9E2Z: DNA replication licensing factor MCM2

Cryo-EM structure of human CMG helicase stalled at G4-containing DNA template. Determined by electron microscopy at 2.6 Å resolution. Released 26 Mar 2025.

Method: Electron microscopy
Resolution: 2.6 Å
Organisms: Homo sapiens, synthetic construct
Chains: 13
Atoms: 41,017
Mol. weight: 738.68 kDa
Ligands: ZN, ATP, MG, ADP
Released: 26 Mar 2025

Explore 9E2Z in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9E2Z contains 269 α-helices and 223 β-strands across 11 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: 37 helices, 29 β-strands

Element	Residues	Length	Sheet
α-helix	182-187	6
α-helix	189-205	17
β-strand	207	1	1
β-strand	213	1	1
α-helix	214-224	11
β-strand	229-233	5	2
α-helix	234-240	7
α-helix	242-250	9
α-helix	252-270	19
α-helix	274-276	3
β-strand	282-286	5	2
β-strand	291	1	3
α-helix	299-301	3
β-strand	305-315	11	3
α-helix	316-318	3
β-strand	319-325	7	3
β-strand	327	1	4
α-helix	336-338	3
β-strand	339-340	2	3
β-strand	363	1	4
β-strand	369-379	11	3
α-helix	386	1
α-helix	390-392	3
β-strand	393-399	7	3
α-helix	400-402	3
β-strand	411-421	11	3
α-helix	424-430	7
β-strand	434	1	5
α-helix	435	1
β-strand	436-445	10	3
α-helix	458-469	12
α-helix	473-479	7
α-helix	489-500	12
β-strand	505	1	6
β-strand	513	1	6
β-strand	519-523	5	7
α-helix	529-539	11
β-strand	543-547	5	7
α-helix	548-550	3
β-strand	551	1	8
α-helix	553-556	4
β-strand	558-562	5	9
β-strand	569-573	5	9
α-helix	575-578	4
β-strand	583-587	5	7
α-helix	589-591	3
α-helix	594-597	4
α-helix	598-600	3
α-helix	601-605	5
β-strand	608-613	6	10
β-strand	616-621	6	10
β-strand	625-630	6	7
α-helix	633-635	3
α-helix	643-646	4
α-helix	651-655	5
β-strand	659-663	5	7
α-helix	669-685	17
α-helix	713-716	4
α-helix	717-730	14
β-strand	734	1	11
α-helix	740-757	18
α-helix	765-780	16
β-strand	785	1	11
α-helix	788-804	17
α-helix	808-817	10
α-helix	819-822	4

Chain 3: 32 helices, 35 β-strands

Element	Residues	Length	Sheet
α-helix	10-23	14
α-helix	31-41	11
β-strand	46-50	5	12
α-helix	51-57	7
α-helix	59-67	9
α-helix	69-87	19
α-helix	89-94	6
β-strand	99-103	5	12
α-helix	107-109	3
β-strand	110-111	2	13
α-helix	118-120	3
β-strand	124-134	11	13
β-strand	138-148	11	13
β-strand	153-158	6	13
α-helix	173-175	3
β-strand	177	1	14
β-strand	183	1	14
β-strand	185-186	2	13
β-strand	192-202	11	13
α-helix	205-207	3
α-helix	209	1
α-helix	213-215	3
β-strand	216-222	7	13
α-helix	223-225	3
β-strand	234-244	11	13
β-strand	248	1	15
β-strand	251-252	2	15
β-strand	255-256	2	16
β-strand	257-267	11	13
α-helix	278-290	13
α-helix	294-301	8
α-helix	308-309	2
α-helix	311-322	12
β-strand	327-328	2	17
β-strand	334-335	2	17
β-strand	341-345	5	18
α-helix	351-361	11
β-strand	365-369	5	18
β-strand	373	1	19
α-helix	375-379	5
β-strand	380-384	5	20
β-strand	391-395	5	20
α-helix	397-400	4
β-strand	405-409	5	18
α-helix	411-413	3
α-helix	416-427	12
β-strand	430-435	6	21
β-strand	438-443	6	21
β-strand	447-452	6	18
α-helix	465-469	5
α-helix	473-477	5
β-strand	481-485	5	18
α-helix	491-506	16
α-helix	513	1
β-strand	515	1	22
β-strand	545	1	23
α-helix	546-548	3
α-helix	550-553	4
β-strand	563	1	23
α-helix	565-576	12
β-strand	581-582	2	24
α-helix	584-599	16
β-strand	607-609	3	25
α-helix	615-631	17
β-strand	636-637	2	24
α-helix	639-652	14
β-strand	654-656	3	25

