LOCUS       NM_001113184            4141 bp    mRNA    linear   ROD 03-APR-2024
DEFINITION  Rattus norvegicus glutamate ionotropic receptor AMPA type subunit 4
            (Gria4), transcript variant 2, mRNA.
ACCESSION   NM_001113184
VERSION     NM_001113184.1
KEYWORDS    RefSeq.
SOURCE      Rattus norvegicus (Norway rat)
  ORGANISM  Rattus norvegicus
            Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi;
            Mammalia; Eutheria; Euarchontoglires; Glires; Rodentia; Myomorpha;
            Muroidea; Muridae; Murinae; Rattus.
REFERENCE   1  (bases 1 to 4141)
  AUTHORS   Rossner,B., Klingler,M., Bulat,T., Sase,A., Zeilinger,A.,
            Spitzwieser,M., Aradska,J., Cichna-Markl,M. and Lubec,G.
  TITLE     Hippocampal GluA2 and GluA4 protein but not corresponding mRNA and
            promoter methylation levels are modulated at retrieval in spatial
            learning of the rat
  JOURNAL   Amino Acids 49 (1), 117-127 (2017)
   PUBMED   27714514
  REMARK    GeneRIF: Hippocampal GluA2 and GluA4 protein but not corresponding
            mRNA and promoter methylation levels are modulated at retrieval in
            spatial learning of the rat.
REFERENCE   2  (bases 1 to 4141)
  AUTHORS   Luchkina,N.V., Coleman,S.K., Huupponen,J., Cai,C., Kivisto,A.,
            Taira,T., Keinanen,K. and Lauri,S.E.
  TITLE     Molecular mechanisms controlling synaptic recruitment of GluA4
            subunit-containing AMPA-receptors critical for functional
            maturation of CA1 glutamatergic synapses
  JOURNAL   Neuropharmacology 112 (Pt A), 46-56 (2017)
   PUBMED   27157711
  REMARK    GeneRIF: Data show that PKA activation leads to insertion of GluA4
            to synaptic sites with initially weak or silent AMPAR-mediated
            transmission. This suggests that the GluA4-mediated activation of
            silent synapses is a critical mechanism facilitating the functional
            maturation of glutamatergic circuitry during the critical period of
            experience-dependent fine-tuning.
REFERENCE   3  (bases 1 to 4141)
  AUTHORS   Herguedas,B., Garcia-Nafria,J., Cais,O., Fernandez-Leiro,R.,
            Krieger,J., Ho,H. and Greger,I.H.
  TITLE     Structure and organization of heteromeric AMPA-type glutamate
            receptors
  JOURNAL   Science 352 (6285), aad3873 (2016)
   PUBMED   26966189
REFERENCE   4  (bases 1 to 4141)
  AUTHORS   Chiasserini,D., van Weering,J.R., Piersma,S.R., Pham,T.V.,
            Malekzadeh,A., Teunissen,C.E., de Wit,H. and Jimenez,C.R.
  TITLE     Proteomic analysis of cerebrospinal fluid extracellular vesicles: a
            comprehensive dataset
  JOURNAL   J Proteomics 106, 191-204 (2014)
   PUBMED   24769233
REFERENCE   5  (bases 1 to 4141)
  AUTHORS   Aruscavage,P.J. and Bass,B.L.
  TITLE     A phylogenetic analysis reveals an unusual sequence conservation
            within introns involved in RNA editing
  JOURNAL   RNA 6 (2), 257-269 (2000)
   PUBMED   10688364
REFERENCE   6  (bases 1 to 4141)
  AUTHORS   Lomeli,H., Mosbacher,J., Melcher,T., Hoger,T., Geiger,J.R.,
            Kuner,T., Monyer,H., Higuchi,M., Bach,A. and Seeburg,P.H.
