9YMX: DNA-directed RNA polymerase subunit beta

De novo initial transcribing RNA polymerase with 5-mer RNA AUCUA (RPitc5b). Determined by electron microscopy at 2.7 Å resolution. Released 29 Apr 2026.

Method
Electron microscopy
Resolution
2.7 Å
Organisms
Escherichia coli, Escherichia phage T7
Chains
9
Atoms
31,075
Mol. weight
522.94 kDa
Ligands
POP, ZN, MG
Released
29 Apr 2026

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

Secondary structure: helices and β-sheets

9YMX contains 178 α-helices and 209 β-strands across 6 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 G: 7 helices, 16 β-strands

ElementResiduesLengthSheet
α-helix8-114
β-strand12-18733
β-strand23-31933
α-helix35-4713
β-strand52-611034
β-strand74135
α-helix78-869
β-strand90-92336
β-strand97-105934
β-strand108-111435
α-helix112-1143
β-strand115134
α-helix116-1172
β-strand121-123336
β-strand129-133535
β-strand139-1501234
β-strand152-153237
α-helix155-1573
β-strand170-172334
β-strand175-176237
β-strand180-1891033
β-strand199-207933
α-helix213-23220
Chain H: 5 helices, 14 β-strands
ElementResiduesLengthSheet
β-strand13-20838
β-strand23-31938
α-helix35-4915
β-strand52-60939
α-helix78-869
β-strand90-92340
β-strand97-105939
β-strand108-111441
α-helix112-1143
α-helix116-1172
β-strand121-123340
β-strand129-133541
β-strand139-1501239
β-strand152-153242
β-strand171-172239
β-strand175-176242
β-strand180-1921338
β-strand195-2071338
α-helix213-22715
Chain I: 63 helices, 93 β-strands
ElementResiduesLengthSheet
α-helix5-84
α-helix11-122
β-strand13-1421
α-helix19-202
α-helix29-3911
α-helix49-568
β-strand59-6022
β-strand66-75102
α-helix76-783
α-helix82-887
β-strand93-105132
β-strand114-128152
β-strand13013
β-strand136-13833
β-strand141-14553
α-helix1461
β-strand147-15154
α-helix1521
β-strand154-15965
α-helix169-1713
β-strand172-17765
β-strand18016
β-strand184-18855
β-strand194-19855
β-strand204-20525
α-helix206-2127
α-helix217-2248
β-strand227-23157
β-strand238-24037
α-helix243-2464
β-strand25018
β-strand255-25739
β-strand260-26349
β-strand26818
α-helix2691
α-helix271-28010
β-strand284-28637
α-helix289-2924
β-strand296110
β-strand29717
β-strand301-302211
β-strand309-311311
β-strand316110
α-helix319-3268
β-strand332-33657
α-helix346-3538
α-helix359-37012
α-helix378-38912
β-strand39616
α-helix398-40811
α-helix422-43716
β-strand451-45444
α-helix456-48126
α-helix489-4924
α-helix496-50813
β-strand512-51433
α-helix515-5162
α-helix520-5278
β-strand529-53134
α-helix552-5543
β-strand558112
β-strand572-57434
β-strand575112
β-strand580-581213
β-strand587-595913
β-strand598-6071013
α-helix608-6114
β-strand616-617213
β-strand623114
β-strand629114
β-strand633-637513
β-strand642-646513
α-helix647-6493
β-strand652-654313
α-helix665-6673
α-helix671-6733
β-strand674115
α-helix676-68510
α-helix686-6883
β-strand689116
α-helix690-6912
β-strand700-70121
α-helix705-7128
β-strand716-717217
β-strand722-727618
β-strand731-736618
β-strand748-752518
α-helix753-7542
β-strand757-758219
β-strand763120
β-strand764-765219
β-strand768-769217
β-strand776-777218
β-strand782-785417
β-strand789-790221
β-strand793-794221
β-strand798-804722
β-strand815-819522
α-helix820-8245
β-strand830-8411220
β-strand842123
β-strand847123
β-strand849124
β-strand865125
β-strand870126
β-strand871125
α-helix8721
β-strand876-878320
β-strand882-884326
β-strand887-889324
α-helix897-90610
β-strand913-915324
β-strand918-919226
α-helix920-9212
β-strand925-9351120
α-helix943-96018
α-helix964-97815
α-helix994-9996
α-helix1006-103631
α-helix1042-10432
β-strand1046-10581320
β-strand1065-1067322
β-strand1073-1080822
α-helix10811
α-helix1082-10843
β-strand1087127
α-helix10921
β-strand1093127
α-helix10941
β-strand1096-1098322
α-helix1100-11023
β-strand1108115
α-helix1110-113324
α-helix1138-115013
α-helix1161-11633
