8RIF: MCM double hexamer loaded onto dsDNA

Cryo-EM structure of the MCM double hexamer loaded onto dsDNA. Determined by electron microscopy at 2.79 Å resolution. Released 1 Jan 2025.

Method
Electron microscopy
Resolution
2.79 Å
Organism
Saccharomyces cerevisiae
Chains
14
Atoms
64,125
Mol. weight
1257.96 kDa
Ligands
ATP, MG, ZN, ADP
Released
1 Jan 2025

Explore 8RIF in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

8RIF contains 366 α-helices and 356 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.

Chains 2 and A: 27 helices, 25 β-strands

ElementResiduesLengthSheet
α-helix183-1864
α-helix194-1996
α-helix201-21616
α-helix226-23712
β-strand241-24441
α-helix246-2527
α-helix254-2629
α-helix264-28219
α-helix286-2894
β-strand294-29741
β-strand30412
α-helix311-3133
β-strand317-327112
β-strand331-341112
β-strand346-35272
β-strand373-37532
β-strand381-391112
α-helix394-3963
α-helix3981
β-strand405-41172
α-helix412-4143
β-strand423-433112
β-strand44613
β-strand448-458112
α-helix478-48811
α-helix493-5008
α-helix509-52012
β-strand525-52624
β-strand532-53324
β-strand539-54355
α-helix549-55911
β-strand563-56755
β-strand579-58246
β-strand589-59246
α-helix595-5984
β-strand603-60755
α-helix614-62613
β-strand628-63367
β-strand636-64167
β-strand645-65065
β-strand65418
β-strand65618
α-helix663-6653
α-helix671-6744
β-strand679-68355
α-helix689-70517
α-helix760-77314
α-helix780-7834
α-helix784-80017
α-helix807-82317
α-helix831-84717
α-helix851-86010
Chains 3 and B: 26 helices, 27 β-strands
ElementResiduesLengthSheet
α-helix19-3416
α-helix39-5113
α-helix91-933
β-strand95-9959
α-helix100-1067
α-helix108-1169
α-helix118-13619
β-strand154-15859
β-strand165-166210
β-strand179-1891110
β-strand193-2031110
β-strand208-213610
β-strand234111
β-strand240111
β-strand241-243310
β-strand249-2591110
α-helix2661
β-strand273-279710
α-helix280-2823
β-strand290-3011210
β-strand318-3301310
α-helix341-3433
α-helix344-35512
α-helix359-3657
α-helix375-38612
β-strand391-392212
β-strand398-399212
β-strand405-408413
α-helix415-42511
β-strand429-433513
α-helix434-4363
α-helix439-4413
β-strand444-448514
β-strand455-459514
α-helix461-4644
β-strand468-473613
α-helix475-4773
α-helix480-49112
β-strand494-498515
β-strand503-507515
β-strand510-516713
α-helix529-5324
α-helix537-5415
β-strand545-548413
α-helix556-56813
α-helix581-5822
β-strand589-590210
α-helix653-66614
β-strand670116
α-helix675-68713
α-helix699-71618
β-strand720116
α-helix723-73715
Chains 4 and C: 33 helices, 33 β-strands
ElementResiduesLengthSheet
α-helix187-19913
α-helix226-23611
β-strand240-243417
α-helix246-2494
α-helix255-2617
α-helix265-28218
β-strand302-305417
α-helix314-3163
α-helix319-3213
β-strand325-3351118
α-helix336-3383
β-strand339-3491118
β-strand355-359518
β-strand361119
β-strand364-367419
β-strand370120
β-strand379120
β-strand382-384318
α-helix386-3883
β-strand390-4011218
β-strand414-420718
β-strand432-4421118
β-strand445121
β-strand453-454221
β-strand456122
β-strand458-4671018
α-helix474-4763
α-helix478-4803
α-helix482-49110
α-helix499-5002
α-helix503-51311
α-helix518-5269
α-helix534-54512
β-strand550-551223
β-strand557-558223
β-strand564-568524
α-helix574-58411
β-strand588-591424
α-helix593-5953
β-strand596125
α-helix598-6025
β-strand603-607526
β-strand614-618526
α-helix620-6234
β-strand627-632624
α-helix634-6363
α-helix639-6435
α-helix645-6506
β-strand653-658627
β-strand661-666627
β-strand669-675724
α-helix676-6772
α-helix688-6903
α-helix696-6994
β-strand704-708524
α-helix714-72613
α-helix744-75613
β-strand761-762228
α-helix764-77916
