9EI4: Human RNA polymerase II Elongation Complex

Cryo-EM structure of Human RNA polymerase II Elongation Complex bound to the RECQL5 helicase in the presence of ADP. Determined by electron microscopy at 3.7 Å resolution. Released 5 Mar 2025.

Method
Electron microscopy
Resolution
3.7 Å
Organism
Homo sapiens
Chains
16
Atoms
35,854
Mol. weight
655.67 kDa
Ligands
ADP, MG, ZN
Released
5 Mar 2025

Explore 9EI4 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9EI4 contains 189 α-helices and 239 β-strands across 13 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: 72 helices, 69 β-strands

ElementResiduesLengthSheet
β-strand1511
β-strand20-2232
β-strand2513
α-helix28-347
β-strand4714
β-strand5214
α-helix531
β-strand86-9495
α-helix97-993
α-helix100-10910
β-strand11016
α-helix1111
β-strand11716
α-helix124-1329
α-helix137-14812
β-strand15317
β-strand18717
β-strand190-19458
β-strand197-20158
β-strand213-21648
α-helix218-2269
α-helix230-2345
α-helix245-2484
β-strand250-25565
β-strand263-26649
β-strand270-27349
α-helix2741
α-helix275-29622
α-helix302-31817
α-helix326-3283
β-strand329110
β-strand335110
α-helix339-3424
α-helix350-3545
β-strand357-358211
β-strand362-369812
β-strand377-380412
α-helix382-3876
β-strand389-393513
α-helix396-3983
α-helix399-40810
β-strand416-420513
β-strand426-428313
β-strand445-449513
α-helix4501
β-strand455-459512
α-helix466-4683
β-strand469-476812
β-strand482-484312
α-helix486-4883
α-helix489-4924
β-strand500-504512
α-helix509-51810
β-strand520112
α-helix522-5243
β-strand526-527214
β-strand532-533214
α-helix540-5489
β-strand554-555215
α-helix557-5659
α-helix573-5775
β-strand579116
β-strand585116
β-strand586-587215
α-helix588-5958
β-strand602-603217
β-strand629-631317
β-strand634-637417
α-helix642-6454
α-helix652-6598
α-helix662-68322
β-strand686118
α-helix689-6913
α-helix696-72126
α-helix726-7283
α-helix733-75826
α-helix765-7728
α-helix778-7814
α-helix782-7865
β-strand789-790219
β-strand792-793220
β-strand796-797220
α-helix817-8204
β-strand822-823219
α-helix833-86836
β-strand872-873221
β-strand879-880221
β-strand886-888321
α-helix891-8933
β-strand896122
β-strand902-905423
α-helix913-9208
α-helix929-9335
β-strand934124
α-helix936-9449
α-helix946-96924
β-strand976-979423
α-helix983-99311
β-strand1002124
α-helix1006-101914
α-helix1028-103710
α-helix1039-104810
α-helix1051-10566
α-helix1062-107918
β-strand1080122
α-helix1087-110115
α-helix1121-11288
β-strand1138-1142525
α-helix1146-11483
α-helix1151-11599
β-strand1164126
β-strand1170-1177827
α-helix1187-11893
α-helix1191-11966
α-helix1202-12043
β-strand1210-1216727
α-helix12171
α-helix1218-12236
α-helix1228-123912
β-strand1246-1247227
β-strand1255-1259527
α-helix1281-129515
β-strand1299126
β-strand1307125
β-strand1310125
β-strand1319-1322428
β-strand1328-1331428
β-strand1335-1339525
α-helix1343-13486
β-strand1358-1359225
α-helix1362-13687
α-helix1371-138717
α-helix1395-140511
β-strand1408-1409229
α-helix1416-14194
α-helix1426-14294
α-helix1436-144510
β-strand1448-144922
α-helix1455-14595
β-strand1471-1475530
α-helix1477-14815
Chain B: 49 helices, 81 β-strands
ElementResiduesLengthSheet
α-helix22-3615
α-helix41-499
α-helix50-545
α-helix55-617
α-helix63-642
β-strand65-67331
β-strand83-931131
β-strand97132
β-strand107132
α-helix110-1156
β-strand121-1331331
β-strand139-1531531
α-helix168-1736
β-strand185-187333
β-strand190-193433
β-strand196-200534
α-helix2011
β-strand205-208435
β-strand218-221435
β-strand235-239535
β-strand256-259435
β-strand263136
β-strand267-268235
α-helix269-2757
α-helix281-2888
α-helix295-30713
α-helix314-3229
β-strand325136
α-helix332-34110
