9RMW: GluA4

GluA4 in complex with TARP-2, open state, structure of TMD/LBD domains. Determined by electron microscopy at 2.9 Å resolution. Released 24 Sept 2025.

Method: Electron microscopy
Resolution: 2.9 Å
Organism: Rattus norvegicus
Chains: 8
Atoms: 18,965
Mol. weight: 546.01 kDa
Ligands: PLM, OLC, CYZ, GLU
Released: 24 Sept 2025

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

Secondary structure: helices and β-sheets

9RMW contains 104 α-helices and 106 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: 18 helices, 18 β-strands

Element	Residues	Length	Sheet
β-strand	418-421	4	1
β-strand	429-430	2	2
α-helix	440-442	3
β-strand	443-444	2	2
α-helix	446-458	13
β-strand	463-466	4	1
β-strand	474-475	2	3
β-strand	482-483	2	3
α-helix	485-490	6
β-strand	497-498	2	4
α-helix	505-508	4
β-strand	511	1	5
β-strand	519-520	2	6
β-strand	522-527	6	7
β-strand	543	1	8
α-helix	545-567	23
α-helix	597-607	11
α-helix	618-646	29
α-helix	658-662	5
β-strand	668-671	4	7
α-helix	676-683	8
α-helix	687-697	11
α-helix	708-718	11
β-strand	722-727	6	7
α-helix	729-731	3
β-strand	742-745	4	7
β-strand	752-753	2	6
β-strand	756-757	2	4
β-strand	759	1	5
α-helix	766-777	12
α-helix	780-787	8
α-helix	808	1
β-strand	809	1	9
α-helix	810	1
α-helix	811-814	4
α-helix	815-845	31

Chains B and D: 23 helices, 24 β-strands

Element	Residues	Length	Sheet
β-strand	418	1	10
β-strand	419	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	463	1	10
β-strand	475	1	13
β-strand	482	1	13
α-helix	486-491	6
β-strand	496	1	11
β-strand	497	1	14
β-strand	502	1	15
β-strand	511	1	16
β-strand	519-520	2	17
β-strand	522-527	6	18
α-helix	529-535	7
α-helix	538-540	3
β-strand	543	1	19
α-helix	545-567	23
α-helix	595-606	12
α-helix	618-640	23
α-helix	642-646	5
α-helix	658-663	6
β-strand	669-671	3	18
β-strand	672	1	20
α-helix	679-682	4
α-helix	687-697	11
α-helix	702	1
β-strand	705	1	20
α-helix	708-717	10
β-strand	723-727	5	18
α-helix	728-736	9
β-strand	742-745	4	18
β-strand	752-753	2	17
β-strand	754	1	15
β-strand	757	1	14
β-strand	759	1	16
α-helix	766-776	11
α-helix	780-785	6
α-helix	786-790	5
α-helix	807-808	2
β-strand	809	1	8
α-helix	811-813	3
α-helix	815-845	31

Chain C: 18 helices, 18 β-strands

Element	Residues	Length	Sheet
β-strand	418-421	4	27
β-strand	429-430	2	28
α-helix	440-442	3
β-strand	443-444	2	28
α-helix	446-458	13
β-strand	463-466	4	27
β-strand	474-475	2	29
β-strand	482-483	2	29
α-helix	485-490	6
β-strand	497-498	2	30
α-helix	505-508	4
β-strand	511	1	31
β-strand	519-520	2	32
β-strand	522-527	6	33
β-strand	543	1	34
α-helix	545-567	23
α-helix	595-607	13
α-helix	618-646	29
α-helix	658-662	5
β-strand	668-671	4	33
α-helix	676-683	8
α-helix	687-697	11
α-helix	708-718	11
β-strand	722-727	6	33
α-helix	729-731	3
β-strand	742-745	4	33
β-strand	752-753	2	32
β-strand	756-757	2	30
β-strand	759	1	31
α-helix	766-777	12
α-helix	780-787	8
α-helix	808	1
β-strand	809	1	19
α-helix	810	1
α-helix	811-814	4
α-helix	815-845	31

Chains E and G: 6 helices, 6 β-strands

Element	Residues	Length	Sheet
α-helix	9-29	21
β-strand	34-38	5	21
α-helix	53-56	4
β-strand	57-61	5	21
β-strand	65-67	3	22
β-strand	68	1	21
β-strand	77-79	3	22
α-helix	97-104	8
α-helix	106-124	19
α-helix	133-160	28
β-strand	175-176	2	21
α-helix	178-216	39

Chains F and H: 5 helices, 5 β-strands

Element	Residues	Length	Sheet
α-helix	8-29	22
β-strand	34-36	3	23
β-strand	59-61	3	23
β-strand	65-67	3	24
β-strand	77-79	3	24
α-helix	93-104	12
α-helix	106-128	23
α-helix	134-161	28
β-strand	174-176	3	23
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

>9RMW_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

>9RMW_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
PLM	Palmitic acid	C16 H32 O2	18
OLC	(2R)-2,3-dihydroxypropyl (9Z)-octadec-9-enoate	C21 H40 O4	2
CYZ	Cyclothiazide	C14 H16 Cl N3 O4 S2	4
GLU	Glutamic acid	C5 H9 N O4	4
CA	Calcium ion	Ca	1

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
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
9P9B 3.31 Å, Activated GluA4 homotetrameric AMPAR.

Browse structure collections

About this viewer

MolViewer shows 9RMW 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.