9V0E: Cullin-4A

Cryo-EM structure of golcadomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub ubiquitylation assembly. Determined by electron microscopy at 3.51 Å resolution. Released 19 Aug 2026.

Method: Electron microscopy
Resolution: 3.51 Å
Organism: Homo sapiens
Chains: 8
Atoms: 20,731
Mol. weight: 318.92 kDa
Ligands: A1AF4, ZN
Released: 19 Aug 2026

Explore 9V0E in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9V0E contains 84 α-helices and 154 β-strands across 8 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 A: 39 helices, 8 β-strands

Element	Residues	Length	Sheet
α-helix	58-74	17
α-helix	83-94	12
α-helix	99-116	18
α-helix	119-122	4
α-helix	128-152	25
α-helix	154-156	3
α-helix	157-161	5
α-helix	168-169	2
α-helix	170-181	12
α-helix	186-204	19
α-helix	211-223	13
α-helix	227-232	6
α-helix	233-251	19
α-helix	256-276	21
α-helix	280-282	3
α-helix	283-294	12
α-helix	296-298	3
α-helix	299-302	4
α-helix	303-307	5
α-helix	308-312	5
α-helix	316-328	13
α-helix	332-352	21
α-helix	356-360	5
α-helix	361-378	18
α-helix	384-398	15
α-helix	404-417	14
α-helix	426-440	15
α-helix	446-462	17
α-helix	469-483	15
α-helix	486-488	3
α-helix	489-512	24
β-strand	523	1	1
β-strand	526-529	4	1
α-helix	534-535	2
α-helix	545-561	17
β-strand	565-578	14	1
β-strand	589-591	3	1
α-helix	593-602	10
α-helix	610-617	8
α-helix	621-632	12
β-strand	640-642	3	2
β-strand	653-655	3	2
α-helix	693-707	15
α-helix	712-720	9
α-helix	729-741	13
β-strand	745-747	3	3
β-strand	755-757	3	3

Chain B: 12 helices, 98 β-strands

Element	Residues	Length	Sheet
β-strand	4-10	7	4
β-strand	18-21	4	5
β-strand	30-34	5	5
β-strand	38-43	6	5
β-strand	50-56	7	5
β-strand	63-67	5	6
β-strand	76-80	5	6
β-strand	85-94	10	6
β-strand	97-107	11	6
β-strand	115	1	7
β-strand	121-124	4	8
β-strand	130-134	5	8
β-strand	136	1	7
β-strand	139-144	6	8
β-strand	155-158	4	8
β-strand	163-167	5	9
β-strand	177-184	8	9
β-strand	187-196	10	9
β-strand	201-204	4	9
β-strand	210-211	2	9
β-strand	218-221	4	10
β-strand	229-232	4	10
β-strand	237-240	4	10
β-strand	245-248	4	10
α-helix	252-255	4
β-strand	258-263	6	11
β-strand	270-275	6	11
β-strand	279-287	9	11
β-strand	297-307	11	11
α-helix	310-311	2
β-strand	313-316	4	12
β-strand	321-325	5	12
β-strand	331-336	6	12
β-strand	347-353	7	12
β-strand	359-365	7	13
α-helix	367-369	3
β-strand	374-379	6	13
β-strand	386-391	6	13
β-strand	394	1	14
β-strand	396-403	8	15
β-strand	410-413	4	16
β-strand	424-428	5	16
β-strand	433-439	7	16
β-strand	442-446	5	16
β-strand	457-463	7	17
β-strand	467-472	6	17
β-strand	476-480	5	17
β-strand	485-490	6	17
β-strand	500-503	4	18
β-strand	507-512	6	18
β-strand	515-522	8	18
β-strand	525-533	9	18
β-strand	538-542	5	19
β-strand	555-560	6	19
β-strand	565-570	6	19
β-strand	575-581	7	19
α-helix	587	1
β-strand	588-595	8	20
β-strand	600-606	7	20
β-strand	610-615	6	20
β-strand	624-630	7	20
β-strand	637-643	7	21
β-strand	646-652	7	21
β-strand	653	1	22
β-strand	657-661	5	21
β-strand	666-670	5	21
β-strand	671	1	14
β-strand	676	1	22
β-strand	678-682	5	15
β-strand	690-694	5	15
β-strand	698-704	7	15
β-strand	711-712	2	13
β-strand	715-716	2	13
β-strand	720-727	8	23
β-strand	732-741	10	23
β-strand	751	1	23
β-strand	762-765	4	23
α-helix	766-768	3
β-strand	786-795	10	23
β-strand	801-806	6	23
α-helix	807-808	2
β-strand	811-819	9	24
β-strand	828-835	8	24
β-strand	845-854	10	24
β-strand	857-866	10	24
β-strand	874-876	3	25
β-strand	879-881	3	25
β-strand	890-893	4	25
β-strand	899-903	5	25
β-strand	913-917	5	26
β-strand	920-924	5	26
β-strand	930-936	7	26
β-strand	941-948	8	26
β-strand	957-961	5	27
β-strand	964-968	5	27
β-strand	974-978	5	27
β-strand	993-998	6	27
β-strand	1007-1009	3	4
β-strand	1028-1030	3	4
β-strand	1037-1042	6	4
α-helix	1046-1061	16
α-helix	1070-1073	4
β-strand	1076-1078	3	28
β-strand	1081-1083	3	28
β-strand	1086	1	27
β-strand	1088-1089	2	4
α-helix	1091-1095	5
α-helix	1096-1099	4
α-helix	1102-1108	7
α-helix	1126-1137	12