Chain 4: 31 helices, 26 β-strands

Element	Residues	Length	Sheet
α-helix	159-172	14
α-helix	192-203	12
β-strand	207-211	5	26
α-helix	212-218	7
α-helix	220-228	9
α-helix	230-248	19
β-strand	259-263	5	26
α-helix	271-273	3
α-helix	276-278	3
β-strand	282-292	11	27
α-helix	293-295	3
β-strand	296-306	11	27
β-strand	312-316	5	27
β-strand	318	1	28
β-strand	321	1	28
β-strand	327	1	29
β-strand	334	1	29
β-strand	337-339	3	27
α-helix	341-343	3
β-strand	345-356	12	27
α-helix	357	1
α-helix	362	1
α-helix	365-368	4
β-strand	369-375	7	27
α-helix	376-378	3
β-strand	387-397	11	27
β-strand	400	1	30
β-strand	408-409	2	30
β-strand	411	1	31
α-helix	412	1
β-strand	413-423	11	27
α-helix	443-454	12
α-helix	458-466	9
α-helix	474-485	12
β-strand	506-510	5	32
α-helix	516-526	11
β-strand	531-533	3	32
α-helix	562-565	4
β-strand	569-573	5	32
α-helix	576-578	3
α-helix	581-593	13
β-strand	595-600	6	33
β-strand	603-608	6	33
β-strand	611-617	7	32
α-helix	630-633	4
α-helix	638-641	4
β-strand	646-650	5	32
α-helix	651-653	3
α-helix	656-670	15
α-helix	674-697	24
β-strand	701-702	2	34
α-helix	704-723	20
α-helix	731-747	17
β-strand	752-753	2	34
α-helix	755-772	18
α-helix	784-788	5
α-helix	792-806	15

Chain 5: 36 helices, 33 β-strands

Element	Residues	Length	Sheet
α-helix	23-41	19
β-strand	44-46	3	35
β-strand	50-52	3	35
α-helix	54-65	12
β-strand	69-73	5	36
α-helix	74-80	7
α-helix	82-90	9
α-helix	92-111	20
α-helix	121-123	3
β-strand	125-129	5	36
α-helix	133-134	2
α-helix	142-144	3
β-strand	148-158	11	16
β-strand	162-172	11	16
β-strand	178-183	6	16
β-strand	189	1	37
β-strand	191-192	2	15
α-helix	193-194	2
α-helix	205-208	4
β-strand	212-214	3	16
α-helix	216-218	3
β-strand	220-230	11	16
α-helix	231-232	2
α-helix	233-235	3
α-helix	237	1
α-helix	241-243	3
β-strand	244-250	7	16
α-helix	251-253	3
β-strand	262-271	10	16
β-strand	273	1	37
β-strand	289	1	3
β-strand	292-301	10	16
α-helix	316-326	11
α-helix	331-337	7
α-helix	347-358	12
β-strand	363-364	2	38
β-strand	370-371	2	38
β-strand	377-381	5	39
α-helix	387-397	11
β-strand	401-405	5	39
β-strand	409	1	10
α-helix	411-414	4
β-strand	416-420	5	40
β-strand	427-431	5	40
α-helix	433-436	4
β-strand	441-445	5	39
α-helix	447-449	3
α-helix	452-456	5
α-helix	459-463	5
β-strand	466-471	6	19
β-strand	474-479	6	19
β-strand	483-489	7	39
β-strand	490	1	41
β-strand	493	1	41
α-helix	500-503	4
α-helix	508-511	4
β-strand	516-520	5	39
α-helix	526-541	16
α-helix	542-544	3
α-helix	546-547	2
α-helix	556-569	14
β-strand	571	1	22
β-strand	573-574	2	42
α-helix	576-600	25
α-helix	610-626	17
β-strand	631-632	2	42
α-helix	634-644	11
α-helix	645-649	5
α-helix	650-652	3

