28VY: SCMGE assembled on ARS1 DNA with Sld2 and RPA

sCMGE assembled on ARS1 DNA with Sld2 and RPA. Determined by electron microscopy at 2.7 Å resolution. Released 13 May 2026.

Method: Electron microscopy
Resolution: 2.7 Å
Organisms: Saccharomyces cerevisiae, synthetic construct
Chains: 15
Atoms: 54,130
Mol. weight: 1147.98 kDa
Ligands: ATP, ADP, ZN, MG
Released: 13 May 2026

Explore 28VY in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

28VY contains 350 α-helices and 289 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 2: 34 helices, 32 β-strands

Element	Residues	Length	Sheet
α-helix	184-186	3
α-helix	194-199	6
α-helix	201-217	17
β-strand	219	1	1
β-strand	225	1	1
α-helix	226-236	11
β-strand	241-245	5	2
α-helix	246-252	7
α-helix	254-262	9
α-helix	264-279	16
α-helix	286-288	3
β-strand	294-298	5	2
β-strand	304	1	3
α-helix	311-313	3
β-strand	317-327	11	3
β-strand	331-337	7	3
β-strand	339-341	3	4
β-strand	347	1	4
β-strand	351-352	2	3
α-helix	353-354	2
β-strand	364	1	5
β-strand	369	1	5
β-strand	373-375	3	4
β-strand	381-391	11	3
α-helix	398	1
β-strand	405-411	7	3
α-helix	412-414	3
β-strand	423-433	11	3
α-helix	436-442	7
β-strand	446	1	6
β-strand	448-458	11	3
α-helix	478-487	10
α-helix	493-499	7
α-helix	509-520	12
β-strand	525-526	2	7
β-strand	532-533	2	7
β-strand	539-543	5	8
α-helix	549-559	11
β-strand	563-567	5	8
β-strand	571	1	9
α-helix	573-576	4
β-strand	578-583	6	10
β-strand	588-593	6	10
α-helix	595-598	4
β-strand	602-607	6	8
α-helix	609-611	3
α-helix	614-617	4
α-helix	618-620	3
α-helix	621-625	5
β-strand	628-633	6	11
β-strand	636-641	6	11
β-strand	644-650	7	8
α-helix	651-652	2
α-helix	653-655	3
α-helix	663-665	3
α-helix	671-674	4
β-strand	679-683	5	8
α-helix	689-705	17
α-helix	740-753	14
α-helix	760-773	14
β-strand	777	1	12
α-helix	783-798	16
α-helix	807-823	17
β-strand	828	1	12
α-helix	831-847	17
α-helix	854-861	8

Chain 3: 28 helices, 27 β-strands

Element	Residues	Length	Sheet
α-helix	18-34	17
α-helix	36-54	19
β-strand	95-99	5	13
α-helix	100-106	7
α-helix	108-116	9
α-helix	118-120	3
α-helix	122-135	14
α-helix	144-148	5
β-strand	154-158	5	13
α-helix	162-164	3
β-strand	165-166	2	14
α-helix	173-175	3
β-strand	178-189	12	14
β-strand	193-203	11	14
β-strand	208-213	6	14
β-strand	234	1	15
β-strand	240	1	15
β-strand	241-243	3	14
β-strand	249-259	11	14
α-helix	262-264	3
α-helix	266	1
β-strand	273-279	7	14
α-helix	280-282	3
β-strand	291-301	11	14
β-strand	318-328	11	14
α-helix	344-354	11
α-helix	359-366	8
α-helix	375-386	12
β-strand	391-392	2	16
β-strand	398-399	2	16
β-strand	405-409	5	17
α-helix	415-425	11
β-strand	429-433	5	17
β-strand	437	1	18
α-helix	439-443	5
β-strand	444-448	5	19
β-strand	455-459	5	19
α-helix	461-464	4
β-strand	469-473	5	17
α-helix	475-477	3
α-helix	480-492	13
β-strand	494-499	6	20
β-strand	502-507	6	20
β-strand	511-516	6	17
α-helix	529-532	4
α-helix	537-540	4
β-strand	545-549	5	17
α-helix	555-569	15
α-helix	580-582	3
α-helix	653-666	14
β-strand	670	1	21
α-helix	673-688	16
α-helix	699-716	18
β-strand	720	1	21
α-helix	723-737	15

