9KD9: DNA-directed RNA polymerase II subunit RPB1

The structure of RNA polymerase II elongation complex paused at N-5 state by actinomycin D. Determined by electron microscopy at 2.88 Å resolution. Released 5 Nov 2025.

Method
Electron microscopy
Resolution
2.88 Å
Organisms
Saccharomyces cerevisiae S288C, Streptomyces sp., synthetic construct
Chains
16
Atoms
32,956
Mol. weight
549.89 kDa
Ligands
MG, ZN
Released
5 Nov 2025

Explore 9KD9 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9KD9 contains 190 α-helices and 233 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: 79 helices, 65 β-strands

ElementResiduesLengthSheet
α-helix101
β-strand1111
β-strand16-1832
α-helix24-307
β-strand3413
β-strand3714
β-strand4115
β-strand4815
α-helix491
β-strand5214
α-helix56-583
β-strand82-91103
α-helix96-1038
β-strand10616
β-strand11316
α-helix122-1276
α-helix131-14212
β-strand151-15337
β-strand161-16337
α-helix171-1722
β-strand173-17758
β-strand180-18568
α-helix186-1872
β-strand198-20148
α-helix204-2129
α-helix216-2216
α-helix231-2344
β-strand235-24173
α-helix261-28222
α-helix286-30419
α-helix312-3132
β-strand31519
β-strand32119
α-helix325-3295
α-helix3351
α-helix336-3405
β-strand343-345310
β-strand348-355811
β-strand363-367511
α-helix368-3736
β-strand375-379512
α-helix385-39410
β-strand402-406512
β-strand412-414312
α-helix419-4213
β-strand431-435512
β-strand441-445511
α-helix452-4543
β-strand455-463911
β-strand468-470311
α-helix475-4784
β-strand486-490511
α-helix495-50410
β-strand506111
α-helix508-5103
β-strand512-513213
β-strand518-519213
α-helix526-5349
β-strand540-541214
α-helix543-5519
α-helix560-5634
β-strand565-567315
β-strand571115
β-strand572-573214
α-helix574-5818
α-helix5831
β-strand588-590316
β-strand605-608416
β-strand611-614416
α-helix619-6224
α-helix629-6368
α-helix639-65820
β-strand663117
α-helix667-6693
α-helix673-69826
α-helix702-7054
α-helix710-73627
α-helix742-7498
α-helix755-7584
α-helix759-7635
β-strand766-767218
β-strand769-770219
β-strand773-774219
α-helix775-7762
α-helix794-7974
β-strand799-800218
α-helix810-84536
β-strand849-850220
β-strand856-857220
β-strand863-865320
α-helix868-8703
β-strand873121
α-helix875-8773
β-strand879-882422
α-helix890-8978
α-helix910-9123
β-strand913123
α-helix924-94623
β-strand953-956422
α-helix960-97112
α-helix9781
β-strand979123
α-helix983-99614
α-helix1005-102521
α-helix1028-10292
α-helix1030-10356
α-helix1039-105517
β-strand1057121
α-helix1058-10592
α-helix1064-107714
α-helix1097-11059
β-strand1115-1119524
β-strand1120125
α-helix11211
α-helix1122-11243
α-helix1128-113811
α-helix1143-11453
β-strand1147-1154826
α-helix1167-117610
α-helix1185-11873
β-strand1188-11971026
α-helix1199-12046
α-helix1209-122012
α-helix1221-12233
β-strand1224-1228526
β-strand1237-1243726
α-helix1244-12452
α-helix1252-126918
β-strand1281-12921224
β-strand1298-13091224
α-helix1313-13175
β-strand1322125
β-strand1328-1329224
α-helix1332-13387
α-helix1341-135818
α-helix1365-137511
α-helix1389-13913
α-helix1396-13994
α-helix1405-141511
β-strand1418-141922
α-helix1424-14296
α-helix1437-14393
β-strand1441-1445527
α-helix1447-14537
Chain B: 53 helices, 84 β-strands
ElementResiduesLengthSheet
α-helix231
β-strand24128
α-helix251
α-helix29-4012
α-helix45-539
α-helix54-585
α-helix59-646
β-strand68-73629
α-helix80-823
β-strand84-971429
β-strand101-102230
β-strand110-111230
α-helix114-1207
β-strand125-1401629
α-helix146-1472
β-strand150-151229
β-strand156129
β-strand164-171829
α-helix186-1916
β-strand203-205331
β-strand208-212531
β-strand214-218532
α-helix2191
β-strand223-227533
β-strand234-242933
β-strand252-259833
β-strand269-272433
β-strand276134
β-strand280-281233
α-helix282-2898
α-helix294-3018
α-helix308-32114
α-helix327-33711
β-strand338134
α-helix345-35410
α-helix355-3595
α-helix371-38919
α-helix394-3974
α-helix401-4033