Chain 6: 34 helices, 29 β-strands

Element	Residues	Length	Sheet
α-helix	21-36	16
β-strand	38	1	43
β-strand	44	1	43
α-helix	45-50	6
α-helix	51-54	4
β-strand	60-63	4	44
α-helix	65-71	7
α-helix	73-81	9
α-helix	83-98	16
β-strand	111-114	4	44
α-helix	123-125	3
α-helix	128-130	3
β-strand	131	1	5
β-strand	134-144	11	5
α-helix	145-147	3
β-strand	148-158	11	5
α-helix	163	1
β-strand	164-169	6	5
β-strand	191-193	3	5
α-helix	195-197	3
β-strand	199-209	11	5
α-helix	216	1
α-helix	220-222	3
β-strand	223-229	7	5
α-helix	230-232	3
β-strand	241-252	12	5
α-helix	254-256	3
β-strand	281	1	45
β-strand	282	1	28
α-helix	285-287	3
β-strand	292	1	45
β-strand	294-305	12	5
α-helix	322-328	7
α-helix	331-342	12
α-helix	346-354	9
α-helix	362-373	12
β-strand	378-379	2	46
β-strand	385-386	2	46
β-strand	392-396	5	47
α-helix	402-412	11
β-strand	416-420	5	47
β-strand	432-435	4	48
β-strand	442-445	4	48
α-helix	448-451	4
β-strand	456-460	5	47
α-helix	462-464	3
α-helix	467-478	12
β-strand	481-486	6	8
β-strand	489-494	6	8
β-strand	498-503	6	47
α-helix	516-519	4
α-helix	524-528	5
β-strand	532-536	5	47
α-helix	542-557	16
α-helix	559-562	4
α-helix	569-579	11
α-helix	584	1
β-strand	585-586	2	49
α-helix	587	1
α-helix	588-604	17
α-helix	618-634	17
β-strand	639-640	2	49
α-helix	642-660	19

Chain 7: 29 helices, 27 β-strands

Element	Residues	Length	Sheet
α-helix	6-19	14
β-strand	22	1	50
β-strand	30	1	50
α-helix	32-42	11
β-strand	47-51	5	51
α-helix	52-58	7
α-helix	60-68	9
α-helix	70-88	19
α-helix	98-104	7
β-strand	135-139	5	51
β-strand	147	1	31
α-helix	149-151	3
α-helix	154-156	3
β-strand	157	1	31
β-strand	160-170	11	31
β-strand	174-184	11	31
β-strand	190-194	5	31
β-strand	199-200	2	30
β-strand	205	1	52
α-helix	209-213	5
β-strand	219	1	52
β-strand	220-222	3	31
β-strand	228-238	11	31
α-helix	241-243	3
α-helix	245	1
β-strand	252-258	7	31
α-helix	259-261	3
β-strand	270-279	10	31
α-helix	280-282	3
α-helix	285-289	5
β-strand	293	1	13
β-strand	296-305	10	31
α-helix	323-329	7
α-helix	332-338	7
α-helix	348-359	12
β-strand	364	1	53
β-strand	370	1	53
β-strand	377-381	5	54
α-helix	387-397	11
β-strand	401-405	5	54
α-helix	433-436	4
β-strand	441-445	5	54
α-helix	447-449	3
α-helix	452-464	13
β-strand	466-471	6	55
β-strand	474-479	6	55
β-strand	483-488	6	54
α-helix	509-512	4
β-strand	517-521	5	54
α-helix	527-543	17
α-helix	546-548	3
α-helix	552-555	4
α-helix	556-567	12
α-helix	577-595	19
α-helix	603-619	17
α-helix	627-640	14

Chain A: 8 helices, 5 β-strands

Element	Residues	Length	Sheet
α-helix	4-14	11
α-helix	22-24	3
α-helix	26-53	28
α-helix	56-58	3
α-helix	59-93	35
α-helix	100-105	6
α-helix	108-127	20
β-strand	147-152	6	56
β-strand	157-160	4	57
β-strand	166-169	4	57
β-strand	174-178	5	56
α-helix	179-187	9
β-strand	191-194	4	56