Chain 4: 37 helices, 35 β-strands

Element	Residues	Length	Sheet
α-helix	187-199	13
α-helix	203-208	6
α-helix	226-235	10
β-strand	240-244	5	22
α-helix	246-249	4
α-helix	252-254	3
α-helix	256-263	8
α-helix	265-285	21
α-helix	293-298	6
β-strand	302-306	5	22
α-helix	311-313	3
α-helix	314-316	3
α-helix	319-321	3
β-strand	325-335	11	23
β-strand	339-349	11	23
β-strand	355-359	5	23
β-strand	361	1	24
β-strand	364	1	24
β-strand	370	1	25
α-helix	374-376	3
β-strand	379	1	25
β-strand	382-384	3	23
α-helix	386-388	3
β-strand	390-401	12	23
α-helix	403-405	3
α-helix	406-407	2
α-helix	411-413	3
β-strand	414-420	7	23
α-helix	421-423	3
β-strand	432-442	11	23
β-strand	445	1	26
β-strand	453-454	2	26
β-strand	456	1	27
β-strand	458-468	11	23
α-helix	503-513	11
α-helix	518-526	9
α-helix	534-545	12
β-strand	550-552	3	28
β-strand	556-558	3	28
β-strand	564-568	5	29
α-helix	574-584	11
β-strand	588-592	5	29
β-strand	604-607	4	30
β-strand	614-617	4	30
α-helix	620-623	4
β-strand	627-632	6	29
α-helix	639-651	13
β-strand	653-658	6	31
β-strand	661-666	6	31
β-strand	669-675	7	29
α-helix	676-677	2
α-helix	688-691	4
α-helix	696-699	4
β-strand	704-708	5	29
α-helix	714-727	14
α-helix	744-757	14
β-strand	761-762	2	32
α-helix	764-778	15
α-helix	795-811	17
β-strand	816-817	2	32
α-helix	819-836	18
β-strand	838	1	33
β-strand	845	1	33
α-helix	847-852	6
α-helix	856-875	20
α-helix	877-879	3
β-strand	880-882	3	34
α-helix	883-893	11
α-helix	900-911	12
β-strand	916-919	4	34
α-helix	922-924	3
β-strand	926-929	4	34

Chain 5: 36 helices, 27 β-strands

Element	Residues	Length	Sheet
α-helix	21-37	17
α-helix	45-55	11
β-strand	60-64	5	35
α-helix	65-71	7
α-helix	73-81	9
α-helix	83-102	20
α-helix	134-135	2
β-strand	136-140	5	35
α-helix	143-147	5
α-helix	148-150	3
α-helix	153-155	3
β-strand	159-169	11	36
β-strand	173-183	11	36
β-strand	189-193	5	36
β-strand	241-259	19	36
α-helix	262-264	3
α-helix	265-266	2
β-strand	273-279	7	36
α-helix	280-282	3
β-strand	291-301	11	36
β-strand	323	1	3
β-strand	325-335	11	36
α-helix	339-343	5
α-helix	351-361	11
α-helix	366-372	7
α-helix	382-393	12
β-strand	398-399	2	37
β-strand	405-406	2	37
β-strand	412-416	5	38
α-helix	422-432	11
β-strand	436-440	5	38
β-strand	444	1	11
α-helix	446-449	4
β-strand	451-455	5	39
β-strand	462-466	5	39
α-helix	468-471	4
β-strand	476-480	5	38
α-helix	482-484	3
α-helix	487-490	4
α-helix	491-493	3
α-helix	494-498	5
β-strand	501-506	6	18
β-strand	509-514	6	18
β-strand	518-523	6	38
α-helix	536-538	3
α-helix	544-547	4
β-strand	552-556	5	38
α-helix	562-576	15
α-helix	580-589	10
α-helix	596-609	14
β-strand	613-614	2	40
α-helix	616-639	24
α-helix	650-666	17
β-strand	671-672	2	40
α-helix	674-684	11
α-helix	685-689	5
α-helix	690-692	3
α-helix	708-721	14
β-strand	727-729	3	41
α-helix	730-737	8
α-helix	745-757	13
β-strand	761-764	4	41
β-strand	769-772	4	41

Chain 6: 30 helices, 29 β-strands

Element	Residues	Length	Sheet
α-helix	106-120	15
β-strand	124-125	2	42
β-strand	132-133	2	42
α-helix	135-146	12
β-strand	150-154	5	43
α-helix	155-159	5
α-helix	162-164	3
α-helix	165-172	8
α-helix	174-192	19
α-helix	194-197	4
β-strand	264-268	5	43
β-strand	274	1	6
α-helix	276-278	3
α-helix	281-283	3
β-strand	287-297	11	6
β-strand	301-311	11	6
β-strand	317-322	6	6
β-strand	344-346	3	6
β-strand	352-362	11	6
α-helix	365-367	3
α-helix	369	1
β-strand	376-382	7	6
α-helix	383-385	3
β-strand	394-405	12	6
α-helix	407-410	4
β-strand	437	1	44
β-strand	438	1	24
β-strand	448	1	44
β-strand	450-461	12	6
α-helix	501-506	6
α-helix	510-520	11
α-helix	525-531	7
α-helix	541-551	11
β-strand	557-558	2	45
β-strand	564-565	2	45
β-strand	571-575	5	46
α-helix	581-591	11
β-strand	596-599	4	46
β-strand	610-613	4	47
β-strand	622-625	4	47
α-helix	627-630	4
β-strand	634-639	6	46
α-helix	646-649	4
α-helix	652-657	6
β-strand	660-665	6	9
β-strand	668-673	6	9
β-strand	676-682	7	46
α-helix	683-684	2
α-helix	695-698	4
α-helix	703-708	6
β-strand	711-715	5	46
α-helix	721-737	17
α-helix	738-741	4
α-helix	748-758	11
β-strand	764-765	2	48
α-helix	767-784	18
α-helix	797-814	18
β-strand	818-819	2	48
α-helix	821-833	13