α-helix1168-11758
β-strand1182-118321
α-helix1192-120110
β-strand1209-1210222
β-strand1213127
α-helix12191
β-strand1220127
α-helix12211
β-strand1225-12341022
β-strand1235116
β-strand1244-1246328
β-strand1251129
β-strand1252130
β-strand1258129
α-helix1267-12682
β-strand1269-1270231
α-helix1272-12809
α-helix1284-12918
α-helix1298-130912
α-helix1321-133111
β-strand1335-1340632
Chain J: 70 helices, 82 β-strands
ElementResiduesLengthSheet
β-strand19-24632
α-helix27-337
β-strand34-37443
β-strand44144
α-helix501
β-strand51144
α-helix521
α-helix59-624
α-helix78-803
β-strand84145
β-strand91145
α-helix95-995
β-strand102-1121143
α-helix115-1184
α-helix1221
α-helix123-1286
α-helix132-1398
β-strand144-148546
α-helix154-1552
β-strand159-160246
α-helix162-17211
β-strand177-179346
α-helix181-19010
α-helix194-20714
α-helix211-22919
α-helix234-2374
β-strand238-244743
α-helix247-2493
β-strand252-255447
β-strand259-262447
α-helix264-28522
α-helix289-30719
α-helix315-3162
β-strand317148
β-strand323148
α-helix324-3252
α-helix328-3303
α-helix3371
α-helix338-3425
β-strand345-346231
β-strand350-354528
β-strand355-357349
β-strand365-369549
α-helix370-3767
α-helix378-38811
α-helix394-40310
α-helix406-41510
β-strand421-424428
α-helix431-4333
β-strand434-437428
β-strand438-442549
β-strand447-449349
α-helix451-4533
α-helix454-4574
β-strand465-469528
α-helix474-4829
β-strand485128
α-helix486-4883
β-strand491150
β-strand498150
α-helix505-5139
β-strand526-527251
α-helix530-5389
β-strand547-5571151
β-strand563-5731151
α-helix574-5796
α-helix580-5823
α-helix5841
α-helix589-5913
α-helix598-61114
α-helix614-63522
α-helix645-6473
α-helix650-66920
α-helix675-70127
β-strand703-707552
β-strand713-717552
α-helix721-7277
β-strand745153
β-strand748154
α-helix7531
β-strand754154
α-helix7551
β-strand759153
α-helix769-80436
β-strand809-811355
β-strand820-822356
β-strand825-827357
β-strand828158
β-strand830-833457
α-helix836-8394
β-strand843159
β-strand844156
β-strand848-850360
β-strand855-858460
β-strand863159
α-helix866-8749
β-strand880-882356
α-helix885-8873
β-strand894-895255
β-strand901161
β-strand908161
α-helix909-9102
α-helix915-92410
α-helix925-9295
β-strand949-951362
β-strand956-961658
β-strand965-967362
β-strand973-975362
β-strand980-985658
β-strand991-997758
β-strand1002-1003262
β-strand1010-1011258
β-strand1016-1019462
β-strand1026-1028363
β-strand1033-1038664
β-strand1042163
β-strand1046-1050563
β-strand1057-1061563
α-helix1069-10724
β-strand1077-1081564
β-strand1087164
β-strand1089165
α-helix10901
β-strand1096165
β-strand1098-1100364
β-strand1106-1107263
β-strand1114-1115264
β-strand1120-1123463
α-helix1138-11469
α-helix1148-11503
α-helix1153-11553
β-strand1156166
β-strand1161-1165567
β-strand1172-1178767
β-strand1186-1191667
β-strand1203-1204267
β-strand1209166
α-helix1214-12163
α-helix1217-12248
α-helix1226-124217
α-helix1250-126011
β-strand1263-1268668
β-strand1279-1281368
α-helix1282-129413
β-strand1301-1305568
α-helix1309-13146
α-helix1319-13257
α-helix1328-133811
β-strand1341-1342232
α-helix1347-13526
α-helix1360-13623
α-helix1363-137412
Chain K: 4 helices, 0 β-strands
ElementResiduesLengthSheet
α-helix7-137
α-helix16-3217
α-helix46-5510
α-helix61-7919
Chain L: 29 helices, 4 β-strands
ElementResiduesLengthSheet
α-helix97-1059
α-helix109-1113
α-helix113-13523
α-helix138-15215
α-helix215-23521
α-helix245-25814
α-helix263-29533
α-helix299-3068
α-helix314-3185
α-helix325-3295
α-helix330-3323
α-helix334-35118
α-helix355-38026
α-helix384-3918
α-helix392-3943
α-helix401-41818
α-helix421-4233
α-helix427-44620
α-helix454-47421
α-helix477-4793
α-helix480-4878
α-helix491-4999
α-helix503-5042
β-strand505-506247
β-strand510169
β-strand518169
α-helix519-5224
β-strand524130
α-helix531-54919
α-helix555-5639
α-helix575-5806
α-helix584-59916
α-helix607-6104