Chain B: 9 helices, 8 β-strands

Element	Residues	Length	Sheet
α-helix	3-10	8
β-strand	14-19	6	58
β-strand	23	1	59
β-strand	26-27	2	60
β-strand	32-33	2	60
α-helix	34-35	2
β-strand	36	1	59
β-strand	42-45	4	58
α-helix	46-54	9
β-strand	58-61	4	58
α-helix	68-80	13
α-helix	84-87	4
α-helix	92-103	12
α-helix	104-106	3
α-helix	110-138	29
β-strand	143-144	2	61
α-helix	150-171	22

3 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	904	Homo sapiens	P49736 (AlphaFold model)
Isoform 2 of DNA replication licensing factor MCM3	3	protein	853	Homo sapiens	P25205 (AlphaFold model)
DNA replication licensing factor MCM4	4	protein	883	Homo sapiens	P33991 (AlphaFold model)
DNA replication licensing factor MCM5	5	protein	734	Homo sapiens	P33992 (AlphaFold model)
DNA replication licensing factor MCM6	6	protein	821	Homo sapiens	Q14566
DNA replication licensing factor MCM7	7	protein	719	Homo sapiens	P33993
DNA replication complex GINS protein PSF1	A	protein	196	Homo sapiens	Q14691
DNA replication complex GINS protein PSF2	B	protein	222	Homo sapiens	Q9Y248
DNA replication complex GINS protein PSF3	C	protein	216	Homo sapiens	Q9BRX5
DNA replication complex GINS protein SLD5	D	protein	223	Homo sapiens	Q9BRT9
Cell division control protein 45 homolog	E	protein	566	Homo sapiens	O75419
Leading strand DNA template	F	DNA	40	synthetic construct

1 more molecules are not listed.

Sequence of entity 1 (2), FASTA

>9E2Z_1 DNA replication licensing factor MCM2 (chains 2)
MAESSESFTMASSPAQRRRGNDPLTSSPGRSSRRTDALTSSPGRDLPPFEDESEGLLGTE
GPLEEEEDGEELIGDGMERDYRAIPELDAYEAEGLALDDEDVEELTASQREAAERAMRQR
DREAGRGLGRMRRGLLYDSDEEDEERPARKRRQVERATEDGEEDEEMIESIENLEDLKGH
SVREWVSMAGPRLEIHHRFKNFLRTHVDSHGHNVFKERISDMCKENRESLVVNYEDLAAR
EHVLAYFLPEAPAELLQIFDEAALEVVLAMYPKYDRITNHIHVRISHLPLVEELRSLRQL
HLNQLIRTSGVVTSCTGVLPQLSMVKYNCNKCNFVLGPFCQSQNQEVKPGSCPECQSAGP
FEVNMEETIYQNYQRIRIQESPGKVAAGRLPRSKDAILLADLVDSCKPGDEIELTGIYHN
NYDGSLNTANGFPVFATVILANHVAKKDNKVAVGELTDEDVKMITSLSKDQQIGEKIFAS
IAPSIYGHEDIKRGLALALFGGEPKNPGGKHKVRGDINVLLCGDPGTAKSQFLKYIEKVS
SRAIFTTGQGASAVGLTAYVQRHPVSREWTLEAGALVLADRGVCLIDEFDKMNDQDRTSI
HEAMEQQSISISKAGIVTSLQARCTVIAAANPIGGRYDPSLTFSENVDLTEPIISRFDIL
CVVRDTVDPVQDEMLARFVVGSHVRHHPSNKEEEGLANGSAAEPAMPNTYGVEPLPQEVL
KKYIIYAKERVHPKLNQMDQDKVAKMYSDLRKESMATGSIPITVRHIESMIRMAEAHARI
HLRDYVIEDDVNMAIRVMLESFIDTQKFSVMRSMRKTFARYLSFRRDNNELLLFILKQLV
AEQVTYQRNRFGAQQDTIEVPEKDLVDKARQINIHNLSAFYDSELFRMNKFSHDLKRKMI
LQQF

