9MU9: Histone H2A

Name: PDB 9MU9: Structure of a native Drosophila melanogaster Nucleosome Elongation Complex (Pol II EC-nucleosome). Composite map
Published: 2025-02-19
License: https://creativecommons.org/publicdomain/zero/1.0/
Keywords: transcription

Structure of a native Drosophila melanogaster Nucleosome Elongation Complex (Pol II EC-nucleosome). Composite map. Determined by electron microscopy at 7.8 Å resolution. Released 19 Feb 2025.

Method: Electron microscopy
Resolution: 7.8 Å
Organism: Drosophila melanogaster
Chains: 21
Atoms: 42,423
Mol. weight: 615.73 kDa
Ligands: ZN
Released: 19 Feb 2025

Explore 9MU9 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9MU9 contains 200 α-helices and 215 β-strands across 18 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 a and e: 4 helices, 2 β-strands

Element	Residues	Length	Sheet
α-helix	46-55	10
α-helix	65-79	15
β-strand	84-85	2	9
α-helix	87-114	28
β-strand	119	1	10
α-helix	122-132	11

Chain A: 73 helices, 67 β-strands

Element	Residues	Length	Sheet
β-strand	11	1	11
β-strand	17-18	2	12
β-strand	21	1	13
α-helix	24-30	7
β-strand	43-44	2	14
β-strand	47-48	2	14
α-helix	49	1
β-strand	82-90	9	15
α-helix	96-103	8
β-strand	106	1	16
β-strand	113	1	16
α-helix	120-128	9
α-helix	133-143	11
β-strand	149	1	17
α-helix	163-166	4
α-helix	169-171	3
β-strand	179	1	17
α-helix	180-181	2
β-strand	182-185	4	18
β-strand	190-193	4	18
β-strand	205-207	3	18
α-helix	210-218	9
α-helix	222-227	6
α-helix	237-240	4
β-strand	242-247	6	15
β-strand	259	1	19
β-strand	261	1	19
α-helix	267-287	21
α-helix	292-310	19
α-helix	318-320	3
β-strand	321	1	20
α-helix	322	1
β-strand	327	1	20
α-helix	331-335	5
α-helix	349-351	3
β-strand	354-361	8	21
β-strand	369-372	4	21
α-helix	374-379	6
β-strand	381-385	5	22
α-helix	386	1
α-helix	388-390	3
α-helix	391-400	10
β-strand	408-412	5	22
β-strand	418-420	3	22
α-helix	427-429	3
β-strand	437-441	5	22
β-strand	447-451	5	21
α-helix	458-460	3
β-strand	461-468	8	21
β-strand	474-476	3	21
α-helix	478-480	3
α-helix	481-484	4
β-strand	492-496	5	21
α-helix	501-506	6
α-helix	507-511	5
β-strand	512	1	21
α-helix	514-516	3
β-strand	518	1	23
β-strand	525	1	23
α-helix	531-540	10
β-strand	546-548	3	24
α-helix	549-556	8
α-helix	566-569	4
β-strand	571-573	3	24
β-strand	577-579	3	24
α-helix	580-587	8
β-strand	594-597	4	25
β-strand	619-623	5	25
β-strand	626-629	4	25
α-helix	644-652	9
α-helix	654-675	22
α-helix	681-684	4
α-helix	688-713	26
α-helix	718-720	3
α-helix	725-751	27
α-helix	757-764	8
α-helix	770-777	8
β-strand	781-782	2	26
β-strand	785	1	27
β-strand	788	1	27
α-helix	789-791	3
α-helix	809-812	4
β-strand	814-815	2	26
α-helix	825-844	20
α-helix	846-859	14
β-strand	864-865	2	28
β-strand	871-872	2	28
β-strand	878-880	3	28
α-helix	883-885	3
β-strand	888	1	29
β-strand	893-897	5	30
α-helix	905-912	8
α-helix	919-925	7
β-strand	926	1	31
α-helix	928-936	9
α-helix	940-961	22
β-strand	968-972	5	30
α-helix	975-985	11
β-strand	994	1	31
α-helix	998-1011	14
α-helix	1020-1029	10
α-helix	1031-1040	10
α-helix	1045-1049	5
α-helix	1054-1070	17
β-strand	1072	1	29
α-helix	1073-1074	2
β-strand	1078	1	28
α-helix	1080-1088	9
α-helix	1090-1092	3
α-helix	1113-1120	8
β-strand	1130-1134	5	32
β-strand	1135	1	33
α-helix	1137-1140	4
α-helix	1143-1153	11
β-strand	1156	1	34
β-strand	1162-1169	8	35
α-helix	1173-1175	3
α-helix	1179-1188	10
β-strand	1199-1208	10	35
α-helix	1210-1215	6
α-helix	1220-1231	12
β-strand	1235-1239	5	35
β-strand	1248-1254	7	35
α-helix	1273-1286	14
β-strand	1291	1	34
β-strand	1299-1302	4	32
β-strand	1311-1314	4	36
β-strand	1320-1323	4	36
β-strand	1327-1331	5	32
β-strand	1344	1	33
β-strand	1350-1351	2	32
α-helix	1354-1360	7
α-helix	1363-1380	18
α-helix	1387-1397	11
α-helix	1418-1422	5
α-helix	1427-1437	11
β-strand	1440-1441	2	12
α-helix	1446-1452	7
α-helix	1459-1461	3
β-strand	1463-1467	5	37