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
DNA-directed RNA polymerase subunit betaIprotein1342Escherichia coliP0A8V2 (AlphaFold model)
DNA-directed RNA polymerase subunit omegaKprotein91Escherichia coliP0A800 (AlphaFold model)
Nascent RNARRNA5Escherichia coli
T7A1 promoter fragment non-template strandPDNA97Escherichia phage T7
DNA-directed RNA polymerase subunit alphaG, Hprotein329Escherichia coliP0A7Z4 (AlphaFold model)
DNA-directed RNA polymerase subunit beta'Jprotein1415Escherichia coliP0A8T7 (AlphaFold model)
RNA polymerase sigma factor RpoDLprotein616Escherichia coliP00579
T7A1 promoter fragment template strandQDNA97Escherichia phage T7
Sequence of entity 1 (I), FASTA
>9YMX_1 DNA-directed RNA polymerase subunit beta (chains I)
MVYSYTEKKRIRKDFGKRPQVLDVPYLLSIQLDSFQKFIEQDPEGQYGLEAAFRSVFPIQ
SYSGNSELQYVSYRLGEPVFDVQECQIRGVTYSAPLRVKLRLVIYEREAPEGTVKDIKEQ
EVYMGEIPLMTDNGTFVINGTERVIVSQLHRSPGVFFDSDKGKTHSSGKVLYNARIIPYR
GSWLDFEFDPKDNLFVRIDRRRKLPATIILRALNYTTEQILDLFFEKVIFEIRDNKLQME
LVPERLRGETASFDIEANGKVYVEKGRRITARHIRQLEKDDVKLIEVPVEYIAGKVVAKD
YIDESTGELICAANMELSLDLLAKLSQSGHKRIETLFTNDLDHGPYISETLRVDPTNDRL
SALVEIYRMMRPGEPPTREAAESLFENLFFSEDRYDLSAVGRMKFNRSLLREEIEGSGIL
SKDDIIDVMKKLIDIRNGKGEVDDIDHLGNRRIRSVGEMAENQFRVGLVRVERAVKERLS
LGDLDTLMPQDMINAKPISAAVKEFFGSSQLSQFMDQNNPLSEITHKRRISALGPGGLTR
ERAGFEVRDVHPTHYGRVCPIETPEGPNIGLINSLSVYAQTNEYGFLETPYRKVTDGVVT
DEIHYLSAIEEGNYVIAQANSNLDEEGHFVEDLVTCRSKGESSLFSRDQVDYMDVSTQQV
VSVGASLIPFLEHDDANRALMGANMQRQAVPTLRADKPLVGTGMERAVAVDSGVTAVAKR
GGVVQYVDASRIVIKVNEDEMYPGEAGIDIYNLTKYTRSNQNTCINQMPCVSLGEPVERG
DVLADGPSTDLGELALGQNMRVAFMPWNGYNFEDSILVSERVVQEDRFTTIHIQELACVS
RDTKLGPEEITADIPNVGEAALSKLDESGIVYIGAEVTGGDILVGKVTPKGETQLTPEEK
LLRAIFGEKASDVKDSSLRVPNGVSGTVIDVQVFTRDGVEKDKRALEIEEMQLKQAKKDL
SEELQILEAGLFSRIRAVLVAGGVEAEKLDKLPRDRWLELGLTDEEKQNQLEQLAEQYDE
LKHEFEKKLEAKRRKITQGDDLAPGVLKIVKVYLAVKRRIQPGDKMAGRHGNKGVISKIN
PIEDMPYDENGTPVDIVLNPLGVPSRMNIGQILETHLGMAAKGIGDKINAMLKQQQEVAK
LREFIQRAYDLGADVRQKVDLSTFSDEEVMRLAENLRKGMPIATPVFDGAKEAEIKELLK