Chain 7: 32 helices, 27 β-strands

Element	Residues	Length	Sheet
β-strand	7-8	2	14
α-helix	14-27	14
α-helix	62-71	10
β-strand	77-81	5	49
α-helix	82-94	13
α-helix	102-108	7
α-helix	110-123	14
α-helix	125-128	4
α-helix	138-151	14
α-helix	194-197	4
β-strand	201-205	5	49
α-helix	206-208	3
α-helix	209-214	6
β-strand	224-225	2	27
α-helix	227-229	3
α-helix	232-234	3
β-strand	238-248	11	27
α-helix	249-251	3
β-strand	252-262	11	27
β-strand	268-272	5	27
β-strand	277-278	2	26
β-strand	283	1	50
α-helix	287-291	5
β-strand	297	1	50
β-strand	298-300	3	27
β-strand	306-316	11	27
α-helix	319-321	3
α-helix	323	1
β-strand	330-336	7	27
β-strand	348-358	11	27
α-helix	363-368	6
β-strand	371	1	14
β-strand	373-383	11	27
α-helix	398-407	10
α-helix	410-417	8
α-helix	426-435	10
β-strand	442	1	51
β-strand	450	1	51
β-strand	456-460	5	52
α-helix	466-474	9
β-strand	480-483	4	52
β-strand	498-499	2	53
β-strand	506-507	2	53
α-helix	512-515	4
β-strand	520-523	4	52
α-helix	531-541	11
β-strand	545-550	6	54
β-strand	553-558	6	54
β-strand	562-567	6	52
α-helix	580-583	4
α-helix	588-592	5
β-strand	596-600	5	52
α-helix	606-622	17
α-helix	632-634	3
α-helix	635-645	11
α-helix	654-674	21
α-helix	686-702	17
α-helix	710-723	14
α-helix	725-728	4

Chain C: 12 helices, 5 β-strands

Element	Residues	Length	Sheet
α-helix	-1-1	3
α-helix	6-12	7
β-strand	15-20	6	55
β-strand	24	1	56
α-helix	28-31	4
β-strand	38	1	56
β-strand	43-47	5	55
α-helix	48-54	7
β-strand	72-74	3	55
α-helix	75-77	3
α-helix	78-80	3
α-helix	82-90	9
α-helix	97-100	4
α-helix	104-115	12
α-helix	119-140	22
α-helix	160-166	7
α-helix	171-191	21

Chain D: 12 helices, 6 β-strands

Element	Residues	Length	Sheet
α-helix	56-72	17
α-helix	76-79	4
α-helix	81-110	30
α-helix	120-122	3
α-helix	123-153	31
α-helix	155-163	9
α-helix	171-174	4
α-helix	177-197	21
α-helix	199-201	3
α-helix	204-206	3
β-strand	227-232	6	57
α-helix	240-242	3
β-strand	247-248	2	58
β-strand	254-260	7	58
β-strand	265-270	6	58
β-strand	274-278	5	57
α-helix	279-287	9
β-strand	291-292	2	57

5 more chain groups are not listed. Open the entry in the viewer and use the sequence panel to see them.

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
DNA replication licensing factor MCM2	2	protein	868	Saccharomyces cerevisiae	P29469 (AlphaFold model)
DNA replication licensing factor MCM3	3	protein	1006	Saccharomyces cerevisiae	P24279 (AlphaFold model)
DNA replication licensing factor MCM4	4	protein	933	Saccharomyces cerevisiae	P30665 (AlphaFold model)
Minichromosome maintenance protein 5	5	protein	775	Saccharomyces cerevisiae	P29496 (AlphaFold model)
DNA replication licensing factor MCM6	6	protein	1017	Saccharomyces cerevisiae	P53091
DNA replication licensing factor MCM7	7	protein	845	Saccharomyces cerevisiae	P38132
DNA (5'-d(pgpaptpcpgpaptpcpgpaptpcpgpaptpcpgpaptpcpgpap*t)-3')	A, B	DNA	27	synthetic construct
DNA replication complex GINS protein PSF3	C	protein	217	Saccharomyces cerevisiae	Q12146
DNA replication complex GINS protein SLD5	D	protein	294	Saccharomyces cerevisiae	Q03406
Cell division control protein 45	E	protein	657	Saccharomyces cerevisiae	Q08032
DNA polymerase epsilon subunit B	F	protein	689	Saccharomyces cerevisiae	P24482
DNA polymerase epsilon catalytic subunit A	G	protein	2222	Saccharomyces cerevisiae	P21951

2 more molecules are not listed.

