTSPNYSPTSPSYSPTSPSYSPT
SPSYSPSSPRYTPQSPTYTPSSPSYSPSSPSYSPTSPKYTPTSPSYSPSSPEYTPTSPKY
SPTSPKYSPTSPKYSPTSPTYSPTTPKYSPTSPTYSPTSPVYTPTSPKYSPTSPTYSPTS
PKYSPTSPTYSPTSPKGSTYSPTSPGYSPTSPTYSLTSPAISPDDSDEEN
Sequence of entity 2 (B), FASTA
>9QEQ_2 DNA-directed RNA polymerase subunit beta (chains B)
MYDADEDMQYDEDDDEITPDLWQEACWIVISSYFDEKGLVRQQLDSFDEFIQMSVQRIVE
DAPPIDLQAEAQHASGEVEEPPRYLLKFEQIYLSKPTHWERDGAPSPMMPNEARLRNLTY
SAPLYVDITKTVIKEGEEQLQTQHQKTFIGKIPIMLRSTYCLLNGLTDRDLCELNECPLD
PGGYFIINGSEKVLIAQEKMATNTVYVFAKKDSKYAYTGECRSCLENSSRPTSTIWVSML
ARGGQGAKKSAIGQRIVATLPYIKQEVPIIIVFRALGFVSDRDILEHIIYDFEDPEMMEM
VKPSLDEAFVIQEQNVALNFIGSRGAKPGVTKEKRIKYAKEVLQKEMLPHVGVSDFCETK
KAYFLGYMVHRLLLAALGRRELDDRDHYGNKRLDLAGPLLAFLFRGMFKNLLKEVRIYAQ
KFIDRGKDFNLELAIKTRIISDGLKYSLATGNWGDQKKAHQARAGVSQVLNRLTFASTLS
HLRRLNSPIGRDGKLAKPRQLHNTLWGMVCPAETPEGHAVGLVKNLALMAYISVGSQPSP
ILEFLEEWSMENLEEISPAAIADATKIFVNGCWVGIHKDPEQLMNTLRKLRRQMDIIVSE
VSMIRDIREREIRIYTDAGRICRPLLIVEKQKLLLKKRHIDQLKEREYNNYSWQDLVASG
VVEYIDTLEEETVMLAMTPDDLQEKEVAYCSTYTHCEIHPSMILGVCASIIPFPDHNQSP
RNTYQSAMGKQAMGVYITNFHVRMDTLAHVLYYPQKPLVTTRSMEYLRFRELPAGINSIV
AIASYTGYNQEDSVIMNRSAVDRGFFRSVFYRSYKEQESKKGFDQEEVFEKPTRETCQGM
RHAIYDKLDDDGLIAPGVRVSGDDVIIGKTVTLPENEDELEGTNRRYTKRDCSTFLRTSE
TGIVDQVMVTLNQEGYKFCKIRVRSVRIPQIGDKFASRHGQKGTCGIQYRQEDMPFTCEG
ITPDIIINPHAIPSRMTIGHLIECLQGKVSANKGEIGDATPFNDAVNVQKISNLLSDYGY
HLRGNEVLYNGFTGRKITSQIFIGPTYYQRLKHMVDDKIHSRARGPIQILNRQPMEGRSR
DGGLRFGEMERDCQIAHGAAQFLRERLFEASDPYQVHVCNLCGIMAIANTRTHTYECRGC
RNKTQISLVRMPYACKLLFQELMSMSIAPRMMSV
Sequence of entity 3 (C), FASTA
>9QEQ_3 DNA-directed RNA polymerase II subunit RPB3 (chains C)
MPYANQPTVRITELTDENVKFIIENTDLAVANSIRRVFIAEVPIIAIDWVQIDANSSVLH
DEFIAHRLGLIPLTSDDIVDKLQYSRDCTCEEFCPECSVEFTLDVRCNEDQTRHVTSRDL
ISNSPRVIPVTSRNRDNDPSDYVEQDDILIVKLRKGQELRLRAYAKKGFGKEHAKWNPTA
GVAFEYDPDNALRHTVYPKPEEWPKSEYSELDEDESQAPYDPNGKPERFYYNVESCGSLR
PETIVLSALSGLKKKLSDLQTQLSHEIQSDVLTIN
Sequence of entity 4 (D), FASTA
>9QEQ_4 RNA polymerase II subunit D (chains D)
MAAGGSDPRAGDVEEDASQLIFPKEFETAETLLNSEVHMLLEHRKQQNESAEDEQELSEV
FMKTLNYTARFSRFKNRETIASVRSLLLQKKLHKFELACLANLCPETAEESKALIPSLEG
RFEDEELQQILDDIQTKRSFQY
Sequence of entity 5 (E), FASTA
>9QEQ_5 DNA-directed RNA polymerase II subunit E (chains E)