Chain C: 13 helices, 21 β-strands

Element	Residues	Length	Sheet
α-helix	54-56	3
β-strand	78-80	3	29
β-strand	83	1	29
β-strand	96-100	5	29
α-helix	104-115	12
β-strand	121-123	3	29
α-helix	124-125	2
β-strand	135-141	7	29
β-strand	156-172	17	29
β-strand	178-184	7	29
β-strand	188	1	30
α-helix	201-203	3
α-helix	207-210	4
α-helix	221-231	11
α-helix	233-237	5
α-helix	242-246	5
α-helix	250-264	15
α-helix	277-285	9
α-helix	292-300	9
β-strand	303	1	30
α-helix	304-315	12
β-strand	321-322	2	31
α-helix	334-336	3
β-strand	337	1	32
β-strand	349-350	2	33
β-strand	356-357	2	33
β-strand	359-362	4	32
β-strand	368-370	3	34
β-strand	375	1	32
β-strand	384-388	5	32
β-strand	389-391	3	34
β-strand	400-404	5	32
β-strand	414-418	5	32
β-strand	423-424	2	31

Chain D: 7 helices, 4 β-strands

Element	Residues	Length	Sheet
α-helix	4-15	12
β-strand	21-25	5	35
β-strand	32-38	7	35
α-helix	39-40	2
β-strand	49-55	7	35
β-strand	66-69	4	35
α-helix	87-90	4
α-helix	99-110	12
α-helix	121-129	9
α-helix	132-141	10
α-helix	142-146	5

Chain I: 4 helices, 2 β-strands

Element	Residues	Length	Sheet
β-strand	146-147	2	36
β-strand	154-155	2	36
α-helix	158-166	9
α-helix	187-188	2
α-helix	189-194	6
α-helix	195-196	2

Chain N: 4 helices, 9 β-strands

Element	Residues	Length	Sheet
β-strand	2	1	37
β-strand	4-6	3	38
β-strand	12-13	2	38
β-strand	16	1	37
β-strand	22	1	39
α-helix	23-34	12
α-helix	38-40	3
β-strand	41-45	5	38
β-strand	48-49	2	38
α-helix	50-51	2
β-strand	55	1	39
α-helix	56-59	4
β-strand	66-71	6	38

Chain R: 3 helices, 5 β-strands

Element	Residues	Length	Sheet
β-strand	26-35	10	1
α-helix	52-53	2
α-helix	69	1
β-strand	70-72	3	40
β-strand	73	1	41
β-strand	78-80	3	40
α-helix	81-90	10
β-strand	103	1	41

Chain U: 2 helices, 7 β-strands

Element	Residues	Length	Sheet
β-strand	2-6	5	42
β-strand	12-16	5	42
β-strand	22	1	43
α-helix	23-34	12
β-strand	41-44	4	42
β-strand	49	1	42
β-strand	55	1	43
α-helix	56-59	4
β-strand	65-71	7	42

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
Cullin-4A	A	protein	775	Homo sapiens	Q13619 (AlphaFold model)
DNA damage-binding protein 1	B	protein	1140	Homo sapiens	Q16531 (AlphaFold model)
Protein cereblon	C	protein	427	Homo sapiens	Q96SW2 (AlphaFold model)
Ubiquitin-conjugating enzyme E2 D1	D	protein	146	Homo sapiens	P51668 (AlphaFold model)
Zinc finger protein Aiolos	I	protein	58	Homo sapiens	Q9UKT9
NEDD8	N	protein	76	Homo sapiens	Q15843
E3 ubiquitin-protein ligase RBX1, N-terminally processed	R	protein	111	Homo sapiens	P62877
Ubiquitin	U	protein	76	Homo sapiens	P62979