Sequence of entity 1 (2), FASTA

>28VY_1 DNA replication licensing factor MCM2 (chains 2)
MSDNRRRRREEDDSDSENELPPSSPQQHFRGGMNPVSSPIGSPDMINPEGDDNEVDDVPD
IDEVEEQMNEVDLMDDNMYEDYAADHNRDRYDPDQVDDREQQELSLSERRRIDAQLNERD
RLLRNVAYIDDEDEEQEGAAQLDEMGLPVQRRRRRRQYEDLENSDDDLLSDMDIDPLREE
LTLESLSNVKANSYSEWITQPNVSRTIARELKSFLLEYTDETGRSVYGARIRTLGEMNSE
SLEVNYRHLAESKAILALFLAKCPEEMLKIFDLVAMEATELHYPDYARIHSEIHVRISDF
PTIYSLRELRESNLSSLVRVTGVVTRRTGVFPQLKYVKFNCLKCGSILGPFFQDSNEEIR
ISFCTNCKSKGPFRVNGEKTVYRNYQRVTLQEAPGTVPPGRLPRHREVILLADLVDVSKP
GEEVEVTGIYKNNYDGNLNAKNGFPVFATIIEANSIKRREGNTANEGEEGLDVFSWTEEE
EREFRKISRDRGIIDKIISSMAPSIYGHRDIKTAVACSLFGGVPKNVNGKHSIRGDINVL
LLGDPGTAKSQILKYVEKTAHRAVFATGQGASAVGLTASVRKDPITKEWTLEGGALVLAD
KGVCLIDEFDKMNDQDRTSIHEAMEQQSISISKAGIVTTLQARCSIIAAANPNGGRYNST
LPLAQNVSLTEPILSRFDILCVVRDLVDEEADERLATFVVDSHVRSHPENDEDREGEELK
NNGESAIEQGEDEINEQLNARQRRLQRQRKKEEEISPIPQELLMKYIHYARTKIYPKLHQ
MDMDKVSRVYADLRRESISTGSFPITVRHLESILRIAESFAKMRLSEFVSSYDLDRAIKV
VVDSFVDAQKVSVRRQLRRSFAIYTLGH

Sequence of entity 2 (3), FASTA

>28VY_2 DNA replication licensing factor MCM3 (chains 3)
MKRRWKKNFIAVSAANRFKKISSSGALENLYFQGEMEGSTGFDGDATTFFAPDAVFGDRV
RRFQEFLDTFTSYRDSVRSIQVYNSNNAANYNDDQDDADERDLLGDDDGDDLEKEKKAAS
STSLNILPHRIIISLDDLREFDRSFWSGILVEPAYFIPPAEKALTDLADSMDDVPHPNAS
AVSSRHPWKLSFKGSFGAHALSPRTLTAQHLNKLVSVEGIVTKTSLVRPKLIRSVHYAAK
TGRFHYRDYTDATTTLTTRIPTPAIYPTEDTEGNKLTTEYGYSTFIDHQRITVQEMPEMA
PAGQLPRSIDVILDDDLVDKTKPGDRVNVVGVFKSLGAGGMNQSNSNTLIGFKTLILGNT
VYPLHARSTGVAARQMLTDFDIRNINKLSKKKDIFDILSQSLAPSIYGHDHIKKAILLML
MGGVEKNLENGSHLRGDINILMVGDPSTAKSQLLRFVLNTASLAIATTGRGSSGVGLTAA
VTTDRETGERRLEAGAMVLADRGVVCIDEFDKMTDVDRVAIHEVMEQQTVTIAKAGIHTT
LNARCSVIAAANPVFGQYDVNRDPHQNIALPDSLLSRFDLLFVVTDDINEIRDRSISEHV
LRTHRYLPPGYLEGEPVRERLNLSLAVGEDADINPEEHSNSGAGVENEGEDDEDHVFEKF
NPLLQAGAKLAKNKGNYNGTEIPKLVTIPFLRKYVQYAKERVIPQLTQEAINVIVKNYTD
LRNDDNTKKSPITARTLETLIRLATAHAKVRLSKTVNKVDAKVAANLLRFALLGEDIGND
IDEEESEYEEALSKRSPQKSPKKRQRVRQPASNSGSPIKSTPRRSTASSVNATPSSARRI
LRFQDDEQNAGEDDNDIMSPLPADEEAELQRRLQLGLRVSPRRREHLHAPEEGSSGPLTE
VGTPRLPNVSSAGQDDEQQQSVISFDNVEPGTISTGRLSLISGIIARLMQTEIFEEESYP
VASLFERINEELPEEEKFSAQEYLAGLKIMSDRNNLMVADDKVWRV