  TITLE     Control of kinetic properties of AMPA receptor channels by nuclear
            RNA editing
  JOURNAL   Science 266 (5191), 1709-1713 (1994)
   PUBMED   7992055
REFERENCE   7  (bases 1 to 4141)
  AUTHORS   Gallo,V., Upson,L.M., Hayes,W.P., Vyklicky,L. Jr., Winters,C.A. and
            Buonanno,A.
  TITLE     Molecular cloning and development analysis of a new glutamate
            receptor subunit isoform in cerebellum
  JOURNAL   J Neurosci 12 (3), 1010-1023 (1992)
   PUBMED   1372042
REFERENCE   8  (bases 1 to 4141)
  AUTHORS   Wenthold,R.J., Yokotani,N., Doi,K. and Wada,K.
  TITLE     Immunochemical characterization of the non-NMDA glutamate receptor
            using subunit-specific antibodies. Evidence for a hetero-oligomeric
            structure in rat brain
  JOURNAL   J Biol Chem 267 (1), 501-507 (1992)
   PUBMED   1309749
REFERENCE   9  (bases 1 to 4141)
  AUTHORS   Sommer,B., Keinanen,K., Verdoorn,T.A., Wisden,W., Burnashev,N.,
            Herb,A., Kohler,M., Takagi,T., Sakmann,B. and Seeburg,P.H.
  TITLE     Flip and flop: a cell-specific functional switch in
            glutamate-operated channels of the CNS
  JOURNAL   Science 249 (4976), 1580-1585 (1990)
   PUBMED   1699275
REFERENCE   10 (bases 1 to 4141)
  AUTHORS   Keinanen,K., Wisden,W., Sommer,B., Werner,P., Herb,A.,
            Verdoorn,T.A., Sakmann,B. and Seeburg,P.H.
  TITLE     A family of AMPA-selective glutamate receptors
  JOURNAL   Science 249 (4968), 556-560 (1990)
   PUBMED   2166337
COMMENT     REVIEWED REFSEQ: This record has been curated by NCBI staff. The
            reference sequence was derived from CB761818.1, BC093608.1,
            M36421.1, CK598329.1, M85037.1, BF413680.1, BF390245.1 and
            BF390683.1.
            
            Summary: Glutamate receptors are the predominant excitatory
            neurotransmitter receptors in the mammalian brain and are activated
            in a variety of normal neurophysiologic processes. These receptors
            are heteromeric protein complexes composed of multiple subunits,
            arranged to form ligand-gated ion channels. The classification of
            glutamate receptors is based on their activation by different
            pharmacologic agonists. The subunit encoded by this gene belongs to
            a family of AMPA (alpha-amino-3-hydroxy-5-methyl-4-isoxazole
            propionate)-sensitive glutamate receptors, and is subject to RNA
            editing (AGA->GGA; R->G). Alternative splicing of this gene results
            in transcript variants encoding different isoforms, which may vary
            in their signal transduction properties. [provided by RefSeq, Jul
            2008].
            
            Transcript Variant: This variant (2) uses an alternate internal
            coding exon compared to transcript variant 1, and encodes an
            isoform (2, also known as flop isoform) that is the same length,
            but with a few amino acid differences compared to isoform 1. RNA
            editing (AGA->GGA) changes Arg765Gly.
            
            Publication Note:  This RefSeq record includes a subset of the
            publications that are available for this gene. Please see the Gene
            record to access additional publications.
            
            ##Evidence-Data-START##
            RNAseq introns :: mixed sample support SAMEA5760383, SAMEA5760386
                              [ECO:0006172]
            ##Evidence-Data-END##
            
            ##RefSeq-Attributes-START##
            undergoes RNA editing :: PMID: 10688364
            ##RefSeq-Attributes-END##
            COMPLETENESS: complete on the 3' end.
