9QEB: DNA-directed RNA polymerase II subunit RPB1

CryoEM structure of transcribing RNA polymerase II elongation complex. Determined by electron microscopy at 2.26 Å resolution. Released 27 May 2026.

Method
Electron microscopy
Resolution
2.26 Å
Organism
Saccharomyces cerevisiae
Chains
15
Atoms
33,807
Mol. weight
545.17 kDa
Ligands
ATP, ZN, MG
Released
27 May 2026

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

Secondary structure: helices and β-sheets

9QEB contains 194 α-helices and 226 β-strands across 12 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, 67 β-strands

ElementResiduesLengthSheet
α-helix101
β-strand1111
β-strand16-1832
β-strand2113
α-helix24-307
β-strand3414
α-helix56-583
β-strand82-91104
α-helix93-953
α-helix96-10510
β-strand10615
β-strand11315
α-helix120-1267
α-helix131-14212
β-strand14716
β-strand17016
α-helix171-1722
β-strand175-17737
β-strand180-18347
β-strand200-20237
α-helix2031
α-helix204-2129
α-helix216-2216
β-strand22518
β-strand22918
α-helix231-2344
β-strand235-24174
β-strand25219
β-strand25619
α-helix261-28222
α-helix289-30416
α-helix312-3143
β-strand315110
β-strand321110
α-helix3221
α-helix325-3295
α-helix335-3395
β-strand343-345311
β-strand348-355812
β-strand363-367512
α-helix368-3736
β-strand375-379513
α-helix385-39410
β-strand402-406513
β-strand412-414313
β-strand431-435513
α-helix4361
β-strand441-445512
α-helix452-4543
β-strand455-463912
β-strand468-470312
α-helix472-4743
α-helix475-4784
β-strand486-490512
α-helix495-50410
β-strand506112
α-helix507-5104
β-strand512-513214
β-strand518-519214
α-helix525-53511
β-strand540-541215
α-helix543-55210
α-helix560-5634
β-strand565116
β-strand571116
β-strand572-573215
α-helix574-5818
α-helix5831
β-strand588-590317
β-strand605-608417
β-strand611-614417
α-helix619-6224
α-helix629-6368
α-helix639-66022
α-helix666-6694
α-helix673-69826
α-helix710-73627
α-helix742-7498
α-helix755-7584
α-helix759-7635
β-strand766-767218
β-strand769-770219
β-strand773-774219
β-strand779120
β-strand782120
β-strand799-800218
α-helix810-82516
α-helix826-8305
α-helix831-84515
β-strand849-850221
β-strand856-857221
β-strand863-865321
α-helix868-8703
β-strand873122
α-helix875-8773
β-strand878-882523
α-helix890-8978
α-helix907-9093
α-helix910-9123
α-helix916-9194
α-helix924-94623
β-strand953-957523
α-helix960-97112
α-helix978-9792
α-helix983-99412
α-helix1005-101410
α-helix1016-102510
α-helix1028-10292
α-helix1030-10356
α-helix1039-105618
β-strand1057122
α-helix1058-10592
α-helix1064-108118
α-helix1083-10864
α-helix10921
α-helix1093-10975
α-helix1098-11069
β-strand1116-1119424
β-strand1120125
α-helix1131-11388
β-strand1141126
α-helix1143-11464
β-strand1147-1153727
α-helix1167-11726
β-strand1192-1197627
α-helix1199-12057
α-helix1209-122012
β-strand1224-1228527
β-strand1236-1242727
α-helix1257-126913
β-strand1274126
β-strand1282-12921124
β-strand1298-13081124
α-helix1313-13164
β-strand1322125
β-strand1328-1329224
α-helix1332-13398
α-helix1341-135616
α-helix1365-137410
α-helix1396-14005
α-helix1405-141511
α-helix14171
β-strand1418-141922
α-helix1424-14296
α-helix1437-14393
β-strand1441-1445528
α-helix1447-14515
Chain B: 60 helices, 81 β-strands
ElementResiduesLengthSheet
α-helix24-263
α-helix27-293
α-helix30-367
α-helix45-539
α-helix54-585
α-helix59-657
β-strand69-73529
α-helix80-823
β-strand84-971429
β-strand101-103330
β-strand109-111330
α-helix114-1207
β-strand125-1401629
β-strand150-155629
α-helix157-1593
α-helix161-1633
β-strand165-171729
α-helix186-1916
β-strand203-205331
β-strand208-210331
β-strand211-212232
α-helix2131
β-strand214-218533
α-helix2191
β-strand224-227434
β-strand234-239634
β-strand242135
β-strand252135
β-strand255-260634
β-strand268-272534
β-strand276136
β-strand280-281234
α-helix282-2898
α-helix294-3029
α-helix309-32012
α-helix327-33610
β-strand338136
α-helix345-35410