Sequence of entity 1 (A), FASTA

>9V0E_1 Cullin-4A (chains A)
GPGSGWSHPQFEKGSGSADEAPRKGSFSALVGRTNGLTKPAALAAAPAKPGGAGGSKKLV
IKNFRDRPRLPDNYTQDTWRKLHEAVRAVQSSTSIRYNLEELYQAVENLCSHKVSPMLYK
QLRQACEDHVQAQILPFREDSLDSVLFLKKINTCWQDHCRQMIMIRSIFLFLDRTYVLQN
STLPSIWDMGLELFRTHIISDKMVQSKTIDGILLLIERERSGEAVDRSLLRSLLGMLSDL
QVYKDSFELKFLEETNCLYAAEGQRLMQEREVPEYLNHVSKRLEEEGDRVITYLDHSTQK
PLIACVEKQLLGEHLTAILQKGLDHLLDENRVPDLAQMYQLFSRVRGGQQALLQHWSEYI
KTFGTAIVINPEKDKDMVQDLLDFKDKVDHVIEVCFQKNERFVNLMKESFETFINKRPNK
PAELIAKHVDSKLRAGNKEATDEELERTLDKIMILFRFIHGKDVFEAFYKKDLAKRLLVG
KSASVDAEKSMLSKLKHECGAAFTSKLEGMFKDMELSKDIMVHFKQHMQNQSDSGPIDLT
VNILTMGYWPTYTPMEVHLTPEMIKLQEVFKAFYLGKHSGRKLQWQTTLGHAVLKAEFKE
GKKEFQVSLFQTLVLLMFNEGDGFSFEEIKMATGIEDSELRRTLQSLACGKARVLIKSPK
GKEVEDGDKFIFNGEFKHKLFRIKINQIQMKETVEEQVSTTERVFQDRQYQIDAAIVRIM
KMRKTLGHNLLVSELYNQLKFPVKPGDLKKRIESLIDRDYMERDKDNPNQYHYVA

Sequence of entity 2 (B), FASTA

>9V0E_2 DNA damage-binding protein 1 (chains B)
MSYNYVVTAQKPTAVNGCVTGHFTSAEDLNLLIAKNTRLEIYVVTAEGLRPVKEVGMYGK
IAVMELFRPKGESKDLLFILTAKYNACILEYKQSGESIDIITRAHGNVQDRIGRPSETGI
IGIIDPECRMIGLRLYDGLFKVIPLDRDNKELKAFNIRLEELHVIDVKFLYGCQAPTICF
VYQDPQGRHVKTYEVSLREKEFNKGPWKQENVEAEASMVIAVPEPFGGAIIIGQESITYH
NGDKYLAIAPPIIKQSTIVCHNRVDPNGSRYLLGDMEGRLFMLLLEKEEQMDGTVTLKDL
RVELLGETSIAECLTYLDNGVVFVGSRLGDSQLVKLNVDSNEQGSYVVAMETFTNLGPIV
DMCVVDLERQGQGQLVTCSGAFKEGSLRIIRNGIGIHEHASIDLPGIKGLWPLRSDPNRE
TDDTLVLSFVGQTRVLMLNGEEVEETELMGFVDDQQTFFCGNVAHQQLIQITSASVRLVS
QEPKALVSEWKEPQAKNISVASCNSSQVVVAVGRALYYLQIHPQELRQISHTEMEHEVAC
LDITPLGDSNGLSPLCAIGLWTDISARILKLPSFELLHKEMLGGEIIPRSILMTTFESSH
YLLCALGDGALFYFGLNIETGLLSDRKKVTLGTQPTVLRTFRSLSTTNVFACSDRPTVIY
SSNHKLVFSNVNLKEVNYMCPLNSDGYPDSLALANNSTLTIGTIDEIQKLHIRTVPLYES
PRKICYQEVSQCFGVLSSRIEVQDTSGGTTALRPSASTQALSSSVSSSKLFSSSTAPHET
SFGEEVEVHNLLIIDQHTFEVLHAHQFLQNEYALSLVSCKLGKDPNTYFIVGTAMVYPEE
AEPKQGRIVVFQYSDGKLQTVAEKEVKGAVYSMVEFNGKLLASINSTVRLYEWTTEKELR
TECNHYNNIMALYLKTKGDFILVGDLMRSVLLLAYKPMEGNFEEIARDFNPNWMSAVEIL
DDDNFLGAENAFNLFVCQKDSAATTDEERQHLQEVGLFHLGEFVNVFCHGSLVMQNLGET
STPTQGSVLFGTVNGMIGLVTSLSESWYNLLLDMQNRLNKVIKSVGKIEHSFWRSFHTER
KTEPATGFIDGDLIESFLDISRPKMQEVVANLQYDDGSGMKREATADDLIKVVEELTRIH