LGDLPTSGQIRLYDGRTGEQFERPVTVGYMYMLKLNHLVDDKMHARSTGSYSLVTQQPLG
GKAQFGGQRFGEMEVWALEAYGAAYTLQEMLTVKSDDVNGRTKMYKNIVDGNHQMEPGMP
ESFNVLLKEIRSLGINIELEDE
Sequence of entity 2 (K), FASTA
>9YMX_2 DNA-directed RNA polymerase subunit omega (chains K)
MARVTVQDAVEKIGNRFDLVLVAARRARQMQVGGKDPLVPEENDKTTVIALREIEEGLIN
NQILDVRERQEQQEQEAAELQAVTAIAEGRR
Sequence of entity 3 (R), FASTA
>9YMX_3 Nascent RNA (chains R)
AUCUA
Sequence of entity 4 (P), FASTA
>9YMX_4 T7A1 promoter fragment non-template strand (chains P)
GCTTAATTTAAAATTTATCAAAAAGAGTATTGACTTAAAGTCTAACCTATAGGATACTTA
CAGCCATCGAGAGGGACACGGCGAATAGCCATCCCGG
Sequence of entity 5 (G, H), FASTA
>9YMX_5 DNA-directed RNA polymerase subunit alpha (chains G, H)
MQGSVTEFLKPRLVDIEQVSSTHAKVTLEPLERGFGHTLGNALRRILLSSMPGCAVTEVE
IDGVLHEYSTKEGVQEDILEILLNLKGLAVRVQGKDEVILTLNKSGIGPVTAADITHDGD
VEIVKPQHVICHLTDENASISMRIKVQRGRGYVPASTRIHSEEDERPIGRLLVDACYSPV
ERIAYNVEAARVEQRTDLDKLVIEMETNGTIDPEEAIRRAATILAEQLEAFVDLRDVRQP
EVKEEKPEFDPILLRPVDDLELTVRSANCLKAEAIHYIGDLVQRTEVELLKTPNLGKKSL
TEIKDVLASRGLSLGMRLENWPPASIADE
Sequence of entity 6 (J), FASTA
>9YMX_6 DNA-directed RNA polymerase subunit beta' (chains J)
MKDLLKFLKAQTKTEEFDAIKIALASPDMIRSWSFGEVKKPETINYRTFKPERDGLFCAR
IFGPVKDYECLCGKYKRLKHRGVICEKCGVEVTQTKVRRERMGHIELASPTAHIWFLKSL
PSRIGLLLDMPLRDIERVLYFESYVVIEGGMTNLERQQILTEEQYLDALEEFGDEFDAKM
GAEAIQALLKSMDLEQECEQLREELNETNSETKRKKLTKRIKLLEAFVQSGNKPEWMILT
VLPVLPPDLRPLVPLDGGRFATSDLNDLYRRVINRNNRLKRLLDLAAPDIIVRNEKRMLQ
EAVDALLDNGRRGRAITGSNKRPLKSLADMIKGKQGRFRQNLLGKRVDYSGRSVITVGPY
LRLHQCGLPKKMALELFKPFIYGKLELRGLATTIKAAKKMVEREEAVVWDILDEVIREHP
VLLNRAPTLHRLGIQAFEPVLIEGKAIQLHPLVCAAYNADFDGDQMAVHVPLTLEAQLEA
RALMMSTNNILSPANGEPIIVPSQDVVLGLYYMTRDCVNAKGEGMVLTGPKEAERLYRSG
LASLHARVKVRITEYEKDANGELVAKTSLKDTTVGRAILWMIVPKGLPYSIVNQALGKKA
ISKMLNTCYRILGLKPTVIFADQIMYTGFAYAARSGASVGIDDMVIPEKKHEIISEAEAE
VAEIQEQFQSGLVTAGERYNKVIDIWAAANDRVSKAMMDNLQTETVINRDGQEEKQVSFN
SIYMMADSGARGSAAQIRQLAGMRGLMAKPDGSIIETPITANFREGLNVLQYFISTHGAR
KGLADTALKTANSGYLTRRLVDVAQDLVVTEDDCGTHEGIMMTPVIEGGDVKEPLRDRVL