α-helix355-3595
α-helix371-38919
α-helix394-3974
α-helix401-4033
β-strand404-407433
α-helix409-43729
α-helix440-4423
α-helix444-4474
α-helix450-46314
β-strand466137
β-strand476137
α-helix4771
β-strand480-481232
α-helix488-4947
β-strand497-499333
α-helix510-5123
α-helix514-5152
α-helix516-5183
β-strand522138
β-strand536-538333
β-strand539138
β-strand544-545239
α-helix552-5609
β-strand564-565240
α-helix566-5683
α-helix5771
β-strand578-582540
β-strand585-590640
α-helix593-60614
β-strand614-618540
β-strand623-627540
β-strand633-639739
β-strand640-641241
β-strand650-651241
α-helix655-66612
α-helix681-6866
β-strand690-694539
α-helix697-7004
β-strand703-704239
α-helix707-7093
α-helix728-7292
β-strand740-741239
α-helix745-7484
α-helix753-7553
α-helix759-7613
β-strand762142
α-helix764-77411
β-strand777143
α-helix785-7873
β-strand792-796544
β-strand800145
β-strand804-805246
α-helix809-8124
β-strand821-827747
β-strand838-842547
α-helix843-8475
β-strand853-861944
β-strand874148
β-strand898149
β-strand904-906344
β-strand910-912349
β-strand914148
β-strand915150
β-strand936150
β-strand939-940249
α-helix941-9422
β-strand946-9561144
β-strand962-9721144
β-strand979-981347
β-strand987-994847
β-strand1001-1002251
β-strand1010-1012347
α-helix1016-10205
β-strand1022142
α-helix1023-103816
α-helix1040-10412
β-strand1042-1043246
α-helix1052-106110
β-strand1069-1070247
β-strand1072-1073251
α-helix10791
β-strand1080151
α-helix10811
β-strand1085-10941047
β-strand1095143
α-helix1099-11013
β-strand1104-1106312
β-strand1111152
β-strand1118152
α-helix11191
α-helix11271
β-strand1128-1130311
α-helix1132-114110
α-helix1144-11529
β-strand1158-116361
β-strand1172-1174353
β-strand1179-1181353
β-strand1182154
β-strand1187154
β-strand1191-119661
α-helix1198-120912
β-strand121213
β-strand1215-121732
Chain C: 13 helices, 12 β-strands
ElementResiduesLengthSheet
β-strand7-13755
β-strand17-23755
α-helix27-3913
β-strand43-541256
α-helix60-689
β-strand72157
α-helix77-793
β-strand96-104956
α-helix1101
β-strand111-114458
α-helix115-1173
β-strand119-120256
α-helix1311
β-strand132157
α-helix1331
β-strand143-147558
β-strand152-1621156
α-helix168-1703
β-strand176-180555
α-helix197-2004
α-helix202-2043
α-helix205-2095
α-helix217-2193
β-strand228-233655
α-helix240-26526
Chain D: 8 helices, 6 β-strands
ElementResiduesLengthSheet
β-strand6128
β-strand23159
β-strand28159
β-strand36-38360
β-strand44-46360
β-strand48-50328
α-helix52-7221
α-helix119-13416
α-helix139-15012
α-helix157-16812
α-helix174-1829
α-helix188-1947
α-helix196-1983
α-helix204-21714
Chain E: 12 helices, 14 β-strands
ElementResiduesLengthSheet
α-helix6-2520
β-strand28161
α-helix32-354
α-helix39-435
β-strand47162
β-strand53162
α-helix55-584
β-strand60-62363
β-strand64161
α-helix66-716
β-strand78-82563
β-strand88164
α-helix92-10312
β-strand107-112663
β-strand116164
α-helix118-1236
β-strand131-136663
α-helix138-1403
α-helix144-1463
β-strand152-155465
α-helix158-16710
α-helix172-1743
β-strand177-178265
α-helix183-1886
β-strand195-201765
β-strand207-214865
Chain F: 4 helices, 5 β-strands
ElementResiduesLengthSheet
β-strand73128
β-strand84166
α-helix85-862
α-helix87-10216
α-helix117-12610
β-strand132-137628
α-helix1381
β-strand143-147528
β-strand152166
Chain G: 4 helices, 15 β-strands
ElementResiduesLengthSheet
β-strand2-131228
α-helix15-173
α-helix22-3413
β-strand38-39228
β-strand43-49728
β-strand55128
α-helix56-583
β-strand60-61228
β-strand68-791228
β-strand86-951067
β-strand98-103667
β-strand106-111667
α-helix112-1143
β-strand120-122368
β-strand129-131368
β-strand136-137268
β-strand142-1521167
β-strand157-162667
β-strand169-170267
Chain H: 1 helix, 8 β-strands
ElementResiduesLengthSheet
β-strand4-161316
β-strand23-30816
β-strand37-43716
β-strand54-60716
α-helix89-924
β-strand95-1061216
β-strand111-118816
β-strand121-128816
β-strand139-145716

