9UUM: Protein cereblon

Cryo-EM structure of mezigdomide-organized CRL4-DDB1-CRBN-IKZF3(ZF2-ZF3)-UbcH5a-Ub ubiquitylation assembly. Determined by electron microscopy at 3.41 Å resolution. Released 10 Jun 2026.

Method: Electron microscopy
Resolution: 3.41 Å
Organism: Homo sapiens
Chains: 8
Atoms: 20,807
Mol. weight: 318.95 kDa
Ligands: ZN, QFC
Released: 10 Jun 2026

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

Secondary structure: helices and β-sheets

9UUM contains 78 α-helices and 165 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: 37 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	170-178	9
α-helix	179-183	5
α-helix	186-204	19
α-helix	211-223	13
α-helix	227-231	5
α-helix	232-244	13
α-helix	246-251	6
α-helix	256-276	21
α-helix	280-282	3
α-helix	283-291	9
α-helix	292-296	5
α-helix	300-312	13
α-helix	316-328	13
α-helix	332-353	22
α-helix	356-359	4
α-helix	361-378	18
α-helix	384-398	15
α-helix	404-416	13
α-helix	426-440	15
α-helix	446-463	18
α-helix	469-483	15
α-helix	489-514	26
β-strand	522-523	2	15
β-strand	526-529	4	15
α-helix	545-561	17
β-strand	568-578	11	15
β-strand	589-591	3	15
α-helix	593-602	10
α-helix	610-616	7
α-helix	621-633	13
β-strand	640-641	2	47
β-strand	654-655	2	47
α-helix	694-707	14
α-helix	712-722	11
α-helix	729-741	13
β-strand	746-747	2	48
β-strand	755-756	2	48

Chain B: 10 helices, 102 β-strands

Element	Residues	Length	Sheet
β-strand	4-10	7	19
β-strand	15-21	7	20
β-strand	30-35	6	20
β-strand	38-43	6	20
β-strand	50-56	7	20
β-strand	63-67	5	21
β-strand	76-80	5	21
β-strand	85-94	10	21
β-strand	97-107	11	21
β-strand	115	1	22
β-strand	121-124	4	23
β-strand	130-134	5	23
β-strand	136	1	22
β-strand	139-144	6	23
β-strand	155-158	4	23
β-strand	163-169	7	24
β-strand	177-184	8	24
β-strand	187-196	10	24
β-strand	201-204	4	24
β-strand	210-211	2	24
β-strand	218-221	4	25
β-strand	229-232	4	25
β-strand	237-240	4	25
β-strand	245-248	4	25
α-helix	253-255	3
β-strand	258-263	6	26
β-strand	270-275	6	26
β-strand	279-287	9	26
β-strand	297-307	11	26
α-helix	310-311	2
β-strand	313-318	6	27
β-strand	321-325	5	27
β-strand	331-336	6	27
β-strand	347-353	7	27
β-strand	359-365	7	28
α-helix	367-369	3
β-strand	374-379	6	28
β-strand	386-387	2	29
β-strand	388-391	4	28
β-strand	394	1	30
β-strand	396-403	8	31
β-strand	411-413	3	32
β-strand	424-429	6	32
β-strand	432-439	8	32
β-strand	442-445	4	32
β-strand	457-463	7	33
β-strand	467-472	6	33
β-strand	476-480	5	33
β-strand	486-490	5	33
β-strand	500-503	4	34
β-strand	507-512	6	34
β-strand	515-522	8	34
β-strand	525-533	9	34
β-strand	538-542	5	35
β-strand	555-560	6	35
β-strand	565-570	6	35
β-strand	578-581	4	35
β-strand	591-595	5	36
β-strand	600-605	6	36
β-strand	610-615	6	36
β-strand	624-630	7	36
β-strand	639-642	4	37
β-strand	647-651	5	37
β-strand	657-661	5	37
β-strand	666-670	5	37
β-strand	671	1	30
β-strand	678	1	31
β-strand	681-682	2	31
β-strand	690-694	5	31
β-strand	698-704	7	31
β-strand	711-712	2	28
β-strand	715-716	2	29
β-strand	720-727	8	38
β-strand	732-741	10	38
β-strand	751	1	38
β-strand	764-765	2	38
β-strand	786-795	10	38
β-strand	801-806	6	38
β-strand	811-819	9	39
β-strand	828-835	8	39
β-strand	845-854	10	39
β-strand	857-866	10	39
β-strand	874-875	2	40
β-strand	880-881	2	40
β-strand	890	1	41
β-strand	893	1	42
β-strand	899	1	42
β-strand	903	1	41
β-strand	913-917	5	43
β-strand	920-924	5	43
β-strand	929	1	44
β-strand	932-936	5	43
β-strand	941-946	6	43
β-strand	949	1	44
β-strand	957-961	5	45
β-strand	964-968	5	45
β-strand	974-979	6	45
α-helix	988-990	3
β-strand	992-998	7	45
β-strand	1007-1009	3	19
β-strand	1028-1030	3	19
β-strand	1037-1042	6	19
α-helix	1045-1061	17
α-helix	1070-1073	4
β-strand	1075-1078	4	46
β-strand	1081-1084	4	46
β-strand	1086	1	45
β-strand	1088-1090	3	19
α-helix	1091-1095	5
α-helix	1096-1099	4
α-helix	1102-1108	7
α-helix	1126-1137	12