Chain b: 3 helices, 3 β-strands

Element	Residues	Length	Sheet
α-helix	32-41	10
β-strand	46	1	10
α-helix	51-76	26
β-strand	81-82	2	9
α-helix	84-93	10
β-strand	97-99	3	3

Chain B: 53 helices, 69 β-strands

Element	Residues	Length	Sheet
α-helix	14-16	3
α-helix	20-37	18
α-helix	42-49	8
α-helix	50-55	6
α-helix	56-62	7
α-helix	64-65	2
β-strand	66-67	2	38
β-strand	84-94	11	38
β-strand	98-100	3	39
β-strand	106-108	3	39
α-helix	111-116	6
β-strand	122-134	13	38
β-strand	141-154	14	38
α-helix	169-174	6
β-strand	186-188	3	40
β-strand	191-195	5	40
α-helix	196	1
β-strand	197-201	5	41
α-helix	202	1
β-strand	207-210	4	42
β-strand	216-224	9	42
β-strand	234-241	8	42
α-helix	249-253	5
β-strand	257-260	4	42
β-strand	268-269	2	42
α-helix	270-276	7
α-helix	282-289	8
α-helix	296-301	6
α-helix	303-308	6
α-helix	315-324	10
α-helix	334-346	13
α-helix	359-376	18
α-helix	382-384	3
α-helix	389-391	3
β-strand	392-395	4	41
α-helix	397-422	26
α-helix	432-435	4
α-helix	439-450	12
α-helix	460-462	3
β-strand	468-470	3	40
α-helix	476-482	7
β-strand	485-487	3	41
α-helix	502-503	2
β-strand	510	1	43
β-strand	525-526	2	41
β-strand	527	1	43
β-strand	532-533	2	44
α-helix	540-549	10
β-strand	552-553	2	45
α-helix	554-556	3
α-helix	559-561	3
β-strand	566-570	5	45
β-strand	573-578	6	45
α-helix	581-593	13
β-strand	603-607	5	45
β-strand	612-616	5	45
β-strand	622-628	7	44
β-strand	629-630	2	46
β-strand	633-634	2	46
α-helix	638-644	7
α-helix	654-659	6
β-strand	663-667	5	44
α-helix	670-672	3
β-strand	676-677	2	44
α-helix	680-683	4
β-strand	697-698	2	44
α-helix	702-705	4
α-helix	716-718	3
α-helix	721-731	11
β-strand	734	1	47
β-strand	749-753	5	48
β-strand	757	1	49
β-strand	761-762	2	50
β-strand	778-784	7	51
β-strand	796-799	4	51
α-helix	800-805	6
β-strand	810-819	10	48
β-strand	829-831	3	52
β-strand	839-840	2	52
α-helix	847-849	3
β-strand	855	1	53
α-helix	856-857	2
β-strand	861-862	2	48
β-strand	867-869	3	53
β-strand	871-874	4	52
α-helix	875-876	2
α-helix	884-886	3
β-strand	891-893	3	52
β-strand	896-897	2	53
β-strand	904-913	10	48
β-strand	919-929	11	48
β-strand	936-938	3	54
β-strand	945-946	2	54
β-strand	947-951	5	51
α-helix	953-955	3
β-strand	958-959	2	55
β-strand	967-969	3	51
α-helix	973-977	5
α-helix	980-995	16
α-helix	998	1
β-strand	999-1000	2	50
α-helix	1010-1018	9
β-strand	1027-1028	2	51
β-strand	1030-1031	2	55
α-helix	1037	1
β-strand	1038	1	55
α-helix	1039	1
β-strand	1043-1050	8	51
β-strand	1051-1052	2	54
β-strand	1053	1	47
β-strand	1062-1064	3	21
α-helix	1090-1099	10
α-helix	1102-1106	5
α-helix	1107-1111	5
β-strand	1115-1121	7	11
β-strand	1127	1	11
β-strand	1129-1131	3	56
β-strand	1136-1138	3	56
β-strand	1148-1154	7	11
α-helix	1155-1166	12
β-strand	1169	1	13
β-strand	1172	1	12

