9E4Z: Isoform Flip of Glutamate receptor 2

GluA2-gamma2 complex bound glutamate and cyclothiazide. Determined by electron microscopy at 3.7 Å resolution. Released 18 Jun 2025.

Method: Electron microscopy
Resolution: 3.7 Å
Organisms: Rattus norvegicus, Mus musculus
Chains: 8
Atoms: 31,168
Mol. weight: 458.05 kDa
Ligands: CYZ, GLU
Released: 18 Jun 2025

Explore 9E4Z in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9E4Z contains 176 α-helices and 179 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: 34 helices, 36 β-strands

Element	Residues	Length	Sheet
β-strand	11-18	8	1
α-helix	23-36	14
β-strand	42-49	8	1
α-helix	55-67	13
β-strand	72-75	4	1
α-helix	82-92	11
β-strand	96-98	3	1
α-helix	102-103	2
β-strand	110-112	3	1
α-helix	115-116	2
α-helix	118-127	10
β-strand	132-137	6	2
α-helix	139-141	3
α-helix	144-155	12
β-strand	159-164	6	2
α-helix	170-172	3
α-helix	175-186	12
β-strand	191-195	5	2
α-helix	198-210	13
β-strand	219-222	4	2
β-strand	226	1	2
α-helix	227-229	3
α-helix	235-237	3
β-strand	241-246	6	2
α-helix	253-262	10
α-helix	280-301	22
α-helix	324-335	12
β-strand	338-340	3	3
β-strand	343-345	3	3
β-strand	346-347	2	4
β-strand	352	1	1
β-strand	353-354	2	4
β-strand	358-364	7	2
β-strand	367-374	8	2
β-strand	381-382	2	2
β-strand	395-399	5	5
β-strand	407-408	2	6
α-helix	412-414	3
β-strand	421-422	2	6
α-helix	424-436	13
β-strand	440-444	5	5
α-helix	464-467	4
β-strand	474-475	2	5
β-strand	480	1	7
α-helix	483-486	4
β-strand	489-491	3	5
β-strand	496-498	3	7
β-strand	500-505	6	8
α-helix	516-518	3
α-helix	523-544	22
α-helix	566-568	3
α-helix	573-582	10
α-helix	596-623	28
β-strand	646-649	4	8
β-strand	650	1	9
α-helix	654-661	8
α-helix	665-676	12
β-strand	683	1	9
α-helix	686-695	10
β-strand	700-705	6	8
α-helix	707-713	7
β-strand	720-723	4	8
β-strand	730-732	3	7
β-strand	735-737	3	5
α-helix	743-755	13
α-helix	758-767	10
α-helix	785-786	2
β-strand	787	1	10
α-helix	788	1
α-helix	789-819	31

Chain B: 39 helices, 40 β-strands

Element	Residues	Length	Sheet
β-strand	11-18	8	11
α-helix	23-36	14
β-strand	42-45	4	11
β-strand	48-49	2	11
α-helix	55-67	13
β-strand	72-75	4	11
α-helix	82-92	11
β-strand	95-98	4	11
α-helix	101-103	3
β-strand	110-112	3	11
α-helix	118-127	10
β-strand	132-137	6	12
α-helix	139-141	3
α-helix	144-155	12
β-strand	159-164	6	12
α-helix	170-172	3
α-helix	175-187	13
β-strand	191-195	5	12
α-helix	198-210	13
β-strand	219-222	4	12
β-strand	226	1	12
α-helix	233-235	3
β-strand	241-246	6	12
α-helix	253-263	11
α-helix	280-301	22
α-helix	324-333	10
β-strand	339-340	2	13
β-strand	343-344	2	13
β-strand	347	1	14
β-strand	353	1	14
β-strand	358-364	7	12
β-strand	367-374	8	12
β-strand	381-382	2	12
α-helix	386-388	3
α-helix	393-394	2
β-strand	395-399	5	15
β-strand	402	1	16
β-strand	406	1	16
β-strand	407-408	2	17
α-helix	412-414	3
β-strand	421-422	2	17
α-helix	424-436	13
β-strand	440-444	5	15
β-strand	453	1	18
β-strand	460	1	18
α-helix	462-468	7
β-strand	474-475	2	15
β-strand	480	1	19
α-helix	483-486	4
β-strand	489-491	3	15
α-helix	492	1
α-helix	494	1
β-strand	496-497	2	19
β-strand	500-505	6	20
α-helix	508-513	6
α-helix	516-518	3
β-strand	521	1	21
α-helix	523-543	21
α-helix	573-584	12
α-helix	599-620	22
α-helix	621-624	4
α-helix	638-641	4
β-strand	646-648	3	20
β-strand	650	1	22
α-helix	654-661	8
α-helix	665-674	10
β-strand	683	1	22
α-helix	686-695	10
β-strand	700-705	6	20
α-helix	706-712	7
β-strand	720-723	4	20
β-strand	731-732	2	19
β-strand	735-737	3	15
α-helix	743-755	13
α-helix	759-763	5
α-helix	764-768	5
α-helix	774-778	5
α-helix	789-791	3
α-helix	793-818	26