Chain C: 15 helices, 19 β-strands

Element	Residues	Length	Sheet
α-helix	54-56	3
β-strand	78-83	6	1
α-helix	90-92	3
β-strand	96-100	5	1
α-helix	104-114	11
β-strand	119-123	5	1
α-helix	124-125	2
β-strand	135-143	9	1
β-strand	156-167	12	1
β-strand	171	1	1
α-helix	172-173	2
β-strand	179-184	6	1
α-helix	201-203	3
α-helix	208-210	3
α-helix	221-231	11
α-helix	232-237	6
α-helix	242-246	5
α-helix	251-264	14
α-helix	277-285	9
α-helix	292-300	9
α-helix	304-316	13
β-strand	321-322	2	2
α-helix	334-336	3
β-strand	337	1	3
β-strand	346	1	3
β-strand	349	1	4
β-strand	357	1	4
β-strand	359-362	4	3
β-strand	368-370	3	3
β-strand	375	1	3
β-strand	384-391	8	3
β-strand	397-404	8	3
β-strand	415-418	4	3
β-strand	423-424	2	2

Chain D: 8 helices, 8 β-strands

Element	Residues	Length	Sheet
α-helix	4-15	12
β-strand	21-25	5	5
β-strand	32-38	7	5
α-helix	39-40	2
β-strand	49-55	7	5
α-helix	64-65	2
β-strand	66-69	4	5
β-strand	78	1	6
β-strand	83	1	5
β-strand	84	1	6
β-strand	86	1	7
α-helix	87-90	4
α-helix	99-111	13
α-helix	121-129	9
α-helix	132-141	10
α-helix	142-146	5

Chain I: 2 helices, 4 β-strands

Element	Residues	Length	Sheet
β-strand	146-147	2	8
β-strand	154-155	2	8
α-helix	158-167	10
β-strand	175	1	9
β-strand	182	1	9
α-helix	187-196	10

Chain N: 3 helices, 9 β-strands

Element	Residues	Length	Sheet
β-strand	2	1	10
β-strand	4-6	3	11
β-strand	12-13	2	11
β-strand	16	1	10
β-strand	22	1	12
α-helix	23-33	11
α-helix	38-40	3
β-strand	41-45	5	11
β-strand	48-49	2	11
α-helix	50-51	2
β-strand	55	1	12
β-strand	66-71	6	11