Sequence of entity 2 (3), FASTA

>9E2Z_2 Isoform 2 of DNA replication licensing factor MCM3 (chains 3)
MLPRSPPLPRGNLWWREEFGSFRAGVESSWEPPRDFGGGSSLAAGMAGTVVLDDVELREA
QRDYLDFLDDEEDQGIYQSKVRELISDNQYRLIVNVNDLRRKNEKRANRLLNNAFEELVA
FQRALKDFVASIDATYAKQYEEFYVGLEGSFGSKHVSPRTLTSCFLSCVVCVEGIVTKCS
LVRPKVVRSVHYCPATKKTIERRYSDLTTLVAFPSSSVYPTKDEENNPLETEYGLSVYKD
HQTITIQEMPEKAPAGQLPRSVDVILDDDLVDKAKPGDRVQVVGTYRCLPGKKGGYTSGT
FRTVLIACNVKQMSKDAQPSFSAEDIAKIKKFSKTRSKDIFDQLAKSLAPSIHGHDYVKK
AILCLLLGGVERDLENGSHIRGDINILLIGDPSVAKSQLLRYVLCTAPRAIPTTGRGSSG
VGLTAAVTTDQETGERRLEAGAMVLADRGVVCIDEFDKMSDMDRTAIHEVMEQGRVTIAK
AGIHARLNARCSVLAAANPVYGRYDQYKTPMENIGLQDSLLSRFDLLFIMLDQMDPEQDR
EISDHVLRMHRYRAPGEQDGDAMPLGSAVDILATDDPNFSQEDQQDTQIYEKHDNLLHGT
KKKKEKMVSAAFMKKYIHVAKIIKPVLTQESATYIAEEYSRLRSQDSMSSDTARTSPVTA
RTLETLIRLATAHAKARMSKTVDLQDAEEAVELVQYAYFKKVLEKEKKRKKRSEDESETE
DEEEKSQEDQEQKRKRRKTRQPDAKDGDSYDPYDFSDTEEEMPQVHTPKTADSQETKESQ
KVELSESRLKAFKVALLDVFREAHAQSIGMNRLTESINRDSEEPFSSVEIQAALSKMQDD
NQVMVSEGIIFLI

Sequence of entity 3 (4), FASTA

>9E2Z_3 DNA replication licensing factor MCM4 (chains 4)
MHHHHHHHHENLYFQGSSATMSSPASTPSRRGSRRGRATPAQTPRSEDARSSPSQRRRGE
DSTSTGELQPMPTSPGVDLQSPAAQDVLFSSPPQMHSSAIPLDFDVSSPLTYGTPSSRVE
GTPRSGVRGTPVRQRPDLGSAQKGLQVDLQSDGAAAEDIVASEQSLGQKLVIWGTDVNVA
ACKENFQRFLQRFIDPLAKEEENVGIDITEPLYMQRLGEINVIGEPFLNVNCEHIKSFDK
NLYRQLISYPQEVIPTFDMAVNEIFFDRYPDSILEHQIQVRPFNALKTKNMRNLNPEDID
QLITISGMVIRTSQLIPEMQEAFFQCQVCAHTTRVEMDRGRIAEPSVCGRCHTTHSMALI
HNRSLFSDKQMIKLQESPEDMPAGQTPHTVILFAHNDLVDKVQPGDRVNVTGIYRAVPIR
VNPRVSNVKSVYKTHIDVIHYRKTDAKRLHGLDEEAEQKLFSEKRVELLKELSRKPDIYE
RLASALAPSIYEHEDIKKGILLQLFGGTRKDFSHTGRGKFRAEINILLCGDPGTSKSQLL
QYVYNLVPRGQYTSGKGSSAVGLTAYVMKDPETRQLVLQTGALVLSDNGICCIDEFDKMN
ESTRSVLHEVMEQQTLSIAKAGIICQLNARTSVLAAANPIESQWNPKKTTIENIQLPHTL
LSRFDLIFLLLDPQDEAYDRRLAHHLVALYYQSEEQAEEELLDMAVLKDYIAYAHSTIMP
RLSEEASQALIEAYVDMRKIGSSRGMVSAYPRQLESLIRLAEAHAKVRLSNKVEAIDVEE
AKRLHREALKQSATDPRTGIVDISILTTGMSATSRKRKEELAEALKKLILSKGKTPALKY
QQLFEDIRGQSDIAITKDMFEEALRALADDDFLTVTGKTVRLL