Sequence of entity 3 (4), FASTA

>28VY_3 DNA replication licensing factor MCM4 (chains 4)
MSQQSSSPTKEDNNSSSPVVPNPDSVPPQLSSPALFYSSSSSQGDIYGRNNSQNLSQGEG
NIRAAIGSSPLNFPSSSQRQNSDVFQSQGRQGRIRSSASASGRSRYHSDLRSDRALPTSS
SSLGRNGQNRVHMRRNDIHTSDLSSPRRIVDFDTRSGVNTLDTSSSSAPPSEASEPLRII
WGTNVSIQECTTNFRNFLMSFKYKFRKILDEREEFINNTTDEELYYIKQLNEMRELGTSN
LNLDARNLLAYKQTEDLYHQLLNYPQEVISIMDQTIKDCMVSLIVDNNLDYDLDEIETKF
YKVRPYNVGSCKGMRELNPNDIDKLINLKGLVLRSTPVIPDMKVAFFKCNVCDHTMAVEI
DRGVIQEPARCERIDCNEPNSMSLIHNRCSFADKQVIKLQETPDFVPDGQTPHSISLCVY
DELVDSCRAGDRIEVTGTFRSIPIRANSRQRVLKSLYKTYVDVVHVKKVSDKRLDVDTST
IEQELMQNKVDHNEVEEVRQITDQDLAKIREVAAREDLYSLLARSIAPSIYELEDVKKGI
LLQLFGGTNKTFTKGGRYRGDINILLCGDPSTSKSQILQYVHKITPRGVYTSGKGSSAVG
LTAYITRDVDTKQLVLESGALVLSDGGVCCIDEFDKMSDSTRSVLHEVMEQQTISIAKAG
IITTLNARSSILASANPIGSRYNPNLPVTENIDLPPPLLSRFDLVYLVLDKVDEKNDREL
AKHLTNLYLEDKPEHISQDDVLPVEFLTMYISYAKEHIHPIITEAAKTELVRAYVGMRKM
GDDSRSDEKRITATTRQLESMIRLAEAHAKMKLKNVVELEDVQEAVRLIRSAIKDYATDP
KTGKIDMNLVQTGKSVIQRKLQEDLSREIMNVLKDQASDSMSFNELIKQINEHSQDRVES
SDIQEALSRLQQEDKVIVLGEGVRRSVRLNNRV

Sequence of entity 4 (5), FASTA

>28VY_4 Minichromosome maintenance protein 5 (chains 5)
MSFDRPEIYSAPVLQGESPNDDDNTEIIKSFKNFILEFRLDSQFIYRDQLRNNILVKNYS
LTVNMEHLIGYNEDIYKKLSDEPSDIIPLFETAITQVAKRISILSRAQSANNNDKDPENT
SMDTDSLLLNSLPTFQLILNSNANQIPLRDLDSEHVSKIVRLSGIIISTSVLSSRATYLS
IMCRNCRHTTSITINNFNSITGNTVSLPRSCLSTIESESSMANESNIGDESTKKNCGPDP
YIIIHESSKFIDQQFLKLQEIPELVPVGEMPRNLTMTCDRYLTNKVIPGTRVTIVGIYSI
YNSKNGAGSGRSGGGNGGSGVAIRTPYIKILGIQSDVETSSIWNSVTMFTEEEEEEFLQL
SRNPKLYEILTNSIAPSIFGNEDIKKAIVCLLMGGSKKILPDGMRLRGDINVLLLGDPGT
AKSQLLKFVEKVSPIAVYTSGKGSSAAGLTASVQRDPMTREFYLEGGAMVLADGGVVCID
EFDKMRDEDRVAIHEAMEQQTISIAKAGITTVLNSRTSVLAAANPIYGRYDDLKSPGDNI
DFQTTILSRFDMIFIVKDDHNEERDISIANHVINIHTGNANAMQNQQEENGSEISIEKMK
RYITYCRLKCAPRLSPQAAEKLSSNFVTIRKQLLINELESTERSSIPITIRQLEAIIRIT
ESLAKLELSPIAQERHVDEAIRLFQASTMDAASQDPIGGLNQASGTSLSEIRRFEQELKR
RLPIGWSTSYQTLRREFVDTHRFSQLALDKALYALEKHETIQLRHQGQNIYRSGV

Sequence of entity 5 (6), FASTA

>28VY_5 DNA replication licensing factor MCM6 (chains 6)
MSSPFPADTPSSNRPSNSSPPPSSIGAGFGSSSGLDSQIGSRLHFPSSSQPHVSNSQTGP
FVNDSTQFSSQRLQTDGSATNDMEGNEPARSFKSRALNHVKKVDDVTGEKVREAFEQFLE
DFSVQSTDTGEVEKVYRAQIEFMKIYDLNTIYIDYQHLSMRENGALAMAISEQYYRFLPF
LQKGLRRVVRKYAPELLNTSDSLKRSEGDEGQADEDEQQDDDMNGSSLPRDSGSSAAPGN
GTSAMATRSITTSTSPEQTERVFQISFFNLPTVHRIRDIRSEKIGSLLSISGTVTRTSEV
RPELYKASFTCDMCRAIVDNVEQSFKYTEPTFCPNPSCENRAFWTLNVTRSRFLDWQKVR
IQENANEIPTGSMPRTLDVILRGDSVERAKPGDRCKFTGVEIVVPDVTQLGLPGVKPSST
LDTRGISKTTEGLNSGVTGLRSLGVRDLTYKISFLACHVISIGSNIGASSPDANSNNRET
ELQMAANLQANNVYQDNERDQEVFLNSLSSDEINELKEMVKDEHIYDKLVRSIAPAVFGH
EAVKKGILLQMLGGVHKSTVEGIKLRGDINICVVGDPSTSKSQFLKYVVGFAPRSVYTSG
KASSAAGLTAAVVRDEEGGDYTIEAGALMLADNGICCIDEFDKMDISDQVAIHEAMEQQT
ISIAKAGIHATLNARTSILAAANPVGGRYNRKLSLRGNLNMTAPIMSRFDLFFVILDDCN
EKIDTELASHIVDLHMKRDEAIEPPFSAEQLRRYIKYARTFKPILTKEARSYLVEKYKEL
RKDDAQGFSRSSYRITVRQLESMIRLSEAIARANCVDEITPSFIAEAYDLLRQSIIRVDV
DDVEMDEEFDNIESQSHAASGNNDDNDDGTGSGVITSEPPADIEEGQSEATARPGTSEKK
KTTVTYDKYVSMMNMIVRKIAEVDREGAEELTAVDIVDWYLLQKENDLGSLAEYWEERRL
AFKVIKRLVKDRILMEIHGTRHNLRDLENEENENNKTVYVIHPNCEVLDQLEPQDSS