α-helix342-3465
α-helix358-37720
α-helix381-3844
α-helix388-3903
β-strand391-394434
α-helix396-42328
α-helix431-4344
α-helix437-45014
β-strand453137
α-helix456-4583
β-strand463137
β-strand468-469233
α-helix475-4828
β-strand484-486334
α-helix497-4993
α-helix503-5053
β-strand509138
β-strand523-525334
β-strand526138
β-strand527139
β-strand531-532240
α-helix539-5479
β-strand551141
α-helix558-5614
β-strand565-569541
β-strand572-577641
α-helix580-59314
β-strand602-606541
β-strand611-615541
β-strand621-627740
β-strand628-629242
β-strand632-633242
α-helix637-6448
α-helix653-6586
β-strand662-666540
β-strand675-676240
α-helix679-6824
β-strand695-696240
α-helix700-7023
β-strand705139
α-helix714-7163
α-helix719-72911
β-strand732143
β-strand748-751444
β-strand755145
β-strand759-760246
α-helix762-7654
β-strand776-782747
β-strand783118
β-strand794-797447
α-helix798-8025
β-strand808-8171044
β-strand820148
β-strand826148
β-strand827-828249
α-helix831-8333
β-strand839149
β-strand853150
α-helix854-8552
β-strand859-860244
β-strand865-867350
β-strand870-872349
β-strand889-891349
β-strand894-895250
α-helix896-8972
β-strand902-9111044
β-strand917-9271144
β-strand934-937447
β-strand942-949847
β-strand956151
β-strand965-967347
α-helix971-9755
α-helix979-99315
β-strand997-998246
α-helix1008-10169
β-strand1025-1026247
β-strand1029151
α-helix10351
β-strand1036151
α-helix10371
β-strand1041-10501047
β-strand1051143
α-helix1055-10573
β-strand1060-1062312
β-strand1067152
β-strand1074152
α-helix10751
β-strand1085-1086211
α-helix1088-109710
α-helix1100-11045
α-helix1105-11095
β-strand1115-111951
β-strand112511
β-strand1127-1129353
β-strand1134-1136353
β-strand1137154
β-strand1142154
β-strand1146-115051
α-helix1153-116311
β-strand116713
β-strand1170-117232
Chain C: 9 helices, 12 β-strands
ElementResiduesLengthSheet
β-strand8-14755
β-strand18-24755
α-helix28-3912
β-strand44-551256
α-helix61-699
β-strand99-106856
α-helix1121
β-strand113-116457
α-helix117-1193
β-strand121-122256
β-strand149-153557
β-strand158-1681156
α-helix174-1763
β-strand179-180255
β-strand185155
α-helix200-2023
α-helix204-2063
α-helix218-2203
β-strand229-230255
β-strand233-235355
α-helix241-27030
Chain D: 9 helices, 1 β-strand
ElementResiduesLengthSheet
α-helix24-263
β-strand30130
α-helix34-4916
α-helix55-573
α-helix59-7012
α-helix77-8812
α-helix94-10310
α-helix109-1146
α-helix116-1183
α-helix124-13613
Chain E: 10 helices, 12 β-strands
ElementResiduesLengthSheet
α-helix4-2320
β-strand26158
α-helix30-334
α-helix37-448
β-strand60-62359
β-strand64158
β-strand72-74359
β-strand81-82260
α-helix84-9613
β-strand101-106659
β-strand109-110260
α-helix112-1209
β-strand127-131559
α-helix133-1353
α-helix139-1413
β-strand147-150429
α-helix153-16210
α-helix167-1693
β-strand172-173229
α-helix178-1836
β-strand190-195629
β-strand203-209729
Chain F: 5 helices, 4 β-strands
ElementResiduesLengthSheet
β-strand56161
α-helix57-582
α-helix59-7416
α-helix77-793
α-helix89-9911
β-strand104-109630
β-strand115-119530
β-strand124161
α-helix1251
Chain G: 3 helices, 14 β-strands
ElementResiduesLengthSheet
β-strand2-131230
α-helix15-173
α-helix22-3413
β-strand38-39230
β-strand43-541230
β-strand57-59330
α-helix601
β-strand66-771230
β-strand81162
β-strand84-92962
β-strand96-101662
β-strand104-109662
β-strand118-119263
β-strand128-129263
β-strand137163
β-strand142-1511062
β-strand158-162562
Chain H: 1 helix, 11 β-strands
ElementResiduesLengthSheet
β-strand4-12916
β-strand22116
β-strand25-32816
β-strand38-44716
β-strand55-61716
β-strand90-1011216
β-strand104164
β-strand107164
β-strand111-118816
β-strand121-128816
α-helix130-1323
β-strand140-148916