Chain C: 38 helices, 40 β-strands

Element	Residues	Length	Sheet
β-strand	11-17	7	23
α-helix	23-36	14
β-strand	42-48	7	23
α-helix	55-67	13
β-strand	72-75	4	23
α-helix	82-92	11
β-strand	96-98	3	23
α-helix	101-103	3
β-strand	110-112	3	23
α-helix	115-116	2
α-helix	118-127	10
β-strand	132-137	6	24
α-helix	139-141	3
α-helix	144-155	12
β-strand	159-164	6	24
α-helix	170-172	3
α-helix	175-185	11
β-strand	191-195	5	24
α-helix	198-210	13
β-strand	219-222	4	24
β-strand	226	1	24
α-helix	227-229	3
α-helix	233-236	4
β-strand	241-246	6	24
α-helix	253-263	11
α-helix	280-301	22
α-helix	324-335	12
β-strand	338-340	3	25
β-strand	343-345	3	25
β-strand	346-347	2	26
β-strand	352	1	23
β-strand	353-354	2	26
β-strand	358-364	7	24
β-strand	367-374	8	24
β-strand	381-382	2	24
β-strand	389	1	27
β-strand	392	1	27
β-strand	395-399	5	28
β-strand	407-408	2	29
α-helix	412-414	3
β-strand	421-422	2	29
α-helix	424-436	13
β-strand	440-444	5	28
β-strand	453	1	30
β-strand	460	1	30
α-helix	462-467	6
β-strand	474-475	2	28
α-helix	479	1
β-strand	480	1	31
α-helix	481	1
α-helix	483-486	4
β-strand	489-491	3	28
β-strand	496-498	3	31
β-strand	500-505	6	32
α-helix	511-513	3
α-helix	516-518	3
α-helix	523-542	20
α-helix	566-568	3
α-helix	573-582	10
α-helix	596-623	28
α-helix	639-641	3
β-strand	646-649	4	32
β-strand	650	1	33
α-helix	654-661	8
α-helix	665-676	12
β-strand	683	1	33
α-helix	686-695	10
β-strand	700-705	6	32
α-helix	706-712	7
β-strand	720-723	4	32
β-strand	730-732	3	31
β-strand	735-737	3	28
α-helix	743-755	13
α-helix	758-767	10
α-helix	785-786	2
β-strand	787	1	21
α-helix	788	1
α-helix	789-817	29