4 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 RPB1Aprotein1733Saccharomyces cerevisiaeP04050 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB2Bprotein1224Saccharomyces cerevisiaeP08518 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB3Cprotein318Saccharomyces cerevisiaeP16370 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB4Dprotein221Saccharomyces cerevisiaeP20433 (AlphaFold model)
DNA-directed RNA polymerases I, II, and III subunit RPABC1Eprotein215Saccharomyces cerevisiaeP20434
DNA-directed RNA polymerases I, II, and III subunit RPABC2Fprotein155Saccharomyces cerevisiaeP20435
DNA-directed RNA polymerase II subunit RPB7Gprotein171Saccharomyces cerevisiaeP34087
DNA-directed RNA polymerases I, II, and III subunit RPABC3Hprotein146Saccharomyces cerevisiaeP20436
DNA-directed RNA polymerase II subunit RPB9Iprotein122Saccharomyces cerevisiaeP27999
DNA-directed RNA polymerases I, II, and III subunit RPABC5Jprotein70Saccharomyces cerevisiaeP22139
DNA-directed RNA polymerase II subunit RPB11Kprotein120Saccharomyces cerevisiaeP38902
DNA-directed RNA polymerases I, II, and III subunit RPABC4Lprotein70Saccharomyces cerevisiaeP40422

3 more molecules are not listed.

