9RN4: GluA4

GluA4 in complex with TARP-2, desensitized state, structure of TMD/LBD domains. Determined by electron microscopy at 3.4 Å resolution. Released 24 Sept 2025.

Method: Electron microscopy
Resolution: 3.4 Å
Organism: Rattus norvegicus
Chains: 8
Atoms: 18,546
Mol. weight: 540.98 kDa
Ligands: OLC, CA, PLM
Released: 24 Sept 2025

Explore 9RN4 in 3D Show helices and sheets RCSB PDB PDBe

Secondary structure: helices and β-sheets

9RN4 contains 113 α-helices and 110 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: 22 helices, 22 β-strands

Element	Residues	Length	Sheet
β-strand	417-421	5	1
β-strand	424	1	2
β-strand	428	1	2
β-strand	429-430	2	3
α-helix	431	1
α-helix	434-436	3
α-helix	439-442	4
β-strand	443-444	2	3
α-helix	446-458	13
β-strand	462-466	5	1
β-strand	475	1	4
β-strand	482	1	4
α-helix	484-490	7
β-strand	496-497	2	1
β-strand	502	1	5
α-helix	505-508	4
β-strand	511-513	3	1
α-helix	514	1
α-helix	516	1
β-strand	518-520	3	5
β-strand	522-526	5	6
β-strand	543	1	7
α-helix	545-567	23
α-helix	595-606	12
α-helix	618-648	31
α-helix	649-651	3
α-helix	658-662	5
β-strand	668-670	3	6
β-strand	672	1	8
α-helix	676-683	8
α-helix	687-698	12
β-strand	705	1	8
α-helix	708-717	10
β-strand	722-727	6	6
α-helix	728-735	8
β-strand	743-745	3	6
β-strand	752-754	3	5
β-strand	757-759	3	1
α-helix	765-777	13
α-helix	780-785	6
α-helix	786-790	5
β-strand	809	1	9
α-helix	816-831	16
α-helix	833-845	13

Chain B: 21 helices, 22 β-strands

Element	Residues	Length	Sheet
β-strand	417-421	5	10
β-strand	424	1	11
β-strand	428	1	11
β-strand	429-430	2	12
α-helix	431	1
α-helix	434-436	3
α-helix	439-442	4
β-strand	443-444	2	12
α-helix	446-458	13
β-strand	462-466	5	10
β-strand	475	1	13
β-strand	482	1	13
α-helix	484-490	7
β-strand	496-497	2	10
β-strand	502	1	14
α-helix	505-508	4
β-strand	511-513	3	10
α-helix	514	1
α-helix	516	1
β-strand	518-520	3	14
β-strand	522-527	6	15
α-helix	536-540	5
β-strand	543	1	16
α-helix	545-567	23
α-helix	597-606	10
α-helix	620-648	29
α-helix	658-662	5
β-strand	668-670	3	15
β-strand	672	1	17
α-helix	676-683	8
α-helix	687-698	12
β-strand	705	1	17
α-helix	708-717	10
β-strand	722-727	6	15
α-helix	728-735	8
β-strand	742-745	4	15
β-strand	752-754	3	14
β-strand	757-759	3	10
α-helix	765-777	13
α-helix	780-785	6
α-helix	786-790	5
β-strand	809	1	7
α-helix	815-843	29

Chain C: 25 helices, 22 β-strands

Element	Residues	Length	Sheet
β-strand	417-421	5	18
β-strand	424	1	19
β-strand	428	1	19
β-strand	429-430	2	20
α-helix	431	1
α-helix	434-436	3
α-helix	439-442	4
β-strand	443-444	2	20
α-helix	446-458	13
β-strand	462-466	5	18
β-strand	475	1	21
β-strand	482	1	21
α-helix	484-490	7
β-strand	496-497	2	18
β-strand	502	1	22
α-helix	505-508	4
β-strand	511-513	3	18
α-helix	514	1
α-helix	516	1
β-strand	518-520	3	22
β-strand	522-527	6	23
α-helix	538-540	3
β-strand	543	1	24
α-helix	545-547	3
α-helix	551-567	17
α-helix	595-605	11
α-helix	618-626	9
α-helix	632-647	16
α-helix	658-662	5
β-strand	668-670	3	23
β-strand	672	1	25
α-helix	676-683	8
α-helix	687-698	12
β-strand	705	1	25
α-helix	708-717	10
β-strand	722-727	6	23
α-helix	728-735	8
β-strand	742-745	4	23
β-strand	752-754	3	22
β-strand	757-759	3	18
α-helix	765-777	13
α-helix	780-785	6
α-helix	786-790	5
α-helix	806-808	3
β-strand	809	1	16
α-helix	816-819	4
α-helix	821-844	24