Chain c: 5 helices, 3 β-strands

Element	Residues	Length	Sheet
α-helix	17-20	4
α-helix	27-35	9
β-strand	42-43	2	4
α-helix	46-72	27
β-strand	77-78	2	5
α-helix	80-88	9
α-helix	91-96	6
β-strand	100-102	3	6

Chain C: 11 helices, 12 β-strands

Element	Residues	Length	Sheet
β-strand	8-14	7	57
β-strand	18-24	7	57
α-helix	28-39	12
β-strand	43-55	13	58
α-helix	61-69	9
β-strand	73	1	59
α-helix	82-84	3
β-strand	99-106	8	58
α-helix	112	1
β-strand	113-116	4	60
α-helix	117-119	3
β-strand	121-122	2	58
β-strand	129	1	59
α-helix	130-132	3
β-strand	150-154	5	60
β-strand	159-170	12	58
α-helix	175-177	3
β-strand	180-187	8	57
α-helix	201-203	3
α-helix	205-207	3
α-helix	219-221	3
β-strand	229-236	8	57
α-helix	242-267	26

Chains d and h: 4 helices, 2 β-strands

Element	Residues	Length	Sheet
α-helix	36-46	11
β-strand	51-52	2	5
α-helix	54-81	28
β-strand	86-87	2	4
α-helix	89-99	11
α-helix	103-121	19

Chain E: 9 helices, 13 β-strands

Element	Residues	Length	Sheet
α-helix	2-23	22
β-strand	26-27	2	67
α-helix	30-33	4
α-helix	37-43	7
β-strand	47	1	68
β-strand	52	1	68
β-strand	60-64	5	67
β-strand	71-75	5	67
β-strand	81-82	2	69
α-helix	84-95	12
β-strand	101-106	6	67
β-strand	109-110	2	69
α-helix	112-120	9
β-strand	126-131	6	67
α-helix	134-136	3
β-strand	147-151	5	70
α-helix	153-163	11
β-strand	171-173	3	70
α-helix	178-183	6
α-helix	185-186	2
β-strand	190-197	8	70
β-strand	201-209	9	70

8 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
RNA	R	RNA	10	Drosophila melanogaster
Template DNA	T	DNA	174	Drosophila melanogaster
Non-template DNA	N	DNA	163	Drosophila melanogaster
Histone H2A	c, g	protein	106	Drosophila melanogaster	P84051 (AlphaFold model)
Histone H2B	d, h	protein	97	Drosophila melanogaster	P02283 (AlphaFold model)
Histone H3	a, e	protein	100	Drosophila melanogaster	P02299 (AlphaFold model)
Histone H4	b, f	protein	80	Drosophila melanogaster	P84040 (AlphaFold model)
DNA-directed RNA polymerase II subunit RPB1	A	protein	1470	Drosophila melanogaster	P04052
DNA-directed RNA polymerase II subunit RPB2	B	protein	1164	Drosophila melanogaster	P08266
DNA-directed RNA polymerase II subunit RPB3	C	protein	269	Drosophila melanogaster	O97183
DNA-directed RNA polymerases I, II, and III subunit RPABC2	F	protein	84	Drosophila melanogaster	Q24320
DNA-directed RNA polymerases I, II, and III subunit RPABC3	H	protein	147	Drosophila melanogaster	Q9VNZ3

5 more molecules are not listed.