5 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 II subunit RPB1Aprotein1970Homo sapiensP24928 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB2Bprotein1174Homo sapiensP30876 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB3Cprotein275Homo sapiensP19387 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB4Dprotein142Homo sapiensO15514 (AlphaFold model)
DNA-directed RNA polymerases I, II, and III subunit RPABC1Eprotein210Homo sapiensP19388
DNA-directed RNA polymerases I, II, and III subunit RPABC2Fprotein127Homo sapiensP61218
DNA-directed RNA polymerase II subunit RPB7Gprotein172Homo sapiensP62487
DNA-directed RNA polymerases I, II, and III subunit RPABC3Hprotein150Homo sapiensP52434
DNA-directed RNA polymerase II subunit RPB9Iprotein125Homo sapiensP36954
DNA-directed RNA polymerases I, II, and III subunit RPABC5Jprotein67Homo sapiensP62875
DNA-directed RNA polymerase II subunit RPB11-aKprotein117Homo sapiensP52435
DNA-directed RNA polymerases I, II, and III subunit RPABC4Lprotein58Homo sapiensP53803

4 more molecules are not listed.

Sequence of entity 1 (A), FASTA
>9EI4_1 DNA-directed RNA polymerase II subunit RPB1 (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
SPSYSPSSPRYTPQSPTYTPSSPSYSPSSPSYSPASPKYTPTSPSYSPSSPEYTPTSPKY
SPTSPKYSPTSPKYSPTSPTYSPTTPKYSPTSPTYSPTSPVYTPTSPKYSPTSPTYSPTS
PKYSPTSPTYSPTSPKGSTYSPTSPGYSPTSPTYSLTSPAISPDDSDEEN
Sequence of entity 2 (B), FASTA
>9EI4_2 DNA-directed RNA polymerase II subunit RPB2 (chains B)
MYDADEDMQYDEDDDEITPDLWQEACWIVISSYFDEKGLVRQQLDSFDEFIQMSVQRIVE
DAPPIDLQAEAQHASGEVEEPPRYLLKFEQIYLSKPTHWERDGAPSPMMPNEARLRNLTY
SAPLYVDITKTVIKEGEEQLQTQHQKTFIGKIPIMLRSTYCLLNGLTDRDLCELNECPLD
PGGYFIINGSEKVLIAQEKMATNTVYVFAKKDSKYAYTGECRSCLENSSRPTSTIWVSML
ARGGQGAKKSAIGQRIVATLPYIKQEVPIIIVFRALGFVSDRDILEHIIYDFEDPEMMEM
VKPSLDEAFVIQEQNVALNFIGSRGAKPGVTKEKRIKYAKEVLQKEMLPHVGVSDFCETK
KAYFLGYMVHRLLLAALGRRELDDRDHYGNKRLDLAGPLLAFLFRGMFKNLLKEVRIYAQ