Chain R: 1 helix, 7 β-strands

Element	Residues	Length	Sheet
β-strand	26-33	8	15
β-strand	42	1	16
β-strand	47	1	16
β-strand	71-72	2	17
β-strand	73	1	18
β-strand	78-79	2	17
α-helix	81-90	10
β-strand	103	1	18

Chain U: 2 helices, 8 β-strands

Element	Residues	Length	Sheet
β-strand	2-6	5	13
β-strand	12-16	5	13
β-strand	22	1	14
α-helix	23-34	12
β-strand	41-44	4	13
β-strand	49	1	13
β-strand	55	1	14
α-helix	57-59	3
β-strand	65-71	7	13
β-strand	75	1	7

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
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 (AlphaFold model)
NEDD8	N	protein	76	Homo sapiens	Q15843 (AlphaFold model)
Ubiquitin	U	protein	76	Homo sapiens	P62979
E3 ubiquitin-protein ligase RBX1, N-terminally processed	R	protein	111	Homo sapiens	P62877
DNA damage-binding protein 1	B	protein	1140	Homo sapiens	Q16531
Cullin-4A	A	protein	775	Homo sapiens	Q13619

Sequence of entity 1 (C), FASTA

>9UUM_1 Protein cereblon (chains C)
GSSHHHHHHHHSSGLEVLFQGPGSSAKKPNIINFDTSLPTSHTYLGADMEEFHGRTLHDD
DSCQVIPVLPQVMMILIPGQTLPLQLFHPQEVSMVRNLIQKDRTFAVLAYSNVQEREAQF
GTTAEIYAYREEQDFGIEIVKVKAIGRQRFKVLELRTQSDGIQQAKVQILPECVLPSTMS
AVQLESLNKCQIFPSKPVSREDQCSYKWWQKYQKRKFHCANLTSWPRWLYSLYDAETLMD
RIKKQLREWDENLKDDSLPSNPIDFSYRVAACLPIDDVLRIQLLKIGSAIQRLRCELDIM
NKCTSLCCKQCQETEITTKNEIFSLSLCGPMAAYVNPHGYVHETLTVYKACNLNLIGRPS
TEHSWFPGYAWTVAQCKICASHIGWKFTATKKDMSPQKFWGLTRSALLPTIPDTEDEISP
DKVILCL

Sequence of entity 2 (D), FASTA

>9UUM_2 Ubiquitin-conjugating enzyme E2 D1 (chains D)
ALKRIQKELSDLQRDPPAHCSAGPVGDDLFHWQATIMGPPDSAYQGGVFFLTVHFPTDYP
FKPPKIAFTTKIYHPNINSNGSICLDILRSQWSPALTVSKVLLSICSLLCDPNPDDPLVP
DIAQIYKSDKEKYNRHAREWTQKYAM

Sequence of entity 3 (I), FASTA

>9UUM_3 Zinc finger protein Aiolos (chains I)
GPGERPFQCNQCGASFTQKGNLLRHICLHTGEKPFKCHLCNYATQRRDALTGHLRTHS

Sequence of entity 4 (N), FASTA

>9UUM_4 NEDD8 (chains N)
MLIKVKTLTGKEIEIDIEPTDKVERIKERVEEKEGIPPQQQRLIYSGKQMNDEKTAADYK
ILGGSVLHLVLALRGG

Sequence of entity 5 (U), FASTA

>9UUM_5 Ubiquitin (chains U)
MQIFVKTLTGKTITLEVEPSDTIENVKAKIQDKEGIPPDQQRLIFAGKQLEDGRTLSDYN
IQKESTLHLVLRLRGG

Sequence of entity 6 (R), FASTA

>9UUM_6 E3 ubiquitin-protein ligase RBX1, N-terminally processed (chains R)
GPGSAAAMDVDTPSGTNSGAGKKRFEVKKWNAVALWAWDIVVDNCAICRNHIMDLCIECQ
ANQASATSEECTVAWGVCNHAFHFHCISRWLKTRQVCPLDNREWEFQKYGH