β-strand404-407432
α-helix409-43729
α-helix450-46213
β-strand480-482331
α-helix483-4842
α-helix488-4958
β-strand497-499332
α-helix516-5183
β-strand522135
β-strand533132
β-strand536-538332
β-strand539135
β-strand540136
β-strand544-545237
α-helix552-5609
α-helix5631
β-strand564-565238
β-strand578-582538
β-strand585-590638
α-helix593-60513
β-strand614-618538
β-strand623-627538
β-strand633-639737
β-strand640-641239
β-strand650-651239
β-strand654128
α-helix655-66814
α-helix681-6866
β-strand690-694537
α-helix697-7004
β-strand703-704237
α-helix707-7104
α-helix728-7292
β-strand740-741237
α-helix745-7484
β-strand750136
α-helix759-7613
α-helix764-77411
β-strand777140
α-helix785-7873
β-strand792-796541
β-strand800142
β-strand804-805243
α-helix807-8093
β-strand821-827744
β-strand828117
β-strand839-842444
α-helix843-8475
β-strand853-8621041
β-strand865145
β-strand871145
β-strand873-874246
β-strand883146
β-strand898147
α-helix899-9002
β-strand904-905241
β-strand910-912347
β-strand914-917446
β-strand924148
β-strand928148
β-strand934-936346
β-strand939-940247
α-helix941-9422
β-strand947-9561041
β-strand962-9721141
β-strand979-982444
β-strand987-994844
β-strand1001149
β-strand1010-1012344
α-helix1016-10205
α-helix1023-103816
β-strand1042-1043243
α-helix1052-106110
β-strand1069-1070244
β-strand1073149
α-helix10791
β-strand1080149
α-helix10811
α-helix10841
β-strand1085-10941044
β-strand1095140
α-helix1099-11013
β-strand1104-1106311
β-strand1111150
β-strand1118150
α-helix1122-11243
β-strand1128-1130310
α-helix1132-11409
α-helix1144-11485
α-helix1149-11535
β-strand1158-116361
β-strand1172-1174351
β-strand1179-1181351
β-strand1182152
β-strand1187152
β-strand1191-119661
α-helix1198-120811
β-strand1215-121732
Chain C: 11 helices, 12 β-strands
ElementResiduesLengthSheet
β-strand7-13753
β-strand17-23753
α-helix27-3913
β-strand43-541254
α-helix60-689
β-strand72-74355
α-helix77-793
β-strand97-104854
β-strand111-114456
β-strand119-120254
β-strand129-132455
α-helix1331
β-strand143-147556
β-strand152-1621154
α-helix168-1703
β-strand176-180553
α-helix197-2004
α-helix202-2043
α-helix205-2095
α-helix211-2133
α-helix218-2192
β-strand228-233653
α-helix240-26627
Chain D: 9 helices, 5 β-strands
ElementResiduesLengthSheet
α-helix20-223
β-strand23157
β-strand28157
β-strand36-38358
β-strand44-46358
β-strand49159
α-helix52-7221
α-helix119-13315
α-helix139-15012
α-helix157-16913
α-helix174-1829
α-helix188-1947
α-helix196-1983
α-helix204-21613
Chain E: 12 helices, 14 β-strands
ElementResiduesLengthSheet
α-helix3-2624
β-strand28160
α-helix32-354
α-helix39-468
β-strand47161
β-strand53161
β-strand60-62362
β-strand64160
α-helix66-716
β-strand78-82562
β-strand87-88263
α-helix90-10213
β-strand107-112662
β-strand115-116263
α-helix118-1214
β-strand131-136662
α-helix144-1463
β-strand152-155464
α-helix156-1572
α-helix158-16710
α-helix1761
β-strand177-178264
α-helix183-1886
α-helix1941
β-strand195-200664
β-strand208-214764
Chain F: 5 helices, 5 β-strands
ElementResiduesLengthSheet
β-strand73127
α-helix76-783
β-strand84165
α-helix85-862
α-helix87-10216
α-helix117-12610
β-strand132-137627
α-helix1381
β-strand143-147527
β-strand152165
Chain G: 4 helices, 15 β-strands
ElementResiduesLengthSheet
β-strand2-131259
α-helix15-173
α-helix22-309
β-strand37-39359
β-strand43-49759
β-strand55159
β-strand61159
α-helix621
β-strand68-791259
β-strand85-941066
β-strand98-103666
β-strand106-111666
α-helix112-1143
β-strand120-122367
β-strand129-131367
β-strand136-137267
β-strand142-1511066
β-strand158-162566
β-strand170166
Chain H: 5 helices, 10 β-strands
ElementResiduesLengthSheet
β-strand4-161315
β-strand23-30815
β-strand37-43715
β-strand54-60715
β-strand64168
α-helix67-682
α-helix78-792
α-helix81-833
β-strand88168
α-helix89-924
β-strand95-1051115
β-strand111-118815
β-strand121-128815
α-helix129-1324
β-strand139-145715