Sequence of entity 1 (A), FASTA
>9QEB_1 DNA-directed RNA polymerase II subunit RPB1 (chains A)
MVGQQYSSAPLRTVKEVQFGLFSPEEVRAISVAKIRFPETMDETQTRAKIGGLNDPRLGS
IDRNLKCQTCQEGMNECPGHFGHIDLAKPVFHVGFIAKIKKVCECVCMHCGKLLLDEHNE
LMRQALAIKDSKKRFAAIWTLCKTKMVCETDVPSEDDPTQLVSRGGCGNTQPTIRKDGLK
LVGSWKKDRATGDADEPELRVLSTEEILNIFKHISVKDFTSLGFNEVFSRPEWMILTCLP
VPPPPVRPSISFNESQRGEDDLTFKLADILKANISLETLEHNGAPHHAIEEAESLLQFHV
ATYMDNDIAGQPQALQKSGRPVKSIRARLKGKEGRIRGNLMGKRVDFSARTVISGDPNLE
LDQVGVPKSIAKTLTYPEVVTPYNIDRLTQLVRNGPNEHPGAKYVIRDSGDRIDLRYSKR
AGDIQLQYGWKVERHIMDNDPVLFNRQPSLHKMSMMAHRVKVIPYSTFRLNLSVTSPYNA
DFDGDEMNLHVPQSEETRAELSQLCAVPLQIVSPQSNKPCMGIVQDTLCGIRKLTLRDTF
IELDQVLNMLYWVPDWDGVIPTPAIIKPKPLWSGKQILSVAIPNGIHLQRFDEGTTLLSP
KDNGMLIIDGQIIFGVVEKKTVGSSNGGLIHVVTREKGPQVCAKLFGNIQKVVNFWLLHN
GFSTGIGDTIADGPTMREITETIAEAKKKVLDVTKEAQANLLTAKHGMTLRESFEDNVVR
FLNEARDKAGRLAEVNLKDLNNVKQMVMAGSKGSFINIAQMSACVGQQSVEGKRIAFGFV
DRTLPHFSKDDYSPESKGFVENSYLRGLTPQEFFFHAMGGREGLIDTAVKTAETGYIQRR
LVKALEDIMVHYDNTTRNSLGNVIQFIYGEDGMDAAHIEKQSLDTIGGSDAAFEKRYRVD
LLNTDHTLDPSLLESGSEILGDLKLQVLLDEEYKQLVKDRKFLREVFVDGEANWPLPVNI
RRIIQNAQQTFHIDHTKPSDLTIKDIVLGVKDLQENLLVLRGKNEIIQNAQRDAVTLFCC
LLRSRLATRRVLQEYRLTKQAFDWVLSNIEAQFLRSVVHPGEMVGVLAAQSIGEPATQMT
LNTFHFAGVASKKVTSGVPRLKEILNVAKNMKTPSLTVYLEPGHAADQEQAKLIRSAIEH
TTLKSVTIASEIYYDPDPRSTVIPEDEEIIQLHFSLLDEEAEQSFDQQSPWLLRLELDRA
AMNDKDLTMGQVGERIKQTFKNDLFVIWSEDNDEKLIIRCRVVRPKSLDAETEAEEDHML
KKIENTMLENITLRGVENIERVVMMKYDRKVPSPTGEYVKEPEWVLETDGVNLSEVMTVP
GIDPTRIYTNSFIDIMEVLGIEAGRAALYKEVYNVIASDGSYVNYRHMALLVDVMTTQGG
LTSVTRHGFNRSNTGALMRCSFEETVEILFEAGASAELDDCRGVSENVILGQMAPIGTGA
FDVMIDEESLVKYMPEQKITEIEDGQDGGVTPYSNESGLVNADLDVKDELMFSPLVDSGS
NDAMAGGFTAYGGADYGEATSPFGAYGEAPTSPGFGVSSPGFSPTSPTYSPTSPAYSPTS
PSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYS
PTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPSYSPTSPAYSPTSPSYSPTSPSYSPTSP
SYSPTSPSYSPTSPNYSPTSPSYSPTSPGYSPGSPAYSPKQDEQKHNENENSR
Sequence of entity 2 (B), FASTA
>9QEB_2 DNA-directed RNA polymerase II subunit RPB2 (chains B)
MSDLANSEKYYDEDPYGFEDESAPITAEDSWAVISAFFREKGLVSQQLDSFNQFVDYTLQ
DIICEDSTLILEQLAQHTTESDNISRKYEISFGKIYVTKPMVNESDGVTHALYPQEARLR
NLTYSSGLFVDVKKRTYEAIDVPGRELKYELIAEESEDDSESGKVFIGRLPIMLRSKNCY