Chain D: 40 helices, 39 β-strands

Element	Residues	Length	Sheet
β-strand	11-17	7	34
α-helix	23-36	14
β-strand	42-48	7	34
α-helix	55-67	13
β-strand	72-74	3	34
α-helix	82-92	11
β-strand	96-98	3	34
α-helix	101-103	3
β-strand	110-112	3	34
α-helix	118-127	10
β-strand	132-137	6	35
α-helix	139-141	3
α-helix	144-155	12
β-strand	159-164	6	35
α-helix	170-172	3
α-helix	175-187	13
β-strand	191-195	5	35
α-helix	198-210	13
β-strand	215	1	36
β-strand	216	1	37
β-strand	218	1	37
β-strand	219-222	4	35
β-strand	226	1	35
α-helix	233-235	3
β-strand	239	1	36
β-strand	241-246	6	35
α-helix	253-263	11
α-helix	280-301	22
α-helix	325-335	11
β-strand	347	1	38
β-strand	353	1	38
β-strand	358-364	7	35
β-strand	367-374	8	35
β-strand	381-382	2	35
α-helix	388-391	4
β-strand	394-399	6	39
β-strand	402	1	40
β-strand	406	1	40
β-strand	407-408	2	41
α-helix	412-414	3
β-strand	421-422	2	41
α-helix	424-436	13
β-strand	439-444	6	39
β-strand	453	1	42
β-strand	460	1	42
α-helix	462-468	7
β-strand	474-475	2	39
α-helix	479	1
β-strand	480	1	39
α-helix	481	1
α-helix	483-486	4
β-strand	489-498	10	39
β-strand	500-505	6	43
α-helix	506-513	8
α-helix	516-518	3
β-strand	521	1	10
α-helix	523-543	21
α-helix	573-584	12
α-helix	598-616	19
α-helix	617-619	3
α-helix	621-623	3
α-helix	638-640	3
β-strand	646-648	3	43
β-strand	650	1	44
α-helix	654-661	8
α-helix	665-676	12
α-helix	680	1
β-strand	683	1	44
α-helix	686-695	10
β-strand	700-705	6	43
α-helix	706-712	7
β-strand	720-723	4	43
β-strand	730-737	8	39
α-helix	743-754	12
α-helix	758-763	6
α-helix	764-768	5
α-helix	775-778	4
α-helix	789-791	3
α-helix	793-818	26

Chain E: 6 helices, 6 β-strands

Element	Residues	Length	Sheet
α-helix	1007-1028	22
β-strand	1033-1037	5	45
β-strand	1056-1060	5	45
β-strand	1064-1065	2	46
β-strand	1067	1	45
β-strand	1077-1078	2	46
α-helix	1093-1102	10
α-helix	1105-1120	16
α-helix	1121-1124	4
α-helix	1133-1159	27
β-strand	1174-1175	2	45
α-helix	1177-1208	32

Chain F: 7 helices, 5 β-strands

Element	Residues	Length	Sheet
α-helix	1005-1028	24
β-strand	1033-1037	5	47
β-strand	1056-1060	5	47
β-strand	1064-1068	5	47
β-strand	1077-1078	2	47
α-helix	1092-1102	11
α-helix	1105-1120	16
α-helix	1121-1124	4
α-helix	1133-1159	27
β-strand	1174-1175	2	47
α-helix	1177-1203	27
α-helix	1205-1208	4

Chain G: 6 helices, 7 β-strands

Element	Residues	Length	Sheet
α-helix	1007-1028	22
β-strand	1033-1037	5	48
β-strand	1056-1060	5	48
β-strand	1064-1068	5	48
β-strand	1077-1078	2	48
β-strand	1084	1	49
β-strand	1087	1	49
α-helix	1092-1101	10
α-helix	1105-1120	16
α-helix	1121-1124	4
α-helix	1133-1159	27
β-strand	1174-1175	2	48
α-helix	1177-1207	31

Chain H: 6 helices, 6 β-strands

Element	Residues	Length	Sheet
α-helix	1008-1028	21
β-strand	1033-1034	2	50
β-strand	1037	1	50
β-strand	1056-1060	5	50
β-strand	1064-1070	7	50
β-strand	1077-1078	2	50
α-helix	1085-1091	7
α-helix	1092-1102	11
α-helix	1105-1120	16
α-helix	1132-1158	27
β-strand	1174-1175	2	50
α-helix	1177-1208	32

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
Isoform Flip of Glutamate receptor 2	A, B, C, D	protein	811	Rattus norvegicus	P19491 (AlphaFold model)
Voltage-dependent calcium channel gamma-2 subunit	E, F, G, H	protein	208	Mus musculus	O88602 (AlphaFold model)