MDDEEETYRLWKIRKTIMQLCHDRGYLVTQDELDQTLEEFKAQFGDKPSEGRPRRTDLTV
LVAHNDDPTDQMFVFFPEEPKVGIKTIKVYCQRMQEENITRALIVVQQGMTPSAKQSLVD
MAPKYILEQFLQQELLINITEHELVPEHVVMTKEEVTELLARYKLRENQLPRIQAGDPVA
RYFGIKRGQVVKIIRPSETAGRYITYRLVQ
Sequence of entity 6 (F), FASTA
>9QEQ_6 DNA-directed RNA polymerases I, II, and III subunit RPABC2 (chains F)
MSDNEDNFDGDDFDDVEEDEGLDDLENAEEEGQENVEILPSGERPQANQKRITTPYMTKY
ERARVLGTRALQIAMCAPVMVELEGETDPLLIAMKELKARKIPIIIRRYLPDGSYEDWGV
DELIISD
Sequence of entity 7 (G), FASTA
>9QEQ_7 DNA-directed RNA polymerase subunit (chains G)
MFYHISLEHEILLHPRYFGPNLLNTVKQKLFTEVEGTCTGKYGFVIAVTTIDNIGAGVIQ
PGRGFVLYPVKYKAIVFRPFKGEVVDAVVTQVNKVGLFTEIGPMSCFISRHSIPSEMEFD
PNSNPPCYKTMDEDIVIQQDDEIRLKIVGTRVDKNDIFAIGSLMDDYLGLVS
Sequence of entity 8 (H), FASTA
>9QEQ_8 DNA-directed RNA polymerases I, II, and III subunit RPABC3 (chains H)
MAGILFEDIFDVKDIDPEGKKFDRVSRLHCESESFKMDLILDVNIQIYPVDLGDKFRLVI
ASTLYEDGTLDDGEYNPTDDRPSRADQFEYVMYGKVYRIEGDETSTEAATRLSAYVSYGG
LLMRLQGDANNLHGFEVDSRVYLLMKKLAF
Sequence of entity 9 (I), FASTA
>9QEQ_9 DNA-directed RNA polymerase II subunit RPB9 (chains I)
MEPDGTYEPGFVGIRFCQECNNMLYPKEDKENRILLYACRNCDYQQEADNSCIYVNKITH
EVDELTQIIADVSQDPTLPRTEDHPCQKCGHKEAVFFQSHSARAEDAMRLYYVCTAPHCG
HRWTE
Sequence of entity 10 (J), FASTA
>9QEQ_10 DNA-directed RNA polymerases I, II, and III subunit RPABC5 (chains J)
MIIPVRCFTCGKIVGNKWEAYLGLLQAEYTEGDALDALGLKRYCCRRMLLAHVDLIEKLL
NYAPLEK
Sequence of entity 11 (K), FASTA
>9QEQ_11 DNA-directed RNA polymerase II subunit RPB11-a (chains K)
MNAPPAFESFLLFEGEKKITINKDTKVPNACLFTINKEDHTLGNIIKSQLLKDPQVLFAG
YKVPHPLEHKIIIRVQTTPDYSPQEAFTNAITDLISELSLLEERFRVAIKDKQEGIE
Sequence of entity 12 (L), FASTA
>9QEQ_12 RNA polymerase II, I and III subunit K (chains L)
MDTQKDVQPPKQQPMIYICGECHTENEIKSRDPIRCRECGYRIMYKKRTKRLVVFDAR

Ligands and cofactors

IDNameFormulaCopies
MGMagnesium ionMg1
ZNZinc ionZn8

Primary citation

Structure of a transcribing Pol II-DSIF-SPT6-U1 snRNP complex. Zhang, L., Batters, C., Aibara, S. et al. Nat Commun (2025) 16:5823-5823. DOI 10.1038/s41467-025-60979-9 · PubMed

Other PDB entries of the same protein (UniProt A0A8D1DPV6), best resolution first:

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