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{{Infobox_gene}}
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'''Glutamate [NMDA] receptor subunit epsilon-2''', also known as '''''N''-methyl D-aspartate receptor subtype 2B''' ('''NMDAR2B''' or '''NR2B'''), is a [[protein]] that in humans is encoded by the ''GRIN2B'' [[gene]].<ref name="pmid1350383">{{cite journal | vauthors = Monyer H, Sprengel R, Schoepfer R, Herb A, Higuchi M, Lomeli H, Burnashev N, Sakmann B, Seeburg PH | title = Heteromeric NMDA receptors: molecular and functional distinction of subtypes | journal = Science | volume = 256 | issue = 5060 | pages = 1217–21 | date = May 1992 | pmid = 1350383 | pmc = | doi = 10.1126/science.256.5060.1217 }}</ref>
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| require_manual_inspection = no
| update_protein_box = yes
| update_summary = yes
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== NMDA receptors ==
{{GNF_Protein_box
''N''-methyl-D-aspartate ([[NMDA receptor|NMDA]]) receptors are a class of [[ionotropic glutamate receptor]]s. The NMDA receptor channel has been shown to be involved in [[long-term potentiation]], an activity-dependent increase in the efficiency of synaptic transmission thought to underlie certain kinds of memory and learning. NMDA receptor channels are heterotetramers composed of two molecules of the key receptor subunit NMDAR1 ([[GRIN1]]) and two drawn from one or more of the four NMDAR2 subunits: NMDAR2A ([[GRIN2A]]), NMDAR2B (GRIN2B), NMDAR2C ([[GRIN2C]]), and NMDAR2D ([[GRIN2D]]). The NR2 subunit acts as the agonist binding site for [[glutamic acid|glutamate]], one of the predominant excitatory neurotransmitter receptors in the mammalian brain.<ref>{{cite web | title = Entrez Gene: GRIN2B glutamate receptor, ionotropic, N-methyl D-aspartate 2B| url = https://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=2904| accessdate = }}</ref>
| image =
| image_source =
| PDB =  
| Name = Glutamate receptor, ionotropic, N-methyl D-aspartate 2B
| HGNCid = 4586
| Symbol = GRIN2B
| AltSymbols =; MGC142178; MGC142180; NMDAR2B; NR2B; hNR3
| OMIM = 138252
| ECnumber = 
| Homologene = 646
| MGIid = 95821
| GeneAtlas_image1 = PBB_GE_GRIN2B_210412_at_tn.png
| GeneAtlas_image2 = PBB_GE_GRIN2B_210411_s_at_tn.png
| Function = {{GNF_GO|id=GO:0000287 |text = magnesium ion binding}} {{GNF_GO|id=GO:0004872 |text = receptor activity}} {{GNF_GO|id=GO:0004972 |text = N-methyl-D-aspartate selective glutamate receptor activity}} {{GNF_GO|id=GO:0005216 |text = ion channel activity}} {{GNF_GO|id=GO:0005234 |text = extracellular-glutamate-gated ion channel activity}} {{GNF_GO|id=GO:0005509 |text = calcium ion binding}} {{GNF_GO|id=GO:0005515 |text = protein binding}}
| Component = {{GNF_GO|id=GO:0005624 |text = membrane fraction}} {{GNF_GO|id=GO:0005887 |text = integral to plasma membrane}} {{GNF_GO|id=GO:0008021 |text = synaptic vesicle}} {{GNF_GO|id=GO:0042734 |text = presynaptic membrane}} {{GNF_GO|id=GO:0045211 |text = postsynaptic membrane}}
| Process = {{GNF_GO|id=GO:0006811 |text = ion transport}} {{GNF_GO|id=GO:0007215 |text = glutamate signaling pathway}} {{GNF_GO|id=GO:0007268 |text = synaptic transmission}} {{GNF_GO|id=GO:0007611 |text = learning and/or memory}}
| Orthologs = {{GNF_Ortholog_box
    | Hs_EntrezGene = 2904
    | Hs_Ensembl = ENSG00000150086
    | Hs_RefseqProtein = NP_000825
    | Hs_RefseqmRNA = NM_000834
    | Hs_GenLoc_db = 
    | Hs_GenLoc_chr = 12
    | Hs_GenLoc_start = 13605411
    | Hs_GenLoc_end = 14024319
    | Hs_Uniprot = Q13224
    | Mm_EntrezGene = 14812
    | Mm_Ensembl = ENSMUSG00000030209
    | Mm_RefseqmRNA = NM_008171
    | Mm_RefseqProtein = NP_032197
    | Mm_GenLoc_db = 
    | Mm_GenLoc_chr = 6
    | Mm_GenLoc_start = 135697791
    | Mm_GenLoc_end = 136009994
    | Mm_Uniprot = Q8CG69
  }}
}}
'''Glutamate receptor, ionotropic, N-methyl D-aspartate 2B''', also known as '''GRIN2B''', is a human [[gene]].