Sequence of entity 6 (7), FASTA

>28VY_6 DNA replication licensing factor MCM7 (chains 7)
MSAALPSIQLPVDYNNLFNEITDFLVTFKQDTLSSDATRNENEDENLDAENIEQHLLEKG
PKYMAMLQKVANRELNSVIIDLDDILQYQNEKFLQGTQADDLVSAIQQNANHFTELFCRA
IDNNMPLPTKEIDYKDDVLDVILNQRRLRNERMLSDRTNEIRSENLMDTTMDPPSSMNDA
LREVVEDETELFPPNLTRRYFLYFKPLSQNCARRYRKKAISSKPLSVRQIKGDFLGQLIT
VRGIITRVSDVKPAVEVIAYTCDQCGYEVFQEVNSRTFTPLSECTSEECSQNQTKGQLFM
STRASKFSAFQECKIQELSQQVPVGHIPRSLNIHVNGTLVRSLSPGDIVDVTGIFLPAPY
TGFKALKAGLLTETYLEAQFVRQHKKKFASFSLTSDVEERVMELITSGDVYNRLAKSIAP
EIYGNLDVKKALLLLLVGGVDKRVGDGMKIRGDINVCLMGDPGVAKSQLLKAICKISPRG
VYTTGKGSSGVGLTAAVMKDPVTDEMILEGGALVLADNGICCIDEFDKMDESDRTAIHEV
MEQQTISISKAGINTTLNARTSILAAANPLYGRYNPRLSPLDNINLPAALLSRFDILFLM
LDIPSRDDDEKLAEHVTYVHMHNKQPDLDFTPVEPSKMREYIAYAKTKRPVMSEAVNDYV
VQAYIRLRQDSKREMDSKFSFGQATPRTLLGIIRLSQALAKLRLADMVDIDDVEEALRLV
RVSKESLYQETNKSKEDESPTTKIFTIIKKMLQETGKNTLSYENIVKTVRLRGFTMLQLS
NCIQEYSYLNVWHLINEGNTLKFVDDGTMDTDQEDSLVSTPKLAPQTTASANVSAQDSDI
DLQDA

Sequence of entity 7 (A, B), FASTA

>28VY_7 DNA (5'-D(P*GP*AP*TP*CP*GP*AP*TP*CP*GP*AP*TP*CP*GP*AP*TP*CP*GP*AP*TP*CP*GP*AP*T)-3') (chains A, B)
GATCGATCGATCGATCGATCGATCGAT

Sequence of entity 8 (C), FASTA

>28VY_8 DNA replication complex GINS protein PSF3 (chains C)
MGSSHHHHHHSSGLVPRGSHMASMGYYDIDDVLADGTEFPCKFQYDIPGLGYLENNPGRP
ITKNTKLNLPLWLARILAIVGGDEALVDEEPVPFVELLPPDMFSTKVMNAIKTDPVALDL
HSINSHFFSLAIKWIMLFSEKELANVVSELLLQRAQELNHHASSLSIDLNADSTGKNSAN
TNIATSTFLLKLEEMEKEIYKKSHESYKDTKRWMFKK

Sequence of entity 9 (D), FASTA

>28VY_9 DNA replication complex GINS protein SLD5 (chains D)
MDINIDDILAELDKETTAVDSTKITQGSSSTTHRDANTIVGSSLDLNDKTQIYVSPQQDF
SDLMKSWKNERCSPELLPYPHQLMKRLLNRISMQSQLIENISMGFLDMQNASNANPPMPN
ESKLPLLCMETELERLKFVIRSYIRCRLSKIDKFSLYLRQLNEDENSLISLTDLLSKDEI
KYHDTHSLIWLKLVNDSILKYMPEELQAINDTEGSVNMIDEPDWNKFVFIHVNGPPDGKW
NEDPLLQENEFGKPCYTVTIPDLKEEVELTIGSIYVMRYEVIRDLLRDDKVALI