α-helix791-7933
α-helix795-81016
β-strand816-817228
α-helix819-83315
β-strand838129
β-strand845129
α-helix847-8493
Chains 5 and D: 30 helices, 28 β-strands
ElementResiduesLengthSheet
β-strand8-12530
α-helix18-203
α-helix24-3714
β-strand40131
β-strand43131
α-helix45-5511
β-strand60-64532
α-helix65-673
α-helix73-819
α-helix83-10422
α-helix134-1352
β-strand136-140532
β-strand156133
β-strand159-1691133
β-strand173-1831133
β-strand189-193533
α-helix222-2254
β-strand241-243333
α-helix245-2473
β-strand249-2591133
α-helix270-2723
β-strand273-279733
α-helix280-2823
β-strand291-3011133
β-strand32312
β-strand325-3351133
α-helix351-36111
α-helix367-3737
α-helix382-39110
β-strand398-399234
β-strand405-406234
β-strand411-416635
α-helix422-43110
β-strand436-440535
β-strand443135
β-strand44617
β-strand452-455436
β-strand462-466536
α-helix468-4714
β-strand476-480535
α-helix487-49913
β-strand501-505537
β-strand510-514537
β-strand518-523635
α-helix524-5252
α-helix537-5393
α-helix544-5496
β-strand552-556535
α-helix562-57716
α-helix579-5813
α-helix587-5904
α-helix596-60712
α-helix616-63924
α-helix643-6442
α-helix650-66516
α-helix674-68714
α-helix688-6936
α-helix695-6973
Chains 6 and E: 31 helices, 34 β-strands
ElementResiduesLengthSheet
α-helix100-1034
α-helix106-12015
β-strand124138
β-strand133138
α-helix135-14612
β-strand150-154539
α-helix155-1606
α-helix166-1727
α-helix174-19219
α-helix194-1974
β-strand198140
α-helix2591
β-strand260140
α-helix261-2622
β-strand264-268539
β-strand27413
α-helix276-2783
α-helix281-2833
β-strand287-297113
β-strand301-311113
β-strand317-32263
β-strand344-34633
β-strand352-362113
α-helix365-3673
β-strand376-38273
α-helix383-3853
β-strand393-405133
α-helix407-4093
β-strand418-421419
β-strand428141
β-strand446-447219
β-strand450-461123
α-helix513-5219
α-helix525-5317
α-helix541-55212
β-strand557-558242
α-helix5631
β-strand564-565242
α-helix5661
β-strand571-574443
α-helix582-59110
β-strand595-599543
β-strand603127
α-helix605-6084
β-strand610-614544
β-strand621-625544
β-strand634-639643
α-helix641-6433
α-helix646-65712
β-strand660-665645
β-strand668-673645
β-strand676-682743
β-strand686146
β-strand688146
α-helix695-6984
α-helix703-7075
β-strand711-714443
α-helix721-73717
α-helix748-75811
β-strand764147
α-helix767-78317
α-helix797-81418
β-strand818147
α-helix821-83414
α-helix837-8393
Chains 7 and F: 36 helices, 31 β-strands
ElementResiduesLengthSheet
β-strand7-8210
α-helix14-2512
α-helix62-7110
β-strand77-81548
α-helix82-9413
α-helix102-1087
α-helix110-12314
α-helix138-16427
α-helix180-1889
α-helix195-1973
β-strand201-205548
α-helix206-2083
α-helix209-2146
α-helix232-2343
β-strand238-2481122
β-strand252-2621122
β-strand267-272622
β-strand277-278221
β-strand283149
α-helix287-2915
β-strand297149
β-strand298-300322
β-strand306-3161122
α-helix319-3213
α-helix3231
β-strand330-336722
α-helix337-3393
β-strand348-3591222
α-helix364-3663
β-strand371110
β-strand372-3831222
α-helix395-40612
α-helix410-4156
α-helix426-43712
β-strand442-445450
β-strand448-450350
β-strand456-460551
α-helix466-47611
β-strand480-484551
α-helix490-4923
β-strand495-499552
β-strand506-510552
α-helix512-5154
β-strand519-524651
α-helix526-5283
α-helix531-54313
β-strand545-550625
β-strand553-558625
β-strand561-567751
α-helix580-5823
α-helix588-5914
β-strand596-600551
α-helix606-62116
α-helix631-6344
α-helix635-64612
β-strand651153
α-helix654-67623
α-helix686-70318
β-strand707153
α-helix710-72213
α-helix725-7284
α-helix741-75212
β-strand759-761354
α-helix762-77211
α-helix776-78813
β-strand792-795454
β-strand800-803454