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 RPB1Aprotein1733Saccharomyces cerevisiae S288CP04050 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB2Bprotein1224Saccharomyces cerevisiae S288CP08518 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB3Cprotein318Saccharomyces cerevisiae S288CP16370 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB4Dprotein221Saccharomyces cerevisiae S288CP20433 (AlphaFold model)
DNA-directed RNA polymerases I, II, and III subunit RPABC1Eprotein215Saccharomyces cerevisiae S288CP20434
DNA-directed RNA polymerases I, II, and III subunit RPABC2Fprotein155Saccharomyces cerevisiae S288CP20435
DNA-directed RNA polymerase II subunit RPB7Gprotein171Saccharomyces cerevisiae S288CP34087
DNA-directed RNA polymerases I, II, and III subunit RPABC3Hprotein146Saccharomyces cerevisiae S288CP20436
DNA-directed RNA polymerase II subunit RPB9Iprotein122Saccharomyces cerevisiae S288CP27999
DNA-directed RNA polymerases I, II, and III subunit RPABC5Jprotein70Saccharomyces cerevisiae S288CP22139
DNA-directed RNA polymerase II subunit RPB11Kprotein120Saccharomyces cerevisiae S288CP38902
DNA-directed RNA polymerases I, II, and III subunit RPABC4Lprotein70Saccharomyces cerevisiae S288CP40422

4 more molecules are not listed.