KFIDRGKDFNLELAIKTRIISDGLKYSLATGNWGDQKKAHQARAGVSQVLNRLTFASTLS
HLRRLNSPIGRDGKLAKPRQLHNTLWGMVCPAETPEGHAVGLVKNLALMAYISVGSQPSP
ILEFLEEWSMENLEEISPAAIADATKIFVNGCWVGIHKDPEQLMNTLRKLRRQMDIIVSE
VSMIRDIREREIRIYTDAGRICRPLLIVEKQKLLLKKRHIDQLKEREYNNYSWQDLVASG
VVEYIDTLEEETVMLAMTPDDLQEKEVAYCSTYTHCEIHPSMILGVCASIIPFPDHNQSP
RNTYQSAMGKQAMGVYITNFHVRMDTLAHVLYYPQKPLVTTRSMEYLRFRELPAGINSIV
AIASYTGYNQEDSVIMNRSAVDRGFFRSVFYRSYKEQESKKGFDQEEVFEKPTRETCQGM
RHAIYDKLDDDGLIAPGVRVSGDDVIIGKTVTLPENEDELESTNRRYTKRDCSTFLRTSE
TGIVDQVMVTLNQEGYKFCKIRVRSVRIPQIGDKFASRHGQKGTCGIQYRQEDMPFTCEG
ITPDIIINPHAIPSRMTIGHLIECLQGKVSANKGEIGDATPFNDAVNVQKISNLLSDYGY
HLRGNEVLYNGFTGRKITSQIFIGPTYYQRLKHMVDDKIHSRARGPIQILNRQPMEGRSR
DGGLRFGEMERDCQIAHGAAQFLRERLFEASDPYQVHVCNLCGIMAIANTRTHTYECRGC
RNKTQISLVRMPYACKLLFQELMSMSIAPRMMSV
Sequence of entity 3 (C), FASTA
>9EI4_3 DNA-directed RNA polymerase II subunit RPB3 (chains C)
MPYANQPTVRITELTDENVKFIIENTDLAVANSIRRVFIAEVPIIAIDWVQIDANSSVLH
DEFIAHRLGLIPLISDDIVDKLQYSRDCTCEEFCPECSVEFTLDVRCNEDQTRHVTSRDL
ISNSPRVIPVTSRNRDNDPNDYVEQDDILIVKLRKGQELRLRAYAKKGFGKEHAKWNPTA
GVAFEYDPDNALRHTVYPKPEEWPKSEYSELDEDESQAPYDPNGKPERFYYNVESCGSLR
PETIVLSALSGLKKKLSDLQTQLSHEIQSDVLTIN
Sequence of entity 4 (D), FASTA
>9EI4_4 DNA-directed RNA polymerase II subunit RPB4 (chains D)
MAAGGSDPRAGDVEEDASQLIFPKEFETAETLLNSEVHMLLEHRKQQNESAEDEQELSEV
FMKTLNYTARFSRFKNRETIASVRSLLLQKKLHKFELACLANLCPETAEESKALIPSLEG
RFEDEELQQILDDIQTKRSFQY
Sequence of entity 5 (E), FASTA
>9EI4_5 DNA-directed RNA polymerases I, II, and III subunit RPABC1 (chains E)
MDDEEETYRLWKIRKTIMQLCHDRGYLVTQDELDQTLEEFKAQFGDKPSEGRPRRTDLTV
LVAHNDDPTDQMFVFFPEEPKVGIKTIKVYCQRMQEENITRALIVVQQGMTPSAKQSLVD
MAPKYILEQFLQQELLINITEHELVPEHVVMTKEEVTELLARYKLRENQLPRIQAGDPVA
RYFGIKRGQVVKIIRPSETAGRYITYRLVQ
Sequence of entity 6 (F), FASTA
>9EI4_6 DNA-directed RNA polymerases I, II, and III subunit RPABC2 (chains F)
MSDNEDNFDGDDFDDVEEDEGLDDLENAEEEGQENVEILPSGERPQANQKRITTPYMTKY