LSEATESDLYKLKECPFDMGGYFIINGSEKVLIAQERSAGNIVQVFKKAAPSPISHVAEI
RSALEKGSRFISTLQVKLYGREGSSARTIKATLPYIKQDIPIVIIFRALGIIPDGEILEH
ICYDVNDWQMLEMLKPCVEDGFVIQDRETALDFIGRRGTALGIKKEKRIQYAKDILQKEF
LPHITQLEGFESRKAFFLGYMINRLLLCALDRKDQDDRDHFGKKRLDLAGPLLAQLFKTL
FKKLTKDIFRYMQRTVEEAHDFNMKLAINAKTITSGLKYALATGNWGEQKKAMSSRAGVS
QVLNRYTYSSTLSHLRRTNTPIGRDGKLAKPRQLHNTHWGLVCPAETPEGQACGLVKNLS
LMSCISVGTDPMPIITFLSEWGMEPLEDYVPHQSPDATRVFVNGVWHGVHRNPARLMETL
RTLRRKGDINPEVSMIRDIREKELKIFTDAGRVYRPLFIVEDDESLGHKELKVRKGHIAK
LMATEYQDIEGGFEDVEEYTWSSLLNEGLVEYIDAEEEESILIAMQPEDLEPAEANEEND
LDVDPAKRIRVSHHATTFTHCEIHPSMILGVAASIIPFPDHNQSPRNTYQSAMGKQAMGV
FLTNYNVRMDTMANILYYPQKPLGTTRAMEYLKFRELPAGQNAIVAIACYSGYNQEDSMI
MNQSSIDRGLFRSLFFRSYMDQEKKYGMSITETFEKPQRTNTLRMKHGTYDKLDDDGLIA
PGVRVSGEDVIIGKTTPISPDEEELGQRTAYHSKRDASTPLRSTENGIVDQVLVTTNQDG
LKFVKVRVRTTKIPQIGDKFASRHGQKGTIGITYRREDMPFTAEGIVPDLIINPHAIPSR
MTVAHLIECLLSKVAALSGNEGDASPFTDITVEGISKLLREHGYQSRGFEVMYNGHTGKK
LMAQIFFGPTYYQRLRHMVDDKIHARARGPMQVLTRQPVEGRSRDGGLRFGEMERDCMIA
HGAASFLKERLMEASDAFRVHICGICGLMTVIAKLNHNQFECKGCDNKIDIYQIHIPYAA
KLLFQELMAMNITPRLYTDRSRDF
Sequence of entity 3 (C), FASTA
>9QEB_3 DNA-directed RNA polymerase II subunit RPB3 (chains C)
MSEEGPQVKIREASKDNVDFILSNVDLAMANSLRRVMIAEIPTLAIDSVEVETNTTVLAD
EFIAHRLGLIPLQSMDIEQLEYSRDCFCEDHCDKCSVVLTLQAFGESESTTNVYSKDLVI
VSNLMGRNIGHPIIQDKEGNGVLICKLRKGQELKLTCVAKKGIAKEHAKWGPAAAIEFEY
DPWNKLKHTDYWYEQDSAKEWPQSKNCEYEDPPNEGDPFDYKAQADTFYMNVESVGSIPV
DQVVVRGIDTLQKKVASILLALTQMDQDKVNFASGDNNTASNMLGSNEDVMMTGAEQDPY
SNASQMGNTGSGGYDNAW
Sequence of entity 4 (D), FASTA
>9QEB_4 DNA-directed RNA polymerase II subunit RPB4 (chains D)
MNVSTSTFQTRRRRLKKVEEEENAATLQLGQEFQLKQINHQGEEEELIALNLSEARLVIK
EALVERRRAFKRSQKKHKKKHLKHENANDETTAVEDEDDDLDEDDVNADDDDFMHSETRE
KELESIDVLLEQTTGGNNKDLKNTMQYLTNFSRFRDQETVGAVIQLLKSTGLHPFEVAQL
GSLACDTADEAKTLIPSLNNKISDDELERILKELSNLETLY
Sequence of entity 5 (E), FASTA
>9QEB_5 DNA-directed RNA polymerases I, II, and III subunit RPABC1 (chains E)
MDQENERNISRLWRAFRTVKEMVKDRGYFITQEEVELPLEDFKAKYCDSMGRPQRKMMSF
QANPTEESISKFPDMGSLWVEFCDEPSVGVKTMKTFVIHIQEKNFQTGIFVYQNNITPSA
MKLVPSIPPATIETFNEAALVVNITHHELVPKHIRLSSDEKRELLKRYRLKESQLPRIQR
ADPVALYLGLKRGEVVKIIRKSETSGRYASYRICM
Sequence of entity 6 (F), FASTA
