9OVU: Composite map of GluA1/A2 in the activated state

Composite map of GluA1/A2 in the activated state, in complex with positive allosteric modulator (R,R)-2b and agonist glutamate (ATD-LBD-TMD). Determined by electron microscopy at 3.2 Å resolution. Released 8 Apr 2026.

Method: Electron microscopy
Resolution: 3.2 Å
Organism: Rattus norvegicus
Chains: 4
Atoms: 25,011
Mol. weight: 383 kDa
Ligands: FWF, GLU
Released: 8 Apr 2026

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

Secondary structure: helices and β-sheets

9OVU contains 138 α-helices and 172 β-strands across 4 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.

Chains A and C: 37 helices, 40 β-strands

Element	Residues	Length	Sheet
β-strand	12-13	2	1
β-strand	17-18	2	2
α-helix	26-35	10
β-strand	42-43	2	1
α-helix	57-66	10
β-strand	72-75	4	2
α-helix	82-92	11
β-strand	95-98	4	2
β-strand	110-112	3	2
α-helix	115-116	2
α-helix	118-125	8
β-strand	134-137	4	3
α-helix	139-141	3
α-helix	142-155	14
β-strand	163-164	2	3
α-helix	172-175	4
α-helix	177-181	5
β-strand	188-190	3	3
α-helix	194-204	11
β-strand	216	1	3
α-helix	228-230	3
β-strand	239	1	4
β-strand	242	1	5
α-helix	250-260	11
α-helix	273-274	2
α-helix	276-298	23
α-helix	322-330	9
β-strand	334-336	3	6
β-strand	339-341	3	6
β-strand	342	1	7
β-strand	350	1	7
β-strand	355	1	5
β-strand	357-359	3	4
β-strand	364-366	3	4
β-strand	392-395	4	8
β-strand	398	1	9
β-strand	402	1	9
β-strand	403-404	2	10
α-helix	414-416	3
β-strand	417-418	2	10
α-helix	420-423	4
α-helix	425-431	7
β-strand	437-440	4	8
β-strand	449-450	2	11
β-strand	455-456	2	11
α-helix	458-464	7
β-strand	470-471	2	8
α-helix	479-482	4
β-strand	485	1	12
β-strand	493-494	2	13
β-strand	496-501	6	14
α-helix	502-503	2
α-helix	505-508	4
β-strand	517	1	15
α-helix	522-529	8
α-helix	534-539	6
α-helix	572-578	7
α-helix	592-615	24
α-helix	632-637	6
β-strand	642-644	3	14
β-strand	646	1	16
α-helix	652-656	5
α-helix	663-671	9
β-strand	679	1	16
α-helix	682-688	7
β-strand	696-701	6	14
α-helix	702-708	7
β-strand	716-718	3	14
β-strand	726-727	2	13
β-strand	733	1	12
α-helix	739-751	13
α-helix	754-757	4
α-helix	759	1
α-helix	760-764	5
β-strand	783	1	17
α-helix	790-805	16
α-helix	808-811	4