<!-- The PBB_Summary template is automatically maintained by Protein Box Bot.  See Template:PBB_Controls to Stop updates. -->
== Function ==
{{PBB_Summary
NR2B has been associated with age- and [[Visual system|visual]]-experience-dependent plasticity in the [[neocortex]] of [[rat]]s, where an increased NR2B/NR2A ratio correlates directly with the stronger excitatory LTP in young animals. This is thought to contribute to experience-dependent refinement of developing [[Cortex (anatomy)|cortical]] circuits.<ref>{{cite journal | vauthors = Yoshimura Y, Ohmura T, Komatsu Y | title = Two forms of synaptic plasticity with distinct dependence on age, experience, and NMDA receptor subtype in rat visual cortex | journal = The Journal of Neuroscience | volume = 23 | issue = 16 | pages = 6557–66 | date = July 2003 | pmid = 12878697 | url = http://www.jneurosci.org/content/23/16/6557 }}</ref>
| section_title =  
| summary_text = N-methyl-D-aspartate (NMDA) receptors are a class of ionotropic glutamate receptors. NMDA receptor channel has been shown to be involved in long-term potentiation, an activity-dependent increase in the efficiency of synaptic transmission thought to underlie certain kinds of memory and learning.  NMDA receptor channels are heteromers composed of the key receptor subunit NMDAR1 (GRIN1) and 1 or more of the 4 NMDAR2 subunits: NMDAR2A (GRIN2A), NMDAR2B (GRIN2B), NMDAR2C (GRIN2C), and NMDAR2D (GRIN2D). GRIN2B may be a candidate gene for the neurodegenerative disorder dentato-rubro-pallidoluysian atrophy (DRPLA)<ref>{{cite web | title = Entrez Gene: GRIN2B glutamate receptor, ionotropic, N-methyl D-aspartate 2B| url = http://www.ncbi.nlm.nih.gov/sites/entrez?Db=gene&Cmd=ShowDetailView&TermToSearch=2904| accessdate = }}</ref>
}}


==See also==
Both mice and rats that were engineered to over-express GRIN2B in their brains have increased mental ability.<ref name="tang1999">{{cite journal | vauthors = Tang YP, Shimizu E, Dube GR, Rampon C, Kerchner GA, Zhuo M, Liu G, Tsien JZ | title = Genetic enhancement of learning and memory in mice | journal = Nature | volume = 401 | issue = 6748 | pages = 63–9 | date = September 1999 | pmid = 10485705 | doi = 10.1038/43432 }}</ref><ref>{{cite journal | vauthors = Wang D, Cui Z, Zeng Q, Kuang H, Wang LP, Tsien JZ, Cao X | title = Genetic enhancement of memory and long-term potentiation but not CA1 long-term depression in NR2B transgenic rats | journal = PLOS ONE | volume = 4 | issue = 10 |page = e7486 | date = October 2009 | pmid = 19838302 | doi = 10.1371/journal.pone.0007486 | pmc=2759522}}</ref>
 