PRIMARY     REFSEQ_SPAN         PRIMARY_IDENTIFIER PRIMARY_SPAN        COMP
            1-28                CB761818.1         1-28
            29-1719             BC093608.1         1-1691
            1720-2650           M36421.1           1270-2200
            2651-3171           CK598329.1         33-553
            3172-3249           M85037.1           2883-2960
            3250-3671           BF413680.1         11-432              c
            3672-3993           BF390245.1         146-467             c
            3994-4141           BF390683.1         1-148               c
FEATURES             Location/Qualifiers
     source          1..4141
                     /organism="Rattus norvegicus"
                     /mol_type="mRNA"
                     /strain="Sprague-Dawley"
                     /db_xref="taxon:10116"
                     /chromosome="8"
                     /map="8q11"
     gene            1..4141
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /note="glutamate ionotropic receptor AMPA type subunit 4"
                     /db_xref="GeneID:29629"
                     /db_xref="RGD:61863"
     exon            1..361
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            362..538
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     CDS             451..3159
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /note="isoform 2 precursor is encoded by transcript
                     variant 2; glutamate receptor, ionotropic, AMPA4 (alpha
                     4); AMPA-selective glutamate receptor 4; glutamate
                     receptor, ionotrophic, AMPA 4; glutamate receptor,
                     ionotropic, AMPA 4"
                     /codon_start=1
                     /product="glutamate receptor 4 isoform 2 precursor"
                     /protein_id="NP_001106655.1"
                     /db_xref="GeneID:29629"
                     /db_xref="RGD:61863"
                     /translation="MRIICRQIVLLFSGFWGLAMGAFPSSVQIGGLFIRNTDQEYTAF
                     RLAIFLHNTSPNASEAPFNLVPHVDNIETANSFAVTNAFCSQYSRGVFAIFGLYDKRS
                     VHTLTSFCSALHISLITPSFPTEGESQFVLQLRPSLRGALLSLLDHYEWNCFVFLYDT
                     DRGYSILQAIMEKAGQNGWHVSAICVENFNDVSYRQLLEELDRRQEKKFVIDCEIERL
                     QNILEQIVSVGKHVKGYHYIIANLGFKDISLERFIHGGANVTGFQLVDFNTPMVTKLM
                     DRWKKLDQREYPGSETPPKYTSALTYDGVLVMAETFRSLRRQKIDISRRGNAGDCLAN
                     PAAPWGQGIDMERTLKQVRIQGLTGNVQFDHYGRRVNYTMDVFELKSTGPRKVGYWND
                     MDKLVLIQDMPTLGNDTAAIENRTVVVTTIMESPYVMYKKNHEMFEGNDKYEGYCVDL
                     ASEIAKHIGIKYKIAIVPDGKYGARDADTKIWNGMVGELVYGKAEIAIAPLTITLVRE
                     EVIDFSKPFMSLGISIMIKKPQKSKPGVFSFLDPLAYEIWMCIVFAYIGVSVVLFLVS
                     RFSPYEWHTEEPEDGKEGPSDQPPNEFGIFNSLWFSLGAFMQQGCDISPRSLSGRIVG
                     GVWWFFTLIIISSYTANLAAFLTVERMVSPIESAEDLAKQTEIAYGTLDSGSTKEFFR
                     RSKIAVYEKMWTYMRSAEPSVFTRTTAEGVARVRKSKGKFAFLLESTMNEYIEQRKPC
                     DTMKVGGNLDSKGYGVATPKGSSLRNAVNLAVLKLNEQGLLDKLKNKWWYDKGECGSG
                     GGDSKDKTSALSLSNVAGVFYILVGGLGLAMLVALIEFCYKSRAEAKRMKLTFSEAIR
                     NKARLSITGSVGENGRVLTPDCPKAVHTGTAIRQSSGLAVIASDLP"
     exon            539..697
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            698..937