Sequence of entity 1 (R), FASTA

>9MU9_1 RNA (chains R)
AAAAAAAAAA

Sequence of entity 2 (T), FASTA

>9MU9_2 Template DNA (chains T)
ATCAGAATCCCGGTGCCGAGGCCGCTCAATTGGTCGTAGACAGCTCTAGCACCGCTTAAA
CGCACGTACGCGCTGTCCCCCGCGTTTTAACCGCCAAGGGGATTACTCCCTAGTCTCCAG
GCACGTGTTTTTTTTTTTTTTTTTTTTTTTTTTTTTCCTTTTTTTTTTTTTTTT

Sequence of entity 3 (N), FASTA

>9MU9_3 Non-template DNA (chains N)
AAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAACACGTGCCTGGAGACTAGGGAGTA
ATCCCCTTGGCGGTTAAAACGCGGGGGACAGCGCGTACGTGCGTTTAAGCGGTGCTAGAG
CTGTCTACGACCAATTGAGCGGCCTCGGCACCGGGATTCTGAT

Sequence of entity 4 (c, g), FASTA

>9MU9_4 Histone H2A (chains c, g)
AKSRSNRAGLQFPVGRIHRLLRKGNYAERVGAGAPVYLAAVMEYLAAEVLELAGNAARDN
KKTRIIPRHLQLAIRNDEELNKLLSGVTIAQGGVLPNIQAVLLPKK

Sequence of entity 5 (d, h), FASTA

>9MU9_5 Histone H2B (chains d, h)
KKRKRKESYAIYIYKVLKQVHPDTGISSKAMSIMNSFVNDIFERIAAEASRLAHYNKRST
ITSREIQTAVRLLLPGELAKHAVSEGTKAVTKYTSSK

Sequence of entity 6 (a, e), FASTA

>9MU9_6 Histone H3 (chains a, e)
KKPHRYRPGTVALREIRRYQKSTELLIRKLPFQRLVREIAQDFKTDLRFQSSAVMALQEA
SEAYLVGLFEDTNLCAIHAKRVTIMPKDIQLARRIRGERA