Sequence of entity 1 (A), FASTA
>9KD9_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
>9KD9_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
>9KD9_3 DNA-directed RNA polymerase II subunit RPB3 (chains C)
MSEEGPQVKIREASKDNVDFILSNVDLAMANSLRRVMIAEIPTLAIDSVEVETNTTVLAD
EFIAHRLGLIPLQSMDIEQLEYSRDCFCEDHCDKCSVVLTLQAFGESESTTNVYSKDLVI
VSNLMGRNIGHPIIQDKEGNGVLICKLRKGQELKLTCVAKKGIAKEHAKWGPAAAIEFEY
DPWNKLKHTDYWYEQDSAKEWPQSKNCEYEDPPNEGDPFDYKAQADTFYMNVESVGSIPV
DQVVVRGIDTLQKKVASILLALTQMDQDKVNFASGDNNTASNMLGSNEDVMMTGAEQDPY
SNASQMGNTGSGGYDNAW
Sequence of entity 4 (D), FASTA
>9KD9_4 DNA-directed RNA polymerase II subunit RPB4 (chains D)
MNVSTSTFQTRRRRLKKVEEEENAATLQLGQEFQLKQINHQGEEEELIALNLSEARLVIK
EALVERRRAFKRSQKKHKKKHLKHENANDETTAVEDEDDDLDEDDVNADDDDFMHSETRE
KELESIDVLLEQTTGGNNKDLKNTMQYLTNFSRFRDQETVGAVIQLLKSTGLHPFEVAQL
GSLACDTADEAKTLIPSLNNKISDDELERILKELSNLETLY
Sequence of entity 5 (E), FASTA
>9KD9_5 DNA-directed RNA polymerases I, II, and III subunit RPABC1 (chains E)
MDQENERNISRLWRAFRTVKEMVKDRGYFITQEEVELPLEDFKAKYCDSMGRPQRKMMSF
QANPTEESISKFPDMGSLWVEFCDEPSVGVKTMKTFVIHIQEKNFQTGIFVYQNNITPSA
MKLVPSIPPATIETFNEAALVVNITHHELVPKHIRLSSDEKRELLKRYRLKESQLPRIQR
ADPVALYLGLKRGEVVKIIRKSETSGRYASYRICM
Sequence of entity 6 (F), FASTA
>9KD9_6 DNA-directed RNA polymerases I, II, and III subunit RPABC2 (chains F)
MSDYEEAFNDGNENFEDFDVEHFSDEETYEEKPQFKDGETTDANGKTIVTGGNGPEDFQQ
HEQIRRKTLKEKAIPKDQRATTPYMTKYERARILGTRALQISMNAPVFVDLEGETDPLRI
AMKELAEKKIPLVIRRYLPDGSFEDWSVEELIVDL
Sequence of entity 7 (G), FASTA
>9KD9_7 DNA-directed RNA polymerase II subunit RPB7 (chains G)
MFFIKDLSLNITLHPSFFGPRMKQYLKTKLLEEVEGSCTGKFGYILCVLDYDNIDIQRGR
ILPTDGSAEFNVKYRAVVFKPFKGEVVDGTVVSCSQHGFEVQVGPMKVFVTKHLMPQDLT
FNAGSNPPSYQSSEDVITIKSRIRVKIEGCISQVSSIHAIGSIKEDYLGAI
Sequence of entity 8 (H), FASTA
>9KD9_8 DNA-directed RNA polymerases I, II, and III subunit RPABC3 (chains H)
MSNTLFDDIFQVSEVDPGRYNKVCRIEAASTTQDQCKLTLDINVELFPVAAQDSLTVTIA
SSLNLEDTPANDSSATRSWRPPQAGDRSLADDYDYVMYGTAYKFEEVSKDLIAVYYSFGG
LLMRLEGNYRNLNNLKQENAYLLIRR
Sequence of entity 9 (I), FASTA
>9KD9_9 DNA-directed RNA polymerase II subunit RPB9 (chains I)
MTTFRFCRDCNNMLYPREDKENNRLLFECRTCSYVEEAGSPLVYRHELITNIGETAGVVQ
DIGSDPTLPRSDRECPKCHSRENVFFQSQQRRKDTSMVLFFVCLSCSHIFTSDQKNKRTQ
FS
Sequence of entity 10 (J), FASTA
>9KD9_10 DNA-directed RNA polymerases I, II, and III subunit RPABC5 (chains J)
MIVPVRCFSCGKVVGDKWESYLNLLQEDELDEGTALSRLGLKRYCCRRMILTHVDLIEKF
LRYNPLEKRD
Sequence of entity 11 (K), FASTA
>9KD9_11 DNA-directed RNA polymerase II subunit RPB11 (chains K)
MNAPDRFELFLLGEGESKLKIDPDTKAPNAVVITFEKEDHTLGNLIRAELLNDRKVLFAA
YKVEHPFFARFKLRIQTTEGYDPKDALKNACNSIINKLGALKTNFETEWNLQTLAADDAF
Sequence of entity 12 (L), FASTA
>9KD9_12 DNA-directed RNA polymerases I, II, and III subunit RPABC4 (chains L)
MSREGFQIPTNLDAAAAGTSQARTATLKYICAECSSKLSLSRTDAVRCKDCGHRILLKAR
TKRLVQFEAR

Ligands and cofactors

IDNameFormulaCopies
MGMagnesium ionMg1
ZNZinc ionZn8

Primary citation

Stepwise transcription stalling by the anti-cancer drug Actinomycin D and insights into short tandem repeat transcription inhibition. Zhao, W., Zhu, L., Liu, Y. et al. Nat Commun (2026) 17. DOI 10.1038/s41467-026-70612-y · 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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