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            938..1122
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            1123..1176
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            1177..1335
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            1336..1503
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            1504..1608
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            1609..1719
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            1720..1926
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            1927..2297
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            2298..2496
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            2497..2744
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            2745..2859
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            2860..2994
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     exon            2995..4125
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
                     /inference="alignment:Splign:2.1.0"
     regulatory      4095..4100
                     /regulatory_class="polyA_signal_sequence"
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
     polyA_site      4125
                     /gene="Gria4"
                     /gene_synonym="GluA4; GluR-D; GluR4"
ORIGIN      
        1 gctgagtgga aagagctagg agccgggagg cagcctgtcc tccctcaagc ctgcaaggat
       61 gcggcagggg actgagacca tggggaggtg ctaacttcct cttcaacccc ccagcccggg
      121 atccgggctg gcatacccgg ccccccgggc tagtttcctg ccaagcgcaa ccaagaagcc
      181 aaaggcgatt tggagggcat aaaaagaaaa agagtggttc tggagagggg tgcccacctc
      241 cagagccggt ccctctgcaa acaggcaaag ataggagaaa ggggatcaga tactggagtg
      301 tacgcaaaat cctctgtctg tggactcgca ccagagcctc ccagaaaacc tgggcgatct
      361 gcgccatcgt cttcaatgcc tctctgaaaa gcctttagca agactgagag aaagagaaaa
      421 gagagcgcgc cagagagagg agcaaagaag atgaggatta tttgcaggca gattgtcttg
      481 ttgttttctg gattttgggg actcgccatg ggagcctttc caagcagcgt tcaaataggt
      541 ggtctcttca tccgaaacac agaccaggaa tacactgctt ttagactggc aatctttctt
      601 cataacacca gccccaatgc atcggaagct cctttcaatt tggtacctca tgtggacaac
      661 attgagactg ccaacagttt tgctgtgaca aacgccttct gttcccagta ttctagaggg
      721 gtgtttgcca tttttggact ctatgacaag agatccgtgc ataccttgac ctcgttctgc
      781 agtgctctgc acatctctct catcacacca agctttccca ctgaagggga gagccagttt
      841 gtgctgcagc taagaccttc actgagaggt gcactcctga gcctcctgga tcactatgag
      901 tggaactgtt tcgtcttcct gtatgataca gacagggggt attcaatact tcaagctata
      961 atggaaaaag caggacaaaa tggatggcat gtcagtgcaa tatgtgtgga aaattttaat
     1021 gatgtcagct acaggcaact gctagaagag cttgacagaa gacaagagaa gaaatttgtg
     1081 atagattgtg agatagagag gcttcaaaac attttagaac aaattgtgag tgttgggaag
     1141 catgtcaaag gctaccatta tatcatcgca aatttgggtt tcaaggatat ttctcttgag
     1201 agatttatac atggaggagc aaatgtaaca ggattccagt tggtagattt taatacaccc
     1261 atggtaacca aactaatgga tcggtggaag aaactagatc agagagaata tccaggttct
     1321 gaaacacctc caaagtacac ctctgctctc acttatgatg gagtcctggt gatggctgaa
     1381 actttccgaa gtctcagaag acagaaaatt gatatttcaa ggagaggaaa tgctggggac
     1441 tgtctggcaa accctgctgc tccctggggc cagggaattg acatggagag gacactgaag
     1501 caggttcgaa ttcaagggct gactgggaat gttcaatttg accattatgg acgtagagtt