Sequence of entity 3 (C), FASTA

>9V0E_3 Protein cereblon (chains C)
GSSHHHHHHHHSSGLEVLFQGPGSSAKKPNIINFDTSLPTSHTYLGADMEEFHGRTLHDD
DSCQVIPVLPQVMMILIPGQTLPLQLFHPQEVSMVRNLIQKDRTFAVLAYSNVQEREAQF
GTTAEIYAYREEQDFGIEIVKVKAIGRQRFKVLELRTQSDGIQQAKVQILPECVLPSTMS
AVQLESLNKCQIFPSKPVSREDQCSYKWWQKYQKRKFHCANLTSWPRWLYSLYDAETLMD
RIKKQLREWDENLKDDSLPSNPIDFSYRVAACLPIDDVLRIQLLKIGSAIQRLRCELDIM
NKCTSLCCKQCQETEITTKNEIFSLSLCGPMAAYVNPHGYVHETLTVYKACNLNLIGRPS
TEHSWFPGYAWTVAQCKICASHIGWKFTATKKDMSPQKFWGLTRSALLPTIPDTEDEISP
DKVILCL

Sequence of entity 4 (D), FASTA

>9V0E_4 Ubiquitin-conjugating enzyme E2 D1 (chains D)
ALKRIQKELSDLQRDPPAHCSAGPVGDDLFHWQATIMGPPDSAYQGGVFFLTVHFPTDYP
FKPPKIAFTTKIYHPNINSNGSICLDILRSQWSPALTVSKVLLSICSLLCDPNPDDPLVP
DIAQIYKSDKEKYNRHAREWTQKYAM

Sequence of entity 5 (I), FASTA

>9V0E_5 Zinc finger protein Aiolos (chains I)
GPGERPFQCNQCGASFTQKGNLLRHICLHTGEKPFKCHLCNYATQRRDALTGHLRTHS

Sequence of entity 6 (N), FASTA

>9V0E_6 NEDD8 (chains N)
MLIKVKTLTGKEIEIDIEPTDKVERIKERVEEKEGIPPQQQRLIYSGKQMNDEKTAADYK
ILGGSVLHLVLALRGG

Sequence of entity 7 (R), FASTA

>9V0E_7 E3 ubiquitin-protein ligase RBX1, N-terminally processed (chains R)
GPGSAAAMDVDTPSGTNSGAGKKRFEVKKWNAVALWAWDIVVDNCAICRNHIMDLCIECQ
ANQASATSEECTVAWGVCNHAFHFHCISRWLKTRQVCPLDNREWEFQKYGH

Sequence of entity 8 (U), FASTA

>9V0E_8 Ubiquitin (chains U)
MQIFVKTLTGKTITLEVEPSDTIENVKAKIQDKEGIPPDQQRLIFAGKQLEDGRTLSDYN
IQKESTLHLVLRLRGG

Ligands and cofactors

ID	Name	Formula	Copies
A1AF4	Golcadomide	C28 H30 F N5 O5	1
ZN	Zinc ion	Zn	6

Primary citation

Cryo-EM structure of golcadomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub ubiquitylation assembly. Deng, Z.H., Ai, H.S., Liu, L. To be published.

Other PDB entries of the same protein (UniProt Q13619 (AlphaFold model), which also has an AlphaFold model), best resolution first:

7OPC 3.0 Å, Pol II-CSB-CRL4CSA-UVSSA-SPT6-PAF (Structure 4)
7OPD 3.0 Å, Pol II-CSB-CRL4CSA-UVSSA-SPT6-PAF (Structure 5)
2HYE 3.1 Å, Crystal Structure of the DDB1-Cul4A-Rbx1-SV5V Complex
9LWI 3.12 Å, The head region of HBx-Smc5/6 ubiquitination complex
9V09 3.25 Å, Cryo-EM structure of lenalidomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub…
9V0C 3.27 Å, Cryo-EM structure of iberdomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub…
9EG8 3.39 Å, Cryo-EM structure of COP9 signalosome precatalytic state with neddylated cullin-4A
9UUM 3.41 Å, Cryo-EM structure of mezigdomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub…
9UUQ 3.41 Å, Cryo-EM structure of thalidomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub…
8B3I 3.5 Å, CRL4CSA-E2-Ub (state 2)
9V0B 3.54 Å, Cryo-EM structure of avadomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub…
9V0A 3.69 Å, Cryo-EM structure of pomalidomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub…

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