ERARVLGTRALQIAMCAPVMVELEGETDPLLIAMKELKARKIPIIIRRYLPDGSYEDWGV
DELIITD
Sequence of entity 7 (G), FASTA
>9EI4_7 DNA-directed RNA polymerase II subunit RPB7 (chains G)
MFYHISLEHEILLHPRYFGPNLLNTVKQKLFTEVEGTCTGKYGFVIAVTTIDNIGAGVIQ
PGRGFVLYPVKYKAIVFRPFKGEVVDAVVTQVNKVGLFTEIGPMSCFISRHSIPSEMEFD
PNSNPPCYKTMDEDIVIQQDDEIRLKIVGTRVDKNDIFAIGSLMDDYLGLVS
Sequence of entity 8 (H), FASTA
>9EI4_8 DNA-directed RNA polymerases I, II, and III subunit RPABC3 (chains H)
MAGILFEDIFDVKDIDPEGKKFDRVSRLHCESESFKMDLILDVNIQIYPVDLGDKFRLVI
ASTLYEDGTLDDGEYNPTDDRPSRADQFEYVMYGKVYRIEGDETSTEAATRLSAYVSYGG
LLMRLQGDANNLHGFEVDSRVYLLMKKLAF
Sequence of entity 9 (I), FASTA
>9EI4_9 DNA-directed RNA polymerase II subunit RPB9 (chains I)
MEPDGTYEPGFVGIRFCQECNNMLYPKEDKENRILLYACRNCDYQQEADNSCIYVNKITH
EVDELTQIIADVSQDPTLPRTEDHPCQKCGHKEAVFFQSHSARAEDAMRLYYVCTAPHCG
HRWTE
Sequence of entity 10 (J), FASTA
>9EI4_10 DNA-directed RNA polymerases I, II, and III subunit RPABC5 (chains J)
MIIPVRCFTCGKIVGNKWEAYLGLLQAEYTEGDALDALGLKRYCCRRMLLAHVDLIEKLL
NYAPLEK
Sequence of entity 11 (K), FASTA
>9EI4_11 DNA-directed RNA polymerase II subunit RPB11-a (chains K)
MNAPPAFESFLLFEGEKKITINKDTKVPNACLFTINKEDHTLGNIIKSQLLKDPQVLFAG
YKVPHPLEHKIIIRVQTTPDYSPQEAFTNAITDLISELSLLEERFRVAIKDKQEGIE
Sequence of entity 12 (L), FASTA
>9EI4_12 DNA-directed RNA polymerases I, II, and III subunit RPABC4 (chains L)
MDTQKDVQPPKQQPMIYICGECHTENEIKSRDPIRCRECGYRIMYKKRTKRLVVFDAR

Ligands and cofactors

IDNameFormulaCopies
ADPAdenosine-5'-diphosphateC10 H15 N5 O10 P21
MGMagnesium ionMg1
ZNZinc ionZn8

Primary citation

Structural insights into transcriptional regulation by the helicase RECQL5. Florez Ariza, A.J., Lue, N.Z., Grob, P. et al. Nat Struct Mol Biol (2025) 32:1721-1730. DOI 10.1038/s41594-025-01611-8 · PubMed

Other PDB entries of the same protein (UniProt P24928 (AlphaFold model), which also has an AlphaFold model), best resolution first:

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