     1561 aattacacaa tggatgtgtt tgaactgaaa agcacaggac ctcgaaaggt tggctactgg
     1621 aatgatatgg ataaattagt cttgattcaa gatatgccta ctcttggcaa tgacacagca
     1681 gctattgaga acagaacagt ggttgtaacc acaattatgg aatctcccta tgttatgtac
     1741 aagaaaaatc atgaaatgtt tgaaggaaat gacaagtacg aaggctactg tgtagatctg
     1801 gcatcggaaa ttgcaaaaca tattggtatc aaatataaaa ttgccattgt tcctgatgga
     1861 aaatatggag caagggacgc agacactaag atctggaatg ggatggtagg agagcttgtg
     1921 tatgggaaag cagagattgc tattgcccct ctgacaatca cattggttcg agaggaagtc
     1981 atcgattttt ctaagccttt tatgagttta ggcatctcta tcatgatcaa aaaacctcag
     2041 aaatctaaac caggagtctt ttccttcttg gaccctctgg cctatgagat ctggatgtgc
     2101 atagtgtttg catacattgg tgtcagtgtg gtcttgttcc tagtcagtag gtttagccca
     2161 tatgagtggc acacagaaga acctgaggat gggaaggaag gacccagtga ccagcctccc
     2221 aatgaatttg gcatctttaa cagcctttgg ttttccctgg gtgcctttat gcaacaagga
     2281 tgtgacattt cacccagatc cctgtcaggt cggattgttg gaggcgtgtg gtggttcttc
     2341 acactcatca ttatatcgtc ctacactgct aatctggctg cattccttac tgtggagaga
     2401 atggtctccc ccatagaaag tgcagaagac ctggccaaac aaacagaaat tgcctatgga
     2461 acacttgatt ctgggtcaac aaaagaattc ttcagaagat caaaaatagc agtgtatgaa
     2521 aagatgtgga cctacatgcg atcggcagag ccgtctgtgt tcactagaac tacagctgag
     2581 ggcgtggctc gtgtccgcaa gtccaagggc aaatttgcct ttctcctgga gtccacgatg
     2641 aatgaataca ttgagcagcg aaagccctgt gacacgatga aagtgggagg aaacctggat
     2701 tccaaaggct atggtgtagc aacgcccaag ggttcctcat taagaaatgc tgttaacctc
     2761 gcagttttaa aactgaatga acaaggcctc ttggacaaat tgaaaaacaa atggtggtac
     2821 gacaaaggag aatgtggcag cgggggaggt gactccaagg acaagacgag tgccttgagc
     2881 ctgagcaacg tagcaggcgt cttctacatt ctggttggcg gcctgggctt ggcaatgctg
     2941 gtggctttga tagagttctg ttacaagtcc agggcagagg cgaagagaat gaagctgact
     3001 ttttccgaag ccataagaaa caaagccagg ttatccatca ctgggagtgt gggagaaaac
     3061 ggccgtgtgc tgacccctga ctgccccaag gccgtacaca caggaactgc aattagacaa
     3121 agttcgggat tggctgtcat tgcatcggac ctaccataaa aaccaaaaaa ataattgagt
     3181 gccttaatca aactgtgttg gtgactgact gaaacgcagc cctgagggaa aggccaagag
     3241 tgggtcttgg ctaaatccat cctttggctt gagagcagga agagcatact gagtgccgga
     3301 cagaaccagt gcatgatttc agcacggaaa tgcagactcg gatttttttt tatcaggaaa
     3361 actcacaatt tagttttttt ggggatagag ggtgggggat agctgggata ggtgtattaa
     3421 cagcaacaaa tttcacatga gcgcacttaa aactaatcag actggtcagt tagtgcatta
     3481 aactgtgaag ttctttctca gaaaggtctt agtcactaca gcaacagatt caacagttac
     3541 tagaggtcaa ggaaccttct acctgtgtct ccagaacacc catttagtcg atggcataat
     3601 ctgcagctga aacaagcagt cactaagatg tgataagaaa ataataaata tgtaaatcct
     3661 gtgaaaaaaa acaacaaaat ttaaagaaag tatttacagt tactttggaa ataaaacaat
     3721 actgaaacat gcttgctttt taactgaagt aaatccagta gaggacaaca caattgtttt
     3781 ttctaaccat cttagggaac aatacattgc aataattgat aataatgcca tcactgtaat
     3841 aaactttaga gacttttttt taatgtaaaa gttgtttgtc accttgtttc ctgtaacctt
     3901 cactctgtca cacaagttgg ctcataggtt gtgtttgtct atcagaaaca acaacaaaaa
     3961 aaatgaagaa aatgttggca tgaagtcatc catctgcaac gtagaattta aaagactacg
     4021 ggtcattcat gttatcaata taatttataa tcctgttctg tgtacaaaat tgtggttttg
     4081 tacccaccca aaagaataaa acaacagttt ttcttacagt acctaaaaaa aaaaaaaaaa
     4141 a
//