== Ligands ==
* [[Besonprodil]]
* [[CERC-301]], a selective NR2B receptor antagonist
* [[Eliprodil]]
* Evt 101, a selective NR2B receptor antagonist. This compound was tested as a potentially fast-acting antidepressant.<ref name="urlThe Effects of a Novel NMDA NR2B-Subtype Selective Antagonist, EVT 101, on Brain Function - Full Text View - ClinicalTrials.gov">{{cite web | url = http://clinicaltrials.gov/ct2/show/NCT00526968 | title = The Effects of a Novel NMDA NR2B-Subtype Selective Antagonist, EVT 101, on Brain Function  | author = | authorlink = | coauthors = | date = 2008-02-14 | format = | work = NCT00526968  | publisher = ClinicalTrials.gov | pages = | archiveurl = | archivedate = | quote = | accessdate = 2010-08-19 }}</ref> In 2011 it was voluntarily withdrawn from a Phase II clinical study in treatment-resistant depression due to an unsatisfactory toxicity profile.<ref name="urlPhase II study with NR2B sub-type selective NMDA antagonist in treatment-resistant depression voluntarily terminated.">{{cite web | url = http://www.evotec.com/archive/en/Press-releases/2011/Phase-II-study-with-NR2B-sub-type-selective-NMDA-antagonist-in-treatment-resistant-depression-voluntarily-terminated/2157/1 | title = Phase II study with NR2B sub-type selective NMDA antagonist in treatment-resistant depression voluntarily terminated  | author = | authorlink = | coauthors = | date = 2011-05-18 | format = | work = | publisher = evotec.com | pages = | archiveurl = | archivedate = | quote = | accessdate = 2015-08-24 }}</ref>
* [[Felbamate]], an anticonvulsant that is also a [[positive allosteric modulator]] for the [[GABAA receptor|GABA<sub>A</sub> receptor]]
* [[Ro-25-6981]] (also known as MI-4), a selective NR2B receptor antagonist
* [[Traxoprodil]], a selective NR2B receptor antagonist
 
== Interactions ==
 
GRIN2B has been shown to [[Protein-protein interaction|interact]] with:
{{div col|colwidth=20em}}
* [[Actinin, alpha 2]],<ref name = "pmid9009191">{{cite journal | vauthors = Wyszynski M, Lin J, Rao A, Nigh E, Beggs AH, Craig AM, Sheng M | title = Competitive binding of alpha-actinin and calmodulin to the NMDA receptor | journal = Nature | volume = 385 | issue = 6615 | pages = 439–42 | date = January 1997 | pmid = 9009191 | doi = 10.1038/385439a0 }}</ref>
* [[DLG2]],<ref name = pmid11997254/><ref name = pmid9278515/>
* [[DLG3]],<ref name = pmid11997254/><ref name = pmid9278515/><ref name = pmid12738960/><ref name = pmid11937501/>
* [[DLG4]],<ref name = "pmid11997254">{{cite journal | vauthors = Inanobe A, Fujita A, Ito M, Tomoike H, Inageda K, Kurachi Y | title = Inward rectifier K+ channel Kir2.3 is localized at the postsynaptic membrane of excitatory synapses | journal = American Journal of Physiology. Cell Physiology | volume = 282 | issue = 6 | pages = C1396-403 | date = June 2002 | pmid = 11997254 | doi = 10.1152/ajpcell.00615.2001 }}</ref><ref name = "pmid9278515">{{cite journal | vauthors = Irie M, Hata Y, Takeuchi M, Ichtchenko K, Toyoda A, Hirao K, Takai Y, Rosahl TW, Südhof TC | title = Binding of neuroligins to PSD-95 | journal = Science | volume = 277 | issue = 5331 | pages = 1511–5 | date = September 1997 | pmid = 9278515 | doi = 10.1126/science.277.5331.1511 }}</ref><ref name = "pmid12738960">{{cite journal | vauthors = Sans N, Prybylowski K, Petralia RS, Chang K, Wang YX, Racca C, Vicini S, Wenthold RJ | title = NMDA receptor trafficking through an interaction between PDZ proteins and the exocyst complex | journal = Nature Cell Biology | volume = 5 | issue = 6 | pages = 520–30 | date = June 2003 | pmid = 12738960 | doi = 10.1038/ncb990 }}</ref><ref name = "pmid11937501">{{cite journal | vauthors = Lim IA, Hall DD