Chains B and D: 32 helices, 46 β-strands

Element	Residues	Length	Sheet
β-strand	13	1	18
β-strand	19	1	19
α-helix	24-36	13
β-strand	44	1	18
β-strand	50	1	19
α-helix	55-66	12
β-strand	72	1	20
α-helix	84-92	9
β-strand	95	1	20
β-strand	98	1	21
β-strand	112	1	21
α-helix	115-116	2
α-helix	118-122	5
β-strand	132	1	22
β-strand	134	1	23
β-strand	135	1	24
α-helix	144-151	8
β-strand	159	1	22
β-strand	162	1	24
α-helix	173-181	9
β-strand	192	1	23
α-helix	198-200	3
α-helix	201-209	9
β-strand	221-222	2	25
β-strand	243-245	3	25
α-helix	253-263	11
α-helix	280-302	23
α-helix	321-323	3
α-helix	326-333	8
β-strand	338	1	26
β-strand	345	1	26
β-strand	346-347	2	27
β-strand	353-354	2	27
β-strand	359-365	7	25
β-strand	367-373	7	25
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	426-435	10
β-strand	440-444	5	28
β-strand	453	1	30
β-strand	460	1	30
α-helix	465-468	4
β-strand	474	1	28
β-strand	475	1	31
β-strand	480	1	32
α-helix	483-486	4
β-strand	489-490	2	33
β-strand	496-498	3	32
β-strand	500	1	34
β-strand	503	1	35
α-helix	507-509	3
β-strand	521	1	36
α-helix	523-541	19
α-helix	573-582	10
α-helix	591-593	3
α-helix	596-598	3
α-helix	605-619	15
β-strand	646-648	3	37
β-strand	650	1	38
α-helix	654-661	8
α-helix	665-676	12
β-strand	683	1	38
α-helix	686-695	10
β-strand	700-702	3	37
β-strand	705	1	34
α-helix	710-713	4
β-strand	722	1	35
β-strand	730-732	3	32
β-strand	735	1	31
β-strand	736-737	2	33
α-helix	743-756	14
α-helix	759-763	5
α-helix	764-768	5
β-strand	787	1	15
α-helix	793-810	18
α-helix	813-817	5

Molecules and chains

Molecule	Chains	Type	Length	Organism	UniProt
Isoform Flip of Glutamate receptor 1	A, C	protein	835	Rattus norvegicus	P19490 (AlphaFold model)
Isoform Flip of Glutamate receptor 2	B, D	protein	842	Rattus norvegicus	P19491 (AlphaFold model)

Sequence of entity 1 (A, C), FASTA

>9OVU_1 Isoform Flip of Glutamate receptor 1 (chains A, C)
MPYIFAFFCTGFLGAVVGANFPNNIQIGGLFPNQQSQEHAAFRFALSQLTEPPKLLPQID
IVNISDSFEMTYRFCSQFSKGVYAIFGFYERRTVNMLTSFCGALHVCFITPSFPVDTSNQ
FVLQLRPELQEALISIIDHYKWQTFVYIYDADRGLSVLQRVLDTAAEKNWQVTAVNILTT
TEEGYRMLFQDLEKKKERLVVVDCESERLNAILGQIVKLEKNGIGYHYILANLGFMDIDL
NKFKESGANVTGFQLVNYTDTIPARIMQQWRTSDSRDHTRVDWKRPKYTSALTYDGVKVM
AEAFQSLRRQRIDISRRGNAGDCLANPAVPWGQGIDIQRALQQVRFEGLTGNVQFNEKGR
RTNYTLHVIEMKHDGIRKIGYWNEDDKFVPAGGDNSSVQNRTYIVTTILEDPYVMLKKNA
NQFEGNDRYEGYCVELAAEIAKHVGYSYRLEIVSDGKYGARDPDTKAWNGMVGELVYGRA
DVAVAPLTITLVREEVIDFSKPFMSLGISIMIKKPQKSKPGVFSFLDPLAYEIWMCIVFA
YIGVSVVLFLVSRFSPYEWHSEEFEEGRDQTTSDQSNEFGIFNSLWFSLGAFMQQGCDIS
PRSLSGRIVGGVWWFFTLIIISSYTANLAAFLTVERMVSPIESAEDLAKQTEIAYGTLEA
GSTKEFFRRSKIAVFEKMWTYMKSAEPSVFVRTTEEGMIRVRKSKGKYAYLLESTMNEYI
EQRKPCDTMKVGGNLDSKGYGIATPKGSALRGPVNLAVLKLSEQGVLDKLKSKWWYDKGE
CGSKDSGSKDKTSALSLSNVAGVFYILIGGLGLAMLVALIEFCYKSRSESKRMKG