Sequence of entity 10 (E), FASTA

>28VY_10 Cell division control protein 45 (chains E)
MYYGISQFSEAYNKILRNSSSHSSCQLVIFVSCLNIDALCATKMLSLLFKKQLVQSQIVP
IFGYSELRRHYSQLDDNINSLLLVGFGGVIDLEAFLEIDPQEYVIDTDEKSGEQSFRRDI
YVLDAHRPWNLDNIFGSQIIQCFDDGTVDDTLGEQKEAYYKLLELDEESGDDELSGDEND
NNGGDDEATDADEVTDEDYKDDDGDYKDDDETISNKRGNSSIGPNDLSKRKQRKKQIHEY
EGVLEEYYSQGTTVVNSISAQIYSLLSAIGETNLSNLWLNILGTTSLDIAYAQVYNRLYP
LLQDEVKRLTPSSRNSVKTPDTLTLNIQPDYYLFLLRHSSLYDSFYYSNYVNAKLSLWNE
NGKKRLHKMFARMGIPLSTAQETWLYMDHSIKRELGIIFDKNLDRYGLQDIIRDGFVRTL
GYRGSISASEFVEALTALLEVGNSTDKDSVKINNDNNDDTDGEEEEDNSAQKLTNLRKRW
VSNFWLSWDALDDRKVELLNRGIQLAQDLQRAIFNTGVAILEKKLIKHLRIYRLCVLQDG
PDLDLYRNPLTLLRLGNWLIECCAESEDKQLLPMVLASIDENTDTYLVAGLTPRYPRGLD
TIHTKKPILNNFSMAFQQITAETDAKVRIDNFESSIIEIRREDLSPFLEKLTLSGLL

Sequence of entity 11 (F), FASTA

>28VY_11 DNA polymerase epsilon subunit B (chains F)
MFGSGNVLPVKIQPPLLRPLAYRVLSRKYGLSIKSDGLSALAEFVGTNIGANWRQGPATI
KFLEQFAAVWKQQERGLFIDQSGVKEVIQEMKEREKVEWSHEHPIQHEENILGRTDDDEN
NSDDEMPIAADSSLQNVSLSSPMRQPTERDEYKQPFKPESSKALDWRDYFKVINASQQQR
FSYNPHKMQFIFVPNKKQNGLGGIAGFLPDIEDKVQMFLTRYYLTNDRVMRNENFQNSDM
FNPLSSMVSLQNELSNTNRQQQSSSMSITPIKNLLGRDAQNFLLLGLLNKNFKGNWSLED
PSGSVEIDISQTIPTQGHYYVPGCMVLVEGIYYSVGNKFHVTSMTLPPGERREITLETIG
NLDLLGIHGISNNNFIARLDKDLKIRLHLLEKELTDHKFVILGANLFLDDLKIMTALSKI
LQKLNDDPPTLLIWQGSFTSVPVFASMSSRNISSSTQFKNNFDALATLLSRFDNLTENTT
MIFIPGPNDLWGSMVSLGASGTLPQDPIPSAFTKKINKVCKNVVWSSNPTRIAYLSQEIV
IFRDDLSGRFKRHRLEFPFNESEDVYTENDNMMSKDTDIVPIDELVKEPDQLPQKVQETR
KLVKTILDQGHLSPFLDSLRPISWDLDHTLTLCPIPSTMVLCDTTSAQFDLTYNGCKVIN
PGSFIHNRRARYMEYVPSSKKTIQEEIYI