Sequence of entity 7 (B), FASTA

>9UUM_7 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 8 (A), FASTA

>9UUM_8 Cullin-4A (chains A)
GPGSGWSHPQFEKGSGSADEAPRKGSFSALVGRTNGLTKPAALAAAPAKPGGAGGSKKLV
IKNFRDRPRLPDNYTQDTWRKLHEAVRAVQSSTSIRYNLEELYQAVENLCSHKVSPMLYK
QLRQACEDHVQAQILPFREDSLDSVLFLKKINTCWQDHCRQMIMIRSIFLFLDRTYVLQN
STLPSIWDMGLELFRTHIISDKMVQSKTIDGILLLIERERSGEAVDRSLLRSLLGMLSDL
QVYKDSFELKFLEETNCLYAAEGQRLMQEREVPEYLNHVSKRLEEEGDRVITYLDHSTQK
PLIACVEKQLLGEHLTAILQKGLDHLLDENRVPDLAQMYQLFSRVRGGQQALLQHWSEYI
KTFGTAIVINPEKDKDMVQDLLDFKDKVDHVIEVCFQKNERFVNLMKESFETFINKRPNK
PAELIAKHVDSKLRAGNKEATDEELERTLDKIMILFRFIHGKDVFEAFYKKDLAKRLLVG
KSASVDAEKSMLSKLKHECGAAFTSKLEGMFKDMELSKDIMVHFKQHMQNQSDSGPIDLT
VNILTMGYWPTYTPMEVHLTPEMIKLQEVFKAFYLGKHSGRKLQWQTTLGHAVLKAEFKE
GKKEFQVSLFQTLVLLMFNEGDGFSFEEIKMATGIEDSELRRTLQSLACGKARVLIKSPK
GKEVEDGDKFIFNGEFKHKLFRIKINQIQMKETVEEQVSTTERVFQDRQYQIDAAIVRIM
KMRKTLGHNLLVSELYNQLKFPVKPGDLKKRIESLIDRDYMERDKDNPNQYHYVA

Ligands and cofactors

ID	Name	Formula	Copies
ZN	Zinc ion	Zn	6
QFC	Mezigdomide	C32 H30 F N5 O4	1

Primary citation

Cryo-EM structure of mezigdomide-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 Q96SW2 (AlphaFold model), which also has an AlphaFold model), best resolution first:

4M91 1.1 Å, crystal structure of hN33/Tusc3-peptide 1
9CUO 1.6 Å, Crystal structure of CRBN with compound 3
9OHQ 1.6 Å, Structure of CRBN:IKZF2:Compound 15
9DOM 1.69 Å, PVTX-405: A Potent, Highly Selective, and Orally Efficacious Molecular Glue Degrader of…
7BQV 1.8 Å, Cereblon in complex with SALL4 and (S)-5-hydroxythalidomide
9O91 1.86 Å, Structure of IKZF2:CRBN:Compound 5 ternary structure
7BQU 1.9 Å, Cereblon in complex with SALL4 and (S)-thalidomide
9GAO 1.95 Å, Crystal structure of CRBNmidi in complex with 2-(4-(2,6-dioxopiperidin-3-yl)phenoxy)-N-me…
9FJX 2.0 Å, Crystal structure of human CRBN-DDB1 in complex with Lenalidomide
8RQC 2.15 Å, Crystal structure of CRBN-midi in complex with mezigdomide and IKZF1 ZF2
8RQ8 2.19 Å, Crystal structure of CRBN-midi in complex with mezigdomide
9OHR 2.34 Å, Structure of CRBN:IKZF2:Compound 35

Browse structure collections

About this viewer

MolViewer shows 9UUM directly in your browser with nothing to install. Switch between cartoon, ball-and-stick, spacefill and surface views, color by chain, secondary structure or B-factor, measure distances, angles and dihedrals, and share or embed the view.