Sequence of entity 2 (B, D), FASTA

>9OVU_2 Isoform Flip of Glutamate receptor 2 (chains B, D)
MGKIMHISVLLSPVLWGLIFGVSSNSIQIGGLFPRGADQEYSAFRVGMVQFSTSEFRLTP
HIDNLEVANSFAVTNAFCSQFSRGVYAIFGFYDKKSVNTITSFCGTLHVSFITPSFPTDG
THPFVIQMRPDLKGALLSLIEYYQWDKFAYLYDSDRGLSTLQAVLDSAAEKKWQVTAINV
GNINNDKKDETYRSLFQDLELKKERRVILDCERDKVNDIVDQVITIGKHVKGYHYIIANL
GFTDGDLLKIQFGGAEVSGFQIVDYDDSLVSKFIERWSTLEEKEYPGAHTATIKYTSALT
YDAVQVMTEAFRNLRKQRIEISRRGNAGDCLANPAVPWGQGVEIERALKQVQVEGLSGNI
KFDQNGKRINYTINIMELKTNGPRKIGYWSEVDKMVLTEDDTSGLEQKTVVVTTILESPY
VMMKKNHEMLEGNERYEGYCVDLAAEIAKHCGFKYKLTIVGDGKYGARDADTKIWNGMVG
ELVYGKADIAIAPLTITLVREEVIDFSKPFMSLGISIMIKKPQKSKPGVFSFLDPLAYEI
WMCIVFAYIGVSVVLFLVSRFSPYEWHTEEFEDGRETQSSESTNEFGIFNSLWFSLGAFM
RQGCDISPRSLSGRIVGGVWWFFTLIIISSYTANLAAFLTVERMVSPIESAEDLSKQTEI
AYGTLDSGSTKEFFRRSKIAVFDKMWTYMRSAEPSVFVRTTAEGVARVRKSKGKYAYLLE
STMNEYIEQRKPCDTMKVGGNLDSKGYGIATPKGSSLGTPVNLAVLKLSEQGVLDKLKNK
WWYDKGECGAKDSGSKEKTSALSLSNVAGVFYILVGGLGLAMLVALIEFCYKSRAEAKRM
KG

Ligands and cofactors

ID	Name	Formula	Copies
FWF	N,N'-[biphenyl-4,4'-diyldi(2R)propane-2,1-diyl]dipropane-2-sulfonamide	C24 H36 N2 O4 S2	2
GLU	Glutamic acid	C5 H9 N O4	4

Water and common crystallization additives (NA) are not listed.

Primary citation

Auxiliary subunits reshape structural asymmetry and functional plasticity in heterotetrameric GluA1/A2 AMPA receptor core. Yen, L.Y., Newton, T.P., Yelshanskaya, M.V. et al. Nat Commun (2026) 17. DOI 10.1038/s41467-026-71063-1 · PubMed

Other PDB entries of the same protein (UniProt P19490 (AlphaFold model), which also has an AlphaFold model), best resolution first:

2AWW 2.21 Å, Synapse associated protein 97 PDZ2 domain variant C378G with C-terminal GluR-A peptide
3SAJ 2.5 Å, Crystal Structure of glutamate receptor GluA1 Amino Terminal Domain
8C2H 2.64 Å, Transmembrane domain of active state homomeric GluA1 AMPA receptor in tandem with TARP…
8C2I 2.7 Å, Transmembrane domain of resting state homomeric GluA1 AMPA receptor in complex with TARP…
8AYN 2.8 Å, Resting state GluA1/A2 AMPA receptor in complex with TARP gamma 8 and ligand LY3130481
8C1Q 2.82 Å, Resting state homomeric GluA1 AMPA receptor in complex with TARP gamma 3
8C1P 2.9 Å, Active state homomeric GluA1 AMPA receptor in complex with TARP gamma 3
7OCE 3.1 Å, Resting state GluA1/A2 AMPA receptor in complex with TARP gamma 8 and CNIH2 (LBD-TMD)
8AYL 3.2 Å, Resting state GluA1/A2 AMPA receptor in complex with TARP gamma 8 and ligand JNJ-61432059
9NR6 3.26 Å, The structure of Noelin 1 with cerebellar GluA1/A4-ATD
8AYM 3.3 Å, Resting state GluA1/A2 AMPA receptor in complex with TARP gamma 8 and ligand JNJ-55511118
8AYO 3.3 Å, Open state GluA1/A2 AMPA receptor in complex with TARP gamma 8 and ligand JNJ-61432059

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