Molecules and chains

MoleculeChainsTypeLengthOrganismUniProt
DNA replication licensing factor MCM22, Aprotein868Saccharomyces cerevisiaeP29469 (AlphaFold model)
DNA replication licensing factor MCM33, Bprotein1006Saccharomyces cerevisiaeP24279 (AlphaFold model)
DNA replication licensing factor MCM44, Cprotein933Saccharomyces cerevisiaeP30665 (AlphaFold model)
Minichromosome maintenance protein 55, Dprotein775Saccharomyces cerevisiaeP29496 (AlphaFold model)
DNA replication licensing factor MCM66, Eprotein1017Saccharomyces cerevisiaeP53091
DNA replication licensing factor MCM77, Fprotein845Saccharomyces cerevisiaeP38132
DNA (53-mer)XDNA53Saccharomyces cerevisiae
DNA (53-mer)YDNA53Saccharomyces cerevisiae
Sequence of entity 1 (2, A), FASTA
>8RIF_1 DNA replication licensing factor MCM2 (chains 2, A)
MSDNRRRRREEDDSDSENELPPSSPQQHFRGGMNPVSSPIGSPDMINPEGDDNEVDDVPD
IDEVEEQMNEVDLMDDNMYEDYAADHNRDRYDPDQVDDREQQELSLSERRRIDAQLNERD
RLLRNVAYIDDEDEEQEGAAQLDEMGLPVQRRRRRRQYEDLENSDDDLLSDMDIDPLREE
LTLESLSNVKANSYSEWITQPNVSRTIARELKSFLLEYTDETGRSVYGARIRTLGEMNSE
SLEVNYRHLAESKAILALFLAKCPEEMLKIFDLVAMEATELHYPDYARIHSEIHVRISDF
PTIYSLRELRESNLSSLVRVTGVVTRRTGVFPQLKYVKFNCLKCGSILGPFFQDSNEEIR
ISFCTNCKSKGPFRVNGEKTVYRNYQRVTLQEAPGTVPPGRLPRHREVILLADLVDVSKP
GEEVEVTGIYKNNYDGNLNAKNGFPVFATIIEANSIKRREGNTANEGEEGLDVFSWTEEE
EREFRKISRDRGIIDKIISSMAPSIYGHRDIKTAVACSLFGGVPKNVNGKHSIRGDINVL
LLGDPGTAKSQILKYVEKTAHRAVFATGQGASAVGLTASVRKDPITKEWTLEGGALVLAD
KGVCLIDEFDKMNDQDRTSIHEAMEQQSISISKAGIVTTLQARCSIIAAANPNGGRYNST
LPLAQNVSLTEPILSRFDILCVVRDLVDEEADERLATFVVDSHVRSHPENDEDREGEELK
NNGESAIEQGEDEINEQLNARQRRLQRQRKKEEEISPIPQELLMKYIHYARTKIYPKLHQ
MDMDKVSRVYADLRRESISTGSFPITVRHLESILRIAESFAKMRLSEFVSSYDLDRAIKV
VVDSFVDAQKVSVRRQLRRSFAIYTLGH
Sequence of entity 2 (3, B), FASTA
>8RIF_2 DNA replication licensing factor MCM3 (chains 3, B)
MKRRWKKNFIAVSAANRFKKISSSGALENLYFQGEMEGSTGFDGDATTFFAPDAVFGDRV
RRFQEFLDTFTSYRDSVRSIQVYNSNNAANYNDDQDDADERDLLGDDDGDDLEKEKKAAS
STSLNILPHRIIISLDDLREFDRSFWSGILVEPAYFIPPAEKALTDLADSMDDVPHPNAS
AVSSRHPWKLSFKGSFGAHALSPRTLTAQHLNKLVSVEGIVTKTSLVRPKLIRSVHYAAK
TGRFHYRDYTDATTTLTTRIPTPAIYPTEDTEGNKLTTEYGYSTFIDHQRITVQEMPEMA
PAGQLPRSIDVILDDDLVDKTKPGDRVNVVGVFKSLGAGGMNQSNSNTLIGFKTLILGNT
VYPLHARSTGVAARQMLTDFDIRNINKLSKKKDIFDILSQSLAPSIYGHDHIKKAILLML
MGGVEKNLENGSHLRGDINILMVGDPSTAKSQLLRFVLNTASLAIATTGRGSSGVGLTAA
VTTDRETGERRLEAGAMVLADRGVVCIDEFDKMTDVDRVAIHEVMEQQTVTIAKAGIHTT
LNARCSVIAAANPVFGQYDVNRDPHQNIALPDSLLSRFDLLFVVTDDINEIRDRSISEHV
LRTHRYLPPGYLEGEPVRERLNLSLAVGEDADINPEEHSNSGAGVENEGEDDEDHVFEKF
NPLLQAGAKLAKNKGNYNGTEIPKLVTIPFLRKYVQYAKERVIPQLTQEAINVIVKNYTD
LRNDDNTKKSPITARTLETLIRLATAHAKVRLSKTVNKVDAKVAANLLRFALLGEDIGND
IDEEESEYEEALSKRSPQKSPKKRQRVRQPASNSGSPIKSTPRRSTASSVNATPSSARRI
LRFQDDEQNAGEDDNDIMSPLPADEEAELQRRLQLGLRVSPRRREHLHAPEEGSSGPLTE
VGTPRLPNVSSAGQDDEQQQSVISFDNVEPGTISTGRLSLISGIIARLMQTEIFEEESYP
VASLFERINEELPEEEKFSAQEYLAGLKIMSDRNNLMVADDKVWRV
Sequence of entity 3 (4, C), FASTA
>8RIF_3 DNA replication licensing factor MCM4 (chains 4, C)
MSQQSSSPTKEDNNSSSPVVPNPDSVPPQLSSPALFYSSSSSQGDIYGRNNSQNLSQGEG
NIRAAIGSSPLNFPSSSQRQNSDVFQSQGRQGRIRSSASASGRSRYHSDLRSDRALPTSS
SSLGRNGQNRVHMRRNDIHTSDLSSPRRIVDFDTRSGVNTLDTSSSSAPPSEASEPLRII
WGTNVSIQECTTNFRNFLMSFKYKFRKILDEREEFINNTTDEELYYIKQLNEMRELGTSN
LNLDARNLLAYKQTEDLYHQLLNYPQEVISIMDQTIKDCMVSLIVDNNLDYDLDEIETKF
YKVRPYNVGSCKGMRELNPNDIDKLINLKGLVLRSTPVIPDMKVAFFKCNVCDHTMAVEI
DRGVIQEPARCERIDCNEPNSMSLIHNRCSFADKQVIKLQETPDFVPDGQTPHSISLCVY
DELVDSCRAGDRIEVTGTFRSIPIRANSRQRVLKSLYKTYVDVVHVKKVSDKRLDVDTST
IEQELMQNKVDHNEVEEVRQITDQDLAKIREVAAREDLYSLLARSIAPSIYELEDVKKGI
LLQLFGGTNKTFTKGGRYRGDINILLCGDPSTSKSQILQYVHKITPRGVYTSGKGSSAVG
LTAYITRDVDTKQLVLESGALVLSDGGVCCIDEFDKMSDSTRSVLHEVMEQQTISIAKAG
IITTLNARSSILASANPIGSRYNPNLPVTENIDLPPPLLSRFDLVYLVLDKVDEKNDREL
AKHLTNLYLEDKPEHISQDDVLPVEFLTMYISYAKEHIHPIITEAAKTELVRAYVGMRKM
GDDSRSDEKRITATTRQLESMIRLAEAHAKMKLKNVVELEDVQEAVRLIRSAIKDYATDP
KTGKIDMNLVQTGKSVIQRKLQEDLSREIMNVLKDQASDSMSFNELIKQINEHSQDRVES
SDIQEALSRLQQEDKVIVLGEGVRRSVRLNNRV
Sequence of entity 4 (5, D), FASTA
>8RIF_4 Minichromosome maintenance protein 5 (chains 5, D)
MSFDRPEIYSAPVLQGESPNDDDNTEIIKSFKNFILEFRLDSQFIYRDQLRNNILVKNYS
LTVNMEHLIGYNEDIYKKLSDEPSDIIPLFETAITQVAKRISILSRAQSANNNDKDPENT
SMDTDSLLLNSLPTFQLILNSNANQIPLRDLDSEHVSKIVRLSGIIISTSVLSSRATYLS
IMCRNCRHTTSITINNFNSITGNTVSLPRSCLSTIESESSMANESNIGDESTKKNCGPDP
YIIIHESSKFIDQQFLKLQEIPELVPVGEMPRNLTMTCDRYLTNKVIPGTRVTIVGIYSI
YNSKNGAGSGRSGGGNGGSGVAIRTPYIKILGIQSDVETSSIWNSVTMFTEEEEEEFLQL
SRNPKLYEILTNSIAPSIFGNEDIKKAIVCLLMGGSKKILPDGMRLRGDINVLLLGDPGT
AKSQLLKFVEKVSPIAVYTSGKGSSAAGLTASVQRDPMTREFYLEGGAMVLADGGVVCID
EFDKMRDEDRVAIHEAMEQQTISIAKAGITTVLNSRTSVLAAANPIYGRYDDLKSPGDNI
DFQTTILSRFDMIFIVKDDHNEERDISIANHVINIHTGNANAMQNQQEENGSEISIEKMK
RYITYCRLKCAPRLSPQAAEKLSSNFVTIRKQLLINELESTERSSIPITIRQLEAIIRIT
ESLAKLELSPIAQERHVDEAIRLFQASTMDAASQDPIGGLNQASGTSLSEIRRFEQELKR
RLPIGWSTSYQTLRREFVDTHRFSQLALDKALYALEKHETIQLRHQGQNIYRSGV
Sequence of entity 5 (6, E), FASTA
>8RIF_5 DNA replication licensing factor MCM6 (chains 6, E)
MSSPFPADTPSSNRPSNSSPPPSSIGAGFGSSSGLDSQIGSRLHFPSSSQPHVSNSQTGP
FVNDSTQFSSQRLQTDGSATNDMEGNEPARSFKSRALNHVKKVDDVTGEKVREAFEQFLE
DFSVQSTDTGEVEKVYRAQIEFMKIYDLNTIYIDYQHLSMRENGALAMAISEQYYRFLPF
LQKGLRRVVRKYAPELLNTSDSLKRSEGDEGQADEDEQQDDDMNGSSLPRDSGSSAAPGN
GTSAMATRSITTSTSPEQTERVFQISFFNLPTVHRIRDIRSEKIGSLLSISGTVTRTSEV
RPELYKASFTCDMCRAIVDNVEQSFKYTEPTFCPNPSCENRAFWTLNVTRSRFLDWQKVR
IQENANEIPTGSMPRTLDVILRGDSVERAKPGDRCKFTGVEIVVPDVTQLGLPGVKPSST
LDTRGISKTTEGLNSGVTGLRSLGVRDLTYKISFLACHVISIGSNIGASSPDANSNNRET
ELQMAANLQANNVYQDNERDQEVFLNSLSSDEINELKEMVKDEHIYDKLVRSIAPAVFGH
EAVKKGILLQMLGGVHKSTVEGIKLRGDINICVVGDPSTSKSQFLKYVVGFAPRSVYTSG
KASSAAGLTAAVVRDEEGGDYTIEAGALMLADNGICCIDEFDKMDISDQVAIHEAMEQQT
ISIAKAGIHATLNARTSILAAANPVGGRYNRKLSLRGNLNMTAPIMSRFDLFFVILDDCN
EKIDTELASHIVDLHMKRDEAIEPPFSAEQLRRYIKYARTFKPILTKEARSYLVEKYKEL
RKDDAQGFSRSSYRITVRQLESMIRLSEAIARANCVDEITPSFIAEAYDLLRQSIIRVDV
DDVEMDEEFDNIESQSHAASGNNDDNDDGTGSGVITSEPPADIEEGQSEATARPGTSEKK
KTTVTYDKYVSMMNMIVRKIAEVDREGAEELTAVDIVDWYLLQKENDLGSLAEYWEERRL
AFKVIKRLVKDRILMEIHGTRHNLRDLENEENENNKTVYVIHPNCEVLDQLEPQDSS
Sequence of entity 6 (7, F), FASTA
>8RIF_6 DNA replication licensing factor MCM7 (chains 7, F)
MSAALPSIQLPVDYNNLFNEITDFLVTFKQDTLSSDATRNENEDENLDAENIEQHLLEKG
PKYMAMLQKVANRELNSVIIDLDDILQYQNEKFLQGTQADDLVSAIQQNANHFTELFCRA
IDNNMPLPTKEIDYKDDVLDVILNQRRLRNERMLSDRTNEIRSENLMDTTMDPPSSMNDA
LREVVEDETELFPPNLTRRYFLYFKPLSQNCARRYRKKAISSKPLSVRQIKGDFLGQLIT
VRGIITRVSDVKPAVEVIAYTCDQCGYEVFQEVNSRTFTPLSECTSEECSQNQTKGQLFM
STRASKFSAFQECKIQELSQQVPVGHIPRSLNIHVNGTLVRSLSPGDIVDVTGIFLPAPY
TGFKALKAGLLTETYLEAQFVRQHKKKFASFSLTSDVEERVMELITSGDVYNRLAKSIAP
EIYGNLDVKKALLLLLVGGVDKRVGDGMKIRGDINVCLMGDPGVAKSQLLKAICKISPRG
VYTTGKGSSGVGLTAAVMKDPVTDEMILEGGALVLADNGICCIDEFDKMDESDRTAIHEV
MEQQTISISKAGINTTLNARTSILAAANPLYGRYNPRLSPLDNINLPAALLSRFDILFLM
LDIPSRDDDEKLAEHVTYVHMHNKQPDLDFTPVEPSKMREYIAYAKTKRPVMSEAVNDYV
VQAYIRLRQDSKREMDSKFSFGQATPRTLLGIIRLSQALAKLRLADMVDIDDVEEALRLV
RVSKESLYQETNKSKEDESPTTKIFTIIKKMLQETGKNTLSYENIVKTVRLRGFTMLQLS
NCIQEYSYLNVWHLINEGNTLKFVDDGTMDTDQEDSLVSTPKLAPQTTASANVSAQDSDI
DLQDA
Sequence of entity 7 (X), FASTA
>8RIF_7 DNA (53-MER) (chains X)
GCATGCATGCGCATGCATGCATGCATGCTGCATGCATGCATGCGCATGCATGC
Sequence of entity 8 (Y), FASTA
>8RIF_8 DNA (53-MER) (chains Y)
GCATGCATGCGCATGCATGCATGCAGCATGCATGCATGCATGCGCATGCATGC

Ligands and cofactors

IDNameFormulaCopies
ATPAdenosine-5'-triphosphateC10 H16 N5 O13 P32
MGMagnesium ionMg8
ZNZinc ionZn10
ADPAdenosine-5'-diphosphateC10 H15 N5 O10 P26

Primary citation

Cell cycle regulation has shaped replication origins in budding yeast. Lim, C.T., Miller, T.C.R., Tan, K.W. et al. Nat Struct Mol Biol (2025) 32:1697-1707. DOI 10.1038/s41594-025-01591-9 · PubMed

Other PDB entries of the same protein (UniProt P29469 (AlphaFold model), which also has an AlphaFold model), best resolution first:

Browse structure collections

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