Sequence of entity 1 (A, B, C, D), FASTA

>9E4Z_1 Isoform Flip of Glutamate receptor 2 (chains A, B, C, D)
NSIQIGGLFPRGADQEYSAFRVGMVQFSTSEFRLTPHIDNLEVANSFAVTNAFCSQFSRG
VYAIFGFYDKKSVNTITSFCGTLHVSFITPSFPTDGTHPFVIQMRPDLKGALLSLIEYYQ
WDKFAYLYDSDRGLSTLQAVLDSAAEKKWQVTAINVGNINNDKKDETYRSLFQDLELKKE
RRVILDCERDKVNDIVDQVITIGKHVKGYHYIIANLGFTDGDLLKIQFGGAEVSGFQIVD
YDDSLVSKFIERWSTLEEKEYPGAHTATIKYTSALTYDAVQVMTEAFRNLRKQRIEISRR
GNAGDCLANPAVPWGQGVEIERALKQVQVEGLSGNIKFDQNGKRINYTINIMELKTNGPR
KIGYWSEVDKMVLTEDDTSGLEQKTVVVTTILESPYVMMKKNHEMLEGNERYEGYCVDLA
AEIAKHCGFKYKLTIVGDGKYGARDADTKIWNGMVGELVYGKADIAIAPLTITLVREEVI
DFSKPFMSLGISIMIKKPQKSKPGVFSFLDPLAYEIWMCIVFAYIGVSVVLFLVSRFSPY
EWHTEEFEDGRETQSSESTNEFGIFNSLWFSLGAFMQQGCDISPRSLSGRIVGGVWWFFT
LIIISSYTANLAAFLTVERMVSPIESAEDLSKQTEIAYGTLDSGSTKEFFRRSKIAVFDK
MWTYMRSAEPSVFVRTTAEGVARVRKSKGKYAYLLESTMNEYIEQRKPCDTMKVGGNLDS
KGYGIATPKGSSLGTPVNLAVLKLSEQGVLDKLKNKWWYDKGECGAKDSGSKEKTSALSL
SNVAGVFYILVGGLGLAMLVALIEFCYKSRA

Sequence of entity 2 (E, F, G, H), FASTA

>9E4Z_2 Voltage-dependent calcium channel gamma-2 subunit (chains E, F, G, H)
LFDRGVQMLLTTVGAFAAFSLMTIAVGTDYWLYSRGVCKTKSVSENETSKKNEEVMTHSG
LWRTCCLEGNFKGLCKQIDHFPEDADYEADTAEYFLRAVRASSIFPILSVILLFMGGLCI
AASEFYKTRHNIILSAGIFFVSAGLSNIIGIIVYISANAGDPSKSDSKKNSYSYGWSFYF
GALSFIIAEMVGVLAVHMFIDRHKQLTG

Ligands and cofactors

ID	Name	Formula	Copies
CYZ	Cyclothiazide	C14 H16 Cl N3 O4 S2	4
GLU	Glutamic acid	C5 H9 N O4	4

Primary citation

Memantine inhibits calcium-permeable AMPA receptors. Carrillo, E., Montano Romero, A., Gonzalez, C.U. et al. Nat Commun (2025) 16:5576-5576. DOI 10.1038/s41467-025-60543-5 · PubMed

Other PDB entries of the same protein (UniProt P19491 (AlphaFold model), which also has an AlphaFold model), best resolution first:

6YK4 1.0 Å, Structure of the AMPA receptor GluA2o ligand-binding domain (S1S2J) in complex with the…
5NG9 1.15 Å, Crystal structure of the GluA2 ligand-binding domain (S1S2J) in complex with agonist…
6YK5 1.15 Å, Structure of the AMPA receptor GluA2o ligand-binding domain (S1S2J) in complex with the…
6YK3 1.2 Å, Structure of the AMPA receptor GluA2o ligand-binding domain (S1S2J) in complex with the…
5JEI 1.23 Å, Crystal structure of the GluA2 LBD in complex with FW
4IGT 1.24 Å, Crystal structure of the GluA2 ligand-binding domain (S1S2J) in complex with the agonist…
4YU0 1.26 Å, Crystal structure of a tetramer of GluA2 TR mutant ligand binding domains bound with…
5NIH 1.3 Å, Crystal structure of the GluA2 ligand-binding domain (S1S2J) in complex with agonist…
1MQI 1.35 Å, Crystal Structure of the GluR2 Ligand Binding Core (S1S2J) in Complex with…
5FTI 1.35 Å, Crystal structure of the GluA2 K738M-T744K LBD in complex with glutamate (lithium form)
4U21 1.39 Å, GluA2flip sLBD complexed with FW and (R,R)-2b crystal form E
4FAT 1.4 Å, Ligand-binding domain of GluA2 (flip) ionotropic glutamate receptor in complex with an…

Browse structure collections

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