>9QEB_6 DNA-directed RNA polymerases I, II, and III subunit RPABC2 (chains F)
MSDYEEAFNDGNENFEDFDVEHFSDEETYEEKPQFKDGETTDANGKTIVTGGNGPEDFQQ
HEQIRRKTLKEKAIPKDQRATTPYMTKYERARILGTRALQISMNAPVFVDLEGETDPLRI
AMKELAEKKIPLVIRRYLPDGSFEDWSVEELIVDL
Sequence of entity 7 (G), FASTA
>9QEB_7 DNA-directed RNA polymerase II subunit RPB7 (chains G)
MFFIKDLSLNITLHPSFFGPRMKQYLKTKLLEEVEGSCTGKFGYILCVLDYDNIDIQRGR
ILPTDGSAEFNVKYRAVVFKPFKGEVVDGTVVSCSQHGFEVQVGPMKVFVTKHLMPQDLT
FNAGSNPPSYQSSEDVITIKSRIRVKIEGCISQVSSIHAIGSIKEDYLGAI
Sequence of entity 8 (H), FASTA
>9QEB_8 DNA-directed RNA polymerases I, II, and III subunit RPABC3 (chains H)
MSNTLFDDIFQVSEVDPGRYNKVCRIEAASTTQDQCKLTLDINVELFPVAAQDSLTVTIA
SSLNLEDTPANDSSATRSWRPPQAGDRSLADDYDYVMYGTAYKFEEVSKDLIAVYYSFGG
LLMRLEGNYRNLNNLKQENAYLLIRR
Sequence of entity 9 (I), FASTA
>9QEB_9 DNA-directed RNA polymerase II subunit RPB9 (chains I)
MTTFRFCRDCNNMLYPREDKENNRLLFECRTCSYVEEAGSPLVYRHELITNIGETAGVVQ
DIGSDPTLPRSDRECPKCHSRENVFFQSQQRRKDTSMVLFFVCLSCSHIFTSDQKNKRTQ
FS
Sequence of entity 10 (J), FASTA
>9QEB_10 DNA-directed RNA polymerases I, II, and III subunit RPABC5 (chains J)
MIVPVRCFSCGKVVGDKWESYLNLLQEDELDEGTALSRLGLKRYCCRRMILTHVDLIEKF
LRYNPLEKRD
Sequence of entity 11 (K), FASTA
>9QEB_11 DNA-directed RNA polymerase II subunit RPB11 (chains K)
MNAPDRFELFLLGEGESKLKIDPDTKAPNAVVITFEKEDHTLGNLIRAELLNDRKVLFAA
YKVEHPFFARFKLRIQTTEGYDPKDALKNACNSIINKLGALKTNFETEWNLQTLAADDAF
Sequence of entity 12 (L), FASTA
>9QEB_12 DNA-directed RNA polymerases I, II, and III subunit RPABC4 (chains L)
MSREGFQIPTNLDAAAAGTSQARTATLKYICAECSSKLSLSRTDAVRCKDCGHRILLKAR
TKRLVQFEAR

Ligands and cofactors

IDNameFormulaCopies
ATPAdenosine-5'-triphosphateC10 H16 N5 O13 P31
ZNZinc ionZn8
MGMagnesium ionMg2

Water and common crystallization additives (K) are not listed.

Primary citation

Sub-2 angstrom cryo-EM structures of transcribing RNA polymerase II reveal critical roles of water molecules in catalysis. Li, Q., Yi, G., Wu, Y. et al. Mol Cell (2026) 86:1896-1910.e7. DOI 10.1016/j.molcel.2026.04.007 · PubMed

Other PDB entries of the same protein (UniProt P04050 (AlphaFold model), which also has an AlphaFold model), best resolution first:

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