Chain D: 24 helices, 22 β-strands

Element	Residues	Length	Sheet
β-strand	417-421	5	26
β-strand	424	1	27
β-strand	428	1	27
β-strand	429-430	2	28
α-helix	431	1
α-helix	434-436	3
α-helix	439-442	4
β-strand	443-444	2	28
α-helix	446-458	13
β-strand	462-466	5	26
β-strand	475	1	29
β-strand	482	1	29
α-helix	484-490	7
β-strand	496-497	2	26
β-strand	502	1	30
α-helix	505-508	4
β-strand	511-513	3	26
α-helix	514	1
α-helix	516	1
β-strand	518-520	3	30
β-strand	522-527	6	31
β-strand	543	1	9
α-helix	545-549	5
α-helix	552-567	16
α-helix	595-606	12
α-helix	618-647	30
α-helix	658-662	5
β-strand	668-670	3	31
β-strand	672	1	32
α-helix	676-683	8
α-helix	687-698	12
β-strand	705	1	32
α-helix	708-717	10
β-strand	722-727	6	31
α-helix	728-735	8
β-strand	742-745	4	31
β-strand	752-754	3	30
β-strand	757-759	3	26
α-helix	765-777	13
α-helix	780-785	6
α-helix	786-790	5
α-helix	808	1
β-strand	809	1	24
α-helix	810	1
α-helix	811-814	4
α-helix	817-845	29

Chain E: 5 helices, 6 β-strands

Element	Residues	Length	Sheet
α-helix	8-29	22
β-strand	34-38	5	33
β-strand	57-61	5	33
β-strand	65-67	3	34
β-strand	68	1	33
β-strand	77-79	3	34
α-helix	94-104	11
α-helix	106-126	21
α-helix	133-162	30
β-strand	174-176	3	33
α-helix	178-211	34

Chain F: 6 helices, 6 β-strands

Element	Residues	Length	Sheet
α-helix	8-29	22
β-strand	34	1	35
β-strand	35-38	4	36
β-strand	57-60	4	36
β-strand	65-68	4	36
β-strand	77-79	3	36
α-helix	93-104	12
α-helix	106-127	22
α-helix	133-152	20
α-helix	155-158	4
β-strand	176	1	35
α-helix	178-211	34

Chain G: 5 helices, 5 β-strands

Element	Residues	Length	Sheet
α-helix	9-29	21
β-strand	34-38	5	37
β-strand	57-61	5	37
β-strand	65-68	4	37
β-strand	77-79	3	37
α-helix	94-104	11
α-helix	106-127	22
α-helix	133-160	28
β-strand	175-176	2	37
α-helix	178-211	34

Chain H: 5 helices, 5 β-strands

Element	Residues	Length	Sheet
α-helix	8-29	22
β-strand	34-38	5	38
β-strand	57-61	5	38
β-strand	65-68	4	38
β-strand	77-79	3	38
α-helix	96-99	4
α-helix	106-128	23
α-helix	133-162	30
β-strand	175-176	2	38
α-helix	178-211	34

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
Isoform 2 of Glutamate receptor 4	A, B, C, D	protein	882	Rattus norvegicus	P19493 (AlphaFold model)
Voltage-dependent calcium channel gamma-2 subunit	E, F, G, H	protein	323	Rattus norvegicus	Q71RJ2 (AlphaFold model)