Sequence of entity 4 (5), FASTA

>9E2Z_4 DNA replication licensing factor MCM5 (chains 5)
MSGFDDPGIFYSDSFGGDAQADEGQARKSQLQRRFKEFLRQYRVGTDRTGFTFKYRDELK
RHYNLGEYWIEVEMEDLASFDEDLADYLYKQPAEHLQLLEEAAKEVADEVTRPRPSGEEV
LQDIQVMLKSDASPSSIRSLKSDMMSHLVKIPGIIIAASAVRAKATRISIQCRSCRNTLT
NIAMRPGLEGYALPRKCNTDQAGRPKCPLDPYFIMPDKCKCVDFQTLKLQELPDAVPHGE
MPRHMQLYCDRYLCDKVVPGNRVTIMGIYSIKKFGLTTSRGRDRVGVGIRSSYIRVLGIQ
VDTDGSGRSFAGAVSPQEEEEFRRLAALPNVYEVISKSIAPSIFGGTDMKKAIACLLFGG
SRKRLPDGLTRRGDINLLMLGDPGTAKSQLLKFVEKCSPIGVYTSGKGSSAAGLTASVMR
DPSSRNFIMEGGAMVLADGGVVCIDEFDKMREDDRVAIHEAMEQQTISIAKAGITTTLNS
RCSVLAAANSVFGRWDETKGEDNIDFMPTILSRFDMIFIVKDEHNEERDVMLAKHVITLH
VSALTQTQAVEGEIDLAKLKKFIAYCRVKCGPRLSAEAAEKLKNRYIIMRSGARQHERDS
DRRSSIPITVRQLEAIVRIAEALSKMKLQPFATEADVEEALRLFQVSTLDAALSGTLSGV
EGFTSQEDQEMLSRIEKQLKRRFAIGSQVSEHSIIKDFTKQKYPEHAIHKVLQLMLRRGE
IQHRMQRKVLYRLK

Sequence of entity 5 (6), FASTA

>9E2Z_5 DNA replication licensing factor MCM6 (chains 6)
MDLAAAAEPGAGSQHLEVRDEVAEKCQKLFLDFLEEFQSSDGEIKYLQLAEELIRPERNT
LVVSFVDLEQFNQQLSTTIQEEFYRVYPYLCRALKTFVKDRKEIPLAKDFYVAFQDLPTR
HKIRELTSSRIGLLTRISGQVVRTHPVHPELVSGTFLCLDCQTVIRDVEQQFKYTQPNIC
RNPVCANRRRFLLDTNKSRFVDFQKVRIQETQAELPRGSIPRSLEVILRAEAVESAQAGD
KCDFTGTLIVVPDVSKLSTPGARAETNSRVSGVDGYETEGIRGLRALGVRDLSYRLVFLA
CCVAPTNPRFGGKELRDEEQTAESIKNQMTVKEWEKVFEMSQDKNLYHNLCTSLFPTIHG
NDEVKRGVLLMLFGGVPKTTGEGTSLRGDINVCIVGDPSTAKSQFLKHVEEFSPRAVYTS
GKASSAAGLTAAVVRDEESHEFVIEAGALMLADNGVCCIDEFDKMDVRDQVAIHEAMEQQ
TISITKAGVKATLNARTSILAAANPISGHYDRSKSLKQNINLSAPIMSRFDLFFILVDEC
NEVTDYAIARRIVDLHSRIEESIDRVYSLDDIRRYLLFARQFKPKISKESEDFIVEQYKH
LRQRDGSGVTKSSWRITVRQLESMIRLSEAMARMHCCDEVQPKHVKEAFRLLNKSIIRVE
TPDVNLDQEEEIQMEVDEGAGGINGHADSPAPVNGINGYNEDINQESAPKASLRLGFSEY
CRISNLIVLHLRKVEEEEDESALKRSELVNWYLKEIESEIDSEEELINKKRIIEKVIHRL
THYDHVLIELTQAGLKGSTEGSESYEEDPYLVVNPNYLLED