Sequence of entity 7 (b, f), FASTA

>9MU9_7 Histone H4 (chains b, f)
RDNIQGITKPAIRRLARRGGVKRISGLIYEETRGVLKVFLENVIRDAVTYTEHAKRKTVT
AMDVVYALKRQGRTLYGFGG

Sequence of entity 8 (A), FASTA

>9MU9_8 DNA-directed RNA polymerase II subunit RPB1 (chains A)
SKAPLRQVKRVQFGILSPDEIRRMSVTEGGVQFAETMEGGRPKLGGLMDPRQGVIDRTSR
CQTCAGNMTECPGHFGHIDLAKPVFHIGFITKTIKILRCVCFYCSKMLVSPHNPKIKEIV
MKSRGQPRKRLAYVYDLCKGKTICEGGEDMDLTKENQQPDPNKKPGHGGCGHYQPSIRRT
GLDLTAEWKHQNEDSQEKKIVVSAERVWEILKHITDEECFILGMDPKYARPDWMIVTVLP
VPPLAVRPAVVMFGAAKNQDDLTHKLSDIIKANNELRKNEASGAAAHVIQENIKMLQFHV
ATLVDNDMPGMPRAMQKSGKPLKAIKARLKGKEGRIRGNLMGKRVDFSARTVITPDPNLR
IDQVGVPRSIAQNLTFPELVTPFNIDRMQELVRRGNSQYPGAKYIVRDNGERIDLRFHPK
SSDLHLQCGYKVERHLRDDDLVIFNRQPTLHKMSMMGHRVKVLPWSTFRMNLSCTSPYNA
DFDGDEMNLHVPQSMETRAEVENIHITPRQIITPQANKPVMGIVQDTLTAVRKMTKRDVF
ITREQVMNLLMFLPTWDAKMPQPCILKPRPLWTGKQIFSLIIPGNVNMIRTHSTHPDEED
EGPYKWISPGDTKVMVEHGELIMGILCKKSLGTSAGSLLHICFLELGHDIAGRFYGNIQT
VINNWLLFEGHSIGIGDTIADPQTYNEIQQAIKKAKDDVINVIQKAHNMELEPTPGNTLR
QTFENKVNRILNDARDKTGGSAKKSLTEYNNLKAMVVSGSKGSNINISQVIACVGQQNVE
GKRIPYGFRKRTLPHFIKDDYGPESRGFVENSYLAGLTPSEFYFHAMGGREGLIDTAVKT
AETGYIQRRLIKAMESVMVNYDGTVRNSVGQLIQLRYGEDGLCGELVEFQNMPTVKLSNK
SFEKRFKFDWSNERLMKKVFTDDVIKEMTDSSEAIQELEAEWDRLVSDRDSLRQIFPNGE
SKVVLPCNLQRMIWNVQKIFHINKRLPTDLSPIRVIKGVKTLLERCVIVTGNDRISKQAN
ENATLLFQCLIRSTLCTKYVSEEFRLSTEAFEWLVGEIETRFQQAQANPGEMVGALAAQS
LGEPATQMTLNTFHFAGVSSKNVTLGVPRLKEIINISKKPKAPSLTVFLTGGAARDAEKA
KNVLCRLEHTTLRKVTANTAIYYDPDPQRTVISEDQEFVNVYYEMPDFDPTRISPWLLRI
ELDRKRMTDKKLTMEQIAEKINVGFGEDLNCIFNDDNADKLVLRIRIMNNEENKFQDEDE
AVDKMEDDMFLRCIEANMLSDMTLQGIEAIGKVYMHLPQTDSKKRIVITETGEFKAIGEW
LLETDGTSMMKVLSERDVDPIRTSSNDICEIFQVLGIEAVRKSVEKEMNAVLQFYGLYVN
YRHLALLCDVMTAKGHLMAITRHGINRQDTGALMRCSFEETVDVLMDAAAHAETDPMRGV
SENIIMGQLPKMGTGCFDLLLDAEKCRFGI

Sequence of entity 9 (B), FASTA

>9MU9_9 DNA-directed RNA polymerase II subunit RPB2 (chains B)
ENAEEISHELWQEACWIVINAYFDEKGLVRQQLDSFDEFIQMSVQRIVEDSPAIELQAEA
QHTSGEVETPPRFSLKFEQIYLSKPTHWEKDGSPSPMMPNEARLRNLTYSAPLYVDITKT
KNVEGLDPVETQHQKTFIGKIPIMLRSTYCLLSQLTDRDLTELNECPLDPGGYFIINGSE
KVLIAQEKMATNTVYVFSMKDGKYAFKTEIRSCLEHSSRPTSTLWVNMMARGSQNIKKSA
IGQRIIAILPYIKQEIPIMIVFRALGFVADRDILEHIIYDFDDPEMMEMVKPSLDEAFVV
QEQNVALNFIGARGARPGVTKDKRIKYAKEILQKEMLPHVGVSDFCETKKAYFLGYMVHR
LLLASLGRRELDDRDHYGNKRLDLAGPLLAFLFRGLFKNLMKEVRMYTQKFIDRGKDFNL
ELAIKTNIITDGLRYSLATGNWGDQKKAHQARAGVSQVLNRLTFASTLSHLRRVNSPIGR
DGKLAKPRQLHNTLWGMLCPAETPEGAAVGLVKNLALMAYISVGSQPSPILEFLEEWSME
NLEEIAPSAIADATKIFVNGCWVGIHRDPEQLMATLRKLRRQMDIIVSEVSMIRDIRDRE
IRIYTDAGRICRPLLIVENGSLLLKKTHVEMLKERDYNNYSWQVLVASGVVEYIDTLEEE
TVMIAMSPYDLKQDKDYAYCTTYTHCEIHPAMILGVCASIIPFPDHNQSPRNTYQSAMGK
QAMGVYITNFHVRMDTLAHVLYYPMKPLVTTRSMEYLRFRELPAGINSIVAILCYTGYNQ
EDSVILNASAVERGFFRSVFYRSYKDSENKRVGDQEENFEKPHRGTCQGMRNAHYDKLDD
DGIIAPGIRVSGDDVVIGKTITLPENDDELDSNTKRFSKRDASTFLRNSETGIVDQVMLT
LNSEGYKFCKIRVRSVRIPQIGDKFASRHGQKGTCGIQYRQEDMAFTCEGLAPDIIINPH
AIPSRMTIGHLIECLQGKLGSNKGEIGDATPFNDAVNVQKISTFLQEYGYHLRGNEVMYN
GHTGRKINAQVFLGPTYYQRLKHMVDDKIHSRARGPVQILVRQPMEGRARDGGLRFGEME
RDCQISHGAAQFLRERLFEVSDPYRVHICNFCGLIAIANLRNNTFECKGCKNKTQISQVR
LPYAAKLLFQELMSMNIAPRLMVT