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

>9RN4_1 Isoform 2 of Glutamate receptor 4 (chains A, B, C, D)
GAFPSSVQIGGLFIRNTDQEYTAFRLAIFLHNTSPNASEAPFNLVPHVDNIETANSFAVT
NAFCSQYSRGVFAIFGLYDKRSVHTLTSFCSALHISLITPSFPTEGESQFVLQLRPSLRG
ALLSLLDHYEWNCFVFLYDTDRGYSILQAIMEKAGQNGWHVSAICVENFNDVSYRQLLEE
LDRRQEKKFVIDCEIERLQNILEQIVSVGKHVKGYHYIIANLGFKDISLERFIHGGANVT
GFQLVDFNTPMVTKLMDRWKKLDQREYPGSETPPKYTSALTYDGVLVMAETFRSLRRQKI
DISRRGNAGDCLANPAAPWGQGIDMERTLKQVRIQGLTGNVQFDHYGRRVNYTMDVFELK
STGPRKVGYWNDMDKLVLIQDMPTLGNDTAAIENRTVVVTTIMESPYVMYKKNHEMFEGN
DKYEGYCVDLASEIAKHIGIKYKIAIVPDGKYGARDADTKIWNGMVGELVYGKAEIAIAP
LTITLVREEVIDFSKPFMSLGISIMIKKPQKSKPGVFSFLDPLAYEIWMCIVFAYIGVSV
VLFLVSRFSPYEWHTEEPEDGKEGPSDQPPNEFGIFNSLWFSLGAFMQQGCDISPRSLSG
RIVGGVWWFFTLIIISSYTANLAAFLTVERMVSPIESAEDLAKQTEIAYGTLDSGSTKEF
FRRSKIAVYEKMWTYMRSAEPSVFTRTTAEGVARVRKSKGKFAFLLESTMNEYIEQRKPC
DTMKVGGNLDSKGYGVATPKGSSLRTPVNLAVLKLSEAGVLDKLKNKWWYDKGECGPKDS
GSKDKTSALSLSNVAGVFYILVGGLGLAMLVALIEFCYKSRAEAKRMKLTFSEATRNKAR
LSITGSVGENGRVLTPDCPKAVHTGTAIRQSSGLAVIASDLP

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

>9RN4_2 Voltage-dependent calcium channel gamma-2 subunit (chains E, F, G, H)
MGLFDRGVQMLLTIVGAFAAFSLMTIAVGTDYWLYSRGVCKTKSVSENETSKKNEEVMTH
SGLWRTCCLEGNFKGLCKQIDHFPEDADYEADTAEYFLRAVRASSIFPILSVILLFMGGL
CIAASEFYKTRHNIILSAGIFFVSAGLSNIIGIIVYISANAGDPSKSDSKKNSYSYGWSF
YFGALSFIIAEMVGVLAVHMFIDRHKQLRATARATDYLQASAITRIPSYRYRYQRRSRSS
SRSTEPSHSRDASPVGVKGFNTLPSTEISMYTLSRDPLKAATTPTATYNSDRDNSFLQVH
NCIQKDSKDSLHANTANRRTTPV

Ligands and cofactors

ID	Name	Formula	Copies
OLC	(2R)-2,3-dihydroxypropyl (9Z)-octadec-9-enoate	C21 H40 O4	4
CA	Calcium ion	Ca	1
PLM	Palmitic acid	C16 H32 O2	4

Primary citation

GluA4 AMPA receptor gating mechanisms and modulation by auxiliary proteins. Vega-Gutierrez, C., Picanol-Parraga, J., Sanchez-Valls, I. et al. Nat Struct Mol Biol (2025) 32:2416-2428. DOI 10.1038/s41594-025-01666-7 · PubMed

Other PDB entries of the same protein (UniProt P19493 (AlphaFold model), which also has an AlphaFold model), best resolution first:

3FAS 1.4 Å, X-ray structure of iGluR4 flip ligand-binding core (S1S2) in complex with (S)-glutamate…
3EPE 1.85 Å, Crystal Structure of the GluR4 Ligand-Binding domain in complex with glutamate
3FAT 1.9 Å, X-ray structure of iGluR4 flip ligand-binding core (S1S2) in complex with (S)-AMPA at…
3KEI 1.9 Å, Crystal Structure of the GluA4 Ligand-Binding domain L651V mutant in complex with…
3KFM 2.2 Å, Crystal Structure of the GluA4 Ligand-Binding domain L651V mutant in complex with kainate
4GPA 2.25 Å, High resolution structure of the GluA4 N-terminal domain (NTD)
3EN3 2.43 Å, Crystal Structure of the GluR4 Ligand-Binding domain in complex with kainate
5FWX 2.5 Å, Crystal structure of the AMPA receptor GluA2/A4 N-terminal domain heterodimer
9QDN 2.71 Å, GluA4 in complex with TARP-2, resting state I, structure of TMD/LBD
9RMW 2.9 Å, GluA4 in complex with TARP-2, open state, structure of TMD/LBD domains
9RN7 3.1 Å, GluA4 in complex with TARP-2, Desensitized state, structure of TMD domain
9NR6 3.26 Å, The structure of Noelin 1 with cerebellar GluA1/A4-ATD

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