Sequence of entity 6 (7), FASTA

>9E2Z_6 DNA replication licensing factor MCM7 (chains 7)
MALKDYALEKEKVKKFLQEFYQDDELGKKQFKYGNQLVRLAHREQVALYVDLDDVAEDDP
ELVDSICENARRYAKLFADAVQELLPQYKEREVVNKDVLDVYIEHRLMMEQRSRDPGMVR
SPQNQYPAELMRRFELYFQGPSSNKPRVIREVRADSVGKLVTVRGIVTRVSEVKPKMVVA
TYTCDQCGAETYQPIQSPTFMPLIMCPSQECQTNRSGGRLYLQTRGSRFIKFQEMKMQEH
SDQVPVGNIPRSITVLVEGENTRIAQPGDHVSVTGIFLPILRTGFRQVVQGLLSETYLEA
HRIVKMNKSEDDESGAGELTREELRQIAEEDFYEKLAASIAPEIYGHEDVKKALLLLLVG
GVDQSPRGMKIRGNINICLMGDPGVAKSQLLSYIDRLAPRSQYTTGRGSSGVGLTAAVLR
DSVSGELTLEGGALVLADQGVCCIDEFDKMAEADRTAIHEVMEQQTISIAKAGILTTLNA
RCSILAAANPAYGRYNPRRSLEQNIQLPAALLSRFDLLWLIQDRPDRDNDLRLAQHITYV
HQHSRQPPSQFEPLDMKLMRRYIAMCREKQPMVPESLADYITAAYVEMRREAWASKDATY
TSARTLLAILRLSTALARLRMVDVVEKEDVNEAIRLMEMSKDSLLGDKGQTARTQRPADV
IFATVRELVSGGRSVRFSEAEQRCVSRGFTPAQFQAALDEYEELNVWQVNASRTRITFV

Sequence of entity 7 (A), FASTA

>9E2Z_7 DNA replication complex GINS protein PSF1 (chains A)
MFCEKAMELIRELHRAPEGQLPAFNEDGLRQVLEEMKALYEQNQSDVNEAKSGGRSDLIP
TIKFRHCSLLRNRRCTVAYLYDRLLRIRALRWEYGSVLPNALRFHMAAEEMEWFNNYKRS
LATYMRSLGGDEGLDITQDMKPPKSLYIEVRCLKDYGEFEVDDGTSVLLKKNSQHFLPRW
KCEQLIRQGVLEHILS

Sequence of entity 8 (B), FASTA

>9E2Z_8 DNA replication complex GINS protein PSF2 (chains B)
MDAAEVEFLAEKELVTIIPNFSLDKIYLIGGDLGPFNPGLPVEVPLWLAINLKQRQKCRL
LPPEWMDVEKLEKMRDHERKEETFTPMPSPYYMELTKLLLNHASDNIPKADEIRTLVKDM
WDTRIAKLRVSADSFVRQQEAHAKLDNLTLMEINTSGTFLTQALNHMYKLRTNLQPLEST
QSQDFENLYFQGSAWSHPQFEKGGGSGGGSGGSAWSHPQFEK

Sequence of entity 9 (C), FASTA

>9E2Z_9 DNA replication complex GINS protein PSF3 (chains C)
MSEAYFRVESGALGPEENFLSLDDILMSHEKLPVRTETAMPRLGAFFLERSAGAETDNAV
PQGSKLELPLWLAKGLFDNKRRILSVELPKIYQEGWRTVFSADPNVVDLHKMGPHFYGFG
SQLLHFDSPENADISQSLLQTFIGRFRRIMDSSQNAYNEDTSALVARLDEMERGLFQTGQ
KGLNDFQCWEKGQASQITASNLVQNYKKRKFTDMED