Sequence of entity 10 (C), FASTA

>9MU9_10 DNA-directed RNA polymerase II subunit RPB3 (chains C)
PYANQPSVQITELTDDNVKFVLEDTELSVANSLRRVFIAETPTLAIDWVQLEANSTVLSD
EFLAHRIGLIPLISDDVVERLQYTRDCICLDFCPECSVEFTLDVKCSEEQTRHVTTADLK
SSNAKVLPVTSRNQGEEDNEYGESNDEILIIKLRKGQELKLRAYAKKGFGKEHAKWNPTA
GVCFEYDPDNSMRHTLYPKPDEWPKSEHTELEDDQYEAPYNWEAKPNKFFFNVESAGALK
PENIVVMGVQVLKNKLSNLQTQLSHESQN

Sequence of entity 11 (F), FASTA

>9MU9_11 DNA-directed RNA polymerases I, II, and III subunit RPABC2 (chains F)
GGVPKSKRITTKYMTKYERARVLGTRALQIAMCAPIMVELDGETDPLQIAMKELKQKKIP
IIIRRYLPDHSYEDWSIDELIMVD

Sequence of entity 12 (H), FASTA

>9MU9_12 DNA-directed RNA polymerases I, II, and III subunit RPABC3 (chains H)
AGVLFEDIFNVKDMDPEGKKFDRVSRLHCESESFKMDLILDINSWLYPMELGDKFRLVLA
TTLREDGCPDSGEYNPMEHEGTRADSFEYVMYGKIYRIEGDEAHNEASRLSAYVSFGGLL
MRLQGDANNLHGFEVDQHMYLLMKRLA

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	8

Primary citation

Structural Characterization of Native RNA Polymerase II Transcription Complexes and Nucleosomes in Drosophila melanogaster. Venette-Smith, N.L., Vishwakarma, R.K., Venkatakrishnan, V. et al. Nat Commun (2026) 17. DOI 10.1038/s41467-026-75963-0 · PubMed

Other PDB entries of the same protein (UniProt P84051 (AlphaFold model), which also has an AlphaFold model), best resolution first:

9ZQB 2.1 Å, Nucleosome with an SSB at SHL -2.8 in complex with human PARP2 and HPF1, Class 1
2NQB 2.3 Å, Drosophila Nucleosome Structure
9ZQC 2.37 Å, Nucleosome with an SSB at SHL -2.8 in complex with human PARP2 and HPF1, Class 2
2PYO 2.43 Å, Drosophila nucleosome core
8UX1 2.5 Å, Cryo-EM structure of Ran bound to RCC1 and the nucleosome core particle
8PP7 2.91 Å, human RYBP-PRC1 bound to mononucleosome
6DZT 2.99 Å, Cryo-EM structure of nucleosome in complex with a single chain antibody fragment
8PP6 3.18 Å, human RYBP-PRC1 bound to H2AK118ub1 nucleosome
9ZQA 3.28 Å, Nucleosome with an SSB at SHL -2.8 in complex with the WGR domain of human PARP2, Class 2
9MU4 3.29 Å, Structure of a native Drosophila melanogaster octameric nucleosome
9ZQ9 3.3 Å, Nucleosome with an SSB at SHL -2.8 in complex with the WGR domain of human PARP2, Class 1
4X23 3.5 Å, Crystal structure of cenp-C in complex with the nucleosome core particle

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