GRVTAEDVLKPGTADILVPRNTLLHEQWCDLLEENSVDAVKVRSVVSCDTDFGVCAHCYG
RDLARGHIINKGEAIGVIAAQSIGEPGTQLTMRTFHIGGAASRAAAESSIQVKNKGSIKL
SNVKSVVNSSGKLVITSRNTELKLIDEFGRTKESYKVPYGAVLAKGDGEQVAGGETVANW
DPHTMPVITEVSGFVRFTDMIDGQTITRQTDELTGLSSLVVLDSAERTAGGKDLRPALKI
VDAQGNDVLIPGTDMPAQYFLPGKAIVQLEDGVQISSGDTLARIPQESGGTKDITGGLPR
VADLFEARRPKEPAILAEISGIVSFGKETKGKRRLVITPVDGSDPYEEMIPKWRQLNVFE
GERVERGDVISDGPEAPHDILRLRGVHAVTRYIVNEVQDVYRLQGVKINDKHIEVIVRQM
LRKATIVNAGSSDFLEGEQVEYSRVKIANRELEANGKVGATYSRDLLGITKASLATESFI
SAASFQETTRVLTEAAVAGKRDELRGLKENVIVGRLIPAGTGYAYHQDRMRRRAAGEAPA
APQVTAEDASASLAELLNAGLGGSDNELELEVLFQ
Sequence of entity 7 (L), FASTA
>9YMX_7 RNA polymerase sigma factor RpoD (chains L)
SEFMEQNPQSQLKLLVTRGKEQGYLTYAEVNDHLPEDIVDSDQIEDIIQMINDMGIQVME
EAPDADDLMLAENTADEDAAEAAAQVLSSVESEIGRTTDPVRMYMREMGTVELLTREGEI
DIAKRIEDGINQVQCSVAEYPEAITYLLEQYDRVEAEEARLSDLITGFVDPNAEEDLAPT
ATHVGSELSQEDLDDDEDEDEEDGDDDSADDDNSIDPELAREKFAELRAQYVVTRDTIKA
KGRSHATAQEEILKLSEVFKQFRLVPKQFDYLVNSMRVMMDRVRTQERLIMKLCVEQCKM
PKKNFITLFTGNETSDTWFNAAIAMNKPWSEKLHDVSEEVHRALQKLQQIEEETGLTIEQ
VKDINRRMSIGEAKARRAKKEMVEANLRLVISIAKKYTNRGLQFLDLIQEGNIGLMKAVD
KFEYRRGYKFSTYATWWIRQAITRSIADQARTIRIPVHMIETINKLNRISRQMLQEMGRE
PTPEELAERMLMPEDKIRKVLKIAKEPISMETPIGDDEDSHLGDFIEDTTLELPLDSATT
ESLRAATHDVLAGLTAREAKVLRMRFGIDMNTDYTLEEVGKQFDVTRERIRQIEAKALRK
LRHPSRSEVLRSFLDD
Sequence of entity 8 (Q), FASTA
>9YMX_8 T7A1 promoter fragment template strand (chains Q)
CCGGGATGGCTATTCGCCGTGTCCCTCTCGATGGCTGTAAGTATCCTATAGGTTAGACTT
TAAGTCAATACTCTTTTTGATAAATTTTAAATTAAGC

Ligands and cofactors

IDNameFormulaCopies
POPPyrophosphate 2-H2 O7 P21
ZNZinc ionZn2
MGMagnesium ionMg1

Primary citation

Structural basis for multi-subunit DNA-dependent RNA polymerase catalytic activity. Mueller, A.U., Darst, S.A. Mol Cell (2026) 86:1881. DOI 10.1016/j.molcel.2026.03.033 · PubMed

Other PDB entries of the same protein (UniProt P0A8V2 (AlphaFold model), which also has an AlphaFold model), best resolution first:

Browse structure collections

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