Sequence of entity 10 (D), FASTA

>9E2Z_10 DNA replication complex GINS protein SLD5 (chains D)
MTEEVDFLGQDSDGGSEEVVLTPAELIERLEQAWMNEKFAPELLESKPEIVECVMEQLEH
MEENLRRAKREDLKVSIHQMEMERIRYVLSSYLRCRLMKIEKFFPHVLEKEKTRPEGEPS
SLSPEELAFAREFMANTESYLKNVALKHMPPNLQKVDLFRAVPKPDLDSYVFLRVRERQE
NILVEPDTDEQRDYVIDLEKGSQHLIRYKTIAPLVASGAVQLI

Sequence of entity 11 (E), FASTA

>9E2Z_11 Cell division control protein 45 homolog (chains E)
MFVSDFRKEFYEVVQSQRVLLFVASDVDALCACKILQALFQCDHVQYTLVPVSGWQELET
AFLEHKEQFHYFILINCGANVDLLDILQPDEDTIFFVCDTHRPVNVVNVYNDTQIKLLIK
QDDDLEVPAYEDIFRDEEEDEEHSGNDSDGSEPSEKRTRLEEEIVEQTMRRRQRREWEAR
RRDILFDYEQYEYHGTSSAMVMFELAWMLSKDLNDMLWWAIVGLTDQWVQDKITQMKYVT
DVGVLQRHVSRHNHRNEDEENTLSVDCTRISFEYDLRLVLYQHWSLHDSLCNTSYTAARF
KLWSVHGQKRLQEFLADMGLPLKQVKQKFQAMDISLKENLREMIEESANKFGMKDMRVQT
FSIHFGFKHKFLASDVVFATMSLMESPEKDGSGTDHFIQALDSLSRSNLDKLYHGLELAK
KQLRATQQTIASCLCTNLVISQGPFLYCSLMEGTPDVMLFSRPASLSLLSKHLLKSFVCS
TKNRRCKLLPLVMAAPLSMEHGTVTVVGIPPETDSSDRKNFFGRAFEKAAESTSSRMLHN
HFDLSVIELKAEDRSKFLDALISLLS

Sequence of entity 12 (F), FASTA

>9E2Z_12 Leading strand DNA template (chains F)
GTGATATCTGCTTTGGGTGGGTGGGTGGGTTGAGGCAATA

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	4
ATP	Adenosine-5'-triphosphate	C10 H16 N5 O13 P3	5
MG	Magnesium ion	Mg	6
ADP	Adenosine-5'-diphosphate	C10 H15 N5 O10 P2	1

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

Primary citation

G-quadruplex-stalled eukaryotic replisome structure reveals helical inchworm DNA translocation. Batra, S., Allwein, B., Kumar, C. et al. Science (2025) 387:eadt1978-eadt1978. DOI 10.1126/science.adt1978 · PubMed

Other PDB entries of the same protein (UniProt P49736 (AlphaFold model), which also has an AlphaFold model), best resolution first:

6YA7 1.67 Å, Cdc7-Dbf4 bound to an Mcm2-S40 derived bivalent substrate
7CIZ 1.8 Å, Crystal structure of DNAJC9 HBD helix2 in complex with H3.3-H4 dimer and MCM2 HBD
5BNX 2.31 Å, Crystal structure of Human MCM2 HBD and ASF1b chaperoning a histone H3.3-H4 dimer
5JA4 2.42 Å, Crystal structure of human TONSL and MCM2 HBDs binding to a histone H3-H4 tetramer
7CJ0 2.5 Å, Crystal structure of DNAJC9 HBD in complex with H3.3-H4 dimer and MCM2 HBD
7W1Y 2.59 Å, Human MCM double hexamer bound to natural DNA duplex (polyAT/polyTA)
5BNV 2.79 Å, Crystal structure of Human MCM2 HBD chaperoning a histone H3-H4 tetramer
7PLO 2.8 Å, H. sapiens replisome-CUL2/LRR1 complex
8W0F 2.8 Å, Cryo-EM structure of a human MCM2-7 double hexamer on dsDNA
4UUZ 2.9 Å, MCM2-histone complex
5BO0 2.91 Å, Crystal structure of Human MCM2 HBD and ASF1b chaperoning a histone H3.2-H4 dimer
8S09 3.1 Å, H. sapiens MCM2-7 double hexamer bound to double stranded DNA

Browse structure collections

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