Sequence of entity 12 (G), FASTA

>28VY_12 DNA polymerase epsilon catalytic subunit A (chains G)
MMFGKKKNNGGSSTARYSAGNKYNTLSNNYALSAQQLLNASKIDDIDSMMGFERYVPPQY
NGRFDAKDIDQIPGRVGWLTNMHATLVSQETLSSGSNGGGNSNDGERVTTNQGISGVDFY
FLDEEGGSFKSTVVYDPYFFIACNDESRVNDVEELVKKYLESCLKSLQIIRKEDLTMDNH
LLGLQKTLIKLSFVNSNQLFEARKLLRPILQDNANNNVQRNIYNVAANGSEKVDAKHLIE
DIREYDVPYHVRVSIDKDIRVGKWYKVTQQGFIEDTRKIAFADPVVMAFDIETTKPPLKF
PDSAVDQIMMISYMIDGEGFLITNREIISEDIEDFEYTPKPEYPGFFTIFNENDEVALLQ
RFFEHIRDVRPTVISTFNGDFFDWPFIHNRSKIHGLDMFDEIGFAPDAEGEYKSSYCSHM
DCFRWVKRDSYLPQGSQGLKAVTQSKLGYNPIELDPELMTPYAFEKPQHLSEYSVSDAVA
TYYLYMKYVHPFIFSLCTIIPLNPDETLRKGTGTLCEMLLMVQAYQHNILLPNKHTDPIE
RFYDGHLLESETYVGGHVESLEAGVFRSDLKNEFKIDPSAIDELLQELPEALKFSVEVEN
KSSVDKVTNFEEIKNQITQKLLELKENNIRNELPLIYHVDVASMYPNIMTTNRLQPDSIK
AERDCASCDFNRPGKTCARKLKWAWRGEFFPSKMDEYNMIKRALQNETFPNKNKFSKKKV
LTFDELSYADQVIHIKKRLTEYSRKVYHRVKVSEIVEREAIVCQRENPFYVDTVKSFRDR
RYEFKGLAKTWKGNLSKIDPSDKHARDEAKKMIVLYDSLQLAHKVILNSFYGYVMRKGSR
WYSMEMAGITCLTGATIIQMARALVERVGRPLELDTDGIWCILPKSFPETYFFTLENGKK
LYLSYPCSMLNYRVHQKFTNHQYQELKDPLNYIYETHSENTIFFEVDGPYKAMILPSSKE
EGKGIKKRYAVFNEDGSLAELKGFELKRRGELQLIKNFQSDIFKVFLEGDTLEGCYSAVA
SVCNRWLDVLDSHGLMLEDEDLVSLICENRSMSKTLKEYEGQKSTSITTARRLGDFLGED
MVKDKGLQCKYIISSKPFNAPVTERAIPVAIFSADIPIKRSFLRRWTLDPSLEDLDIRTI
IDWGYYRERLGSAIQKIITIPAALQGVSNPVPRVEHPDWLKRKIATKEDKFKQTSLTKFF
SKTKNVPTMGKIKDIEDLFEPTVEEDNAKIKIARTTKKKAVSKRKRNQLTNEEDPLVLPS
EIPSMDEDYVGWLNYQKIKWKIQARDRKRRDQLFGNTNSSRERSALGSMIRKQAESYANS
TWEVLQYKDSGEPGVLEVFVTINGKVQNITFHIPKTIYMKFKSQTMPLQKIKNCLIEKSS
ASLPNNPKTSNPAGGQLFKITLPESVFLEEKENCTSIFNDENVLGVFEGTITPHQRAIMD
LGASVTFRSKAMGALGKGIQQGFEMKDLSMAENERYLSGFSMDIGYLLHFPTSIGYEFFS
LFKSWGDTITILVLKPSNQAQEINASSLGQIYKQMFEKKKGKIETYSYLVDIKEDINFEF
VYFTDISKLYRRLSQETTKLKEERGLQFLLLLQSPFITKLLGTIRLLNQMPIVKLSLNEV
LLPQLNWQPTLLKKLVNHVLSSGSWISHLIKLSQYSNIPICNLRLDSMDYIIDVLYARKL
KKENIVLWWNEKAPLPDHGGIQNDFDLNTSWIMNDSEFPKINNSGVYDNVVLDVGVDNLT
VNTILTSALINDAEGSDLVNNNMGIDDKDAVINSPSEFVHDAFSNDALNVLRGMLKEWWD
EALKENSTADLLVNSLASWVQNPNAKLFDGLLRYHVHNLTKKALLQLVNEFSALGSTIVY
ADRNQILIKTNKYSPENCYAYSQYMMKAVRTNPMFSYLDLNIKRYWDLLIWMDKFNFSGL
ACIEIEEKENQDYTAVSQWQLKKFLSPIYQPEFEDWMMIILDSMLKTKQSYLKLNSGTQR
PTQIVNVKKQDKEDSVENSLNGFSHLFSKPLMKRVKKLFKNQQEFILDPQYEADYVIPVL
PGSHLNVKNPLLELVKSLCHVMLLSKSTILEIRTLRKELLKIFELREFAKVAEFKDPSLS
LVVPDFLCEYCFFISDIDFCKAAPESIFSCVRCHKAFNQVLLQEHLIQKLRSDIESYLIQ
DLRCSRCHKVKRDYMSAHCPCAGAWEGTLPRESIVQKLNVFKQVAKYYGFDILLSCIADL
TI

Ligands and cofactors

ID	Name	Formula	Copies
ATP	Adenosine-5'-triphosphate	C10 H16 N5 O13 P3	5
ADP	Adenosine-5'-diphosphate	C10 H15 N5 O10 P2	1
ZN	Zinc ion	Zn	7
MG	Magnesium ion	Mg	6

Primary citation

Structure of the pre-initiation complex explains CMGE biogenesis. Puhringer, T., Canal, B., Palm, G. et al. Nature (2026) 655:1330-1338. DOI 10.1038/s41586-026-10657-7 · PubMed

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

7W8G 2.52 Å, Cryo-EM structure of MCM double hexamer
8RIF 2.79 Å, Cryo-EM structure of the MCM double hexamer loaded onto dsDNA.
7V3V 2.9 Å, Cryo-EM structure of MCM double hexamer bound with DDK in State I
9TH8 2.9 Å, Cryo-EM structure of MCM2-7 DH bound to Sld3-Sld7, Cdc45 and DNA
7P30 3.0 Å, 3.0 A resolution structure of a DNA-loaded MCM double hexamer
8KG6 3.07 Å, Yeast replisome in state I
9RHL 3.1 Å, Phospho-DH bound by Sld3-MBD on ARS1 DNA
7PMK 3.2 Å, S. cerevisiae replisome-SCF(Dia2) complex bound to double-stranded DNA (conformation I)
7PMN 3.2 Å, S. cerevisiae replisome-SCF(Dia2) complex bound to double-stranded DNA (conformation II)
7PT6 3.2 Å, Structure of MCM2-7 DH complexed with Cdc7-Dbf4 in the presence of ATPgS, state III
7V3U 3.2 Å, Cryo-EM structure of MCM double hexamer with structured Mcm4-NSD
9E2Y 3.2 Å, Cryo-EM structure of yeast CMG helicase stalled at G4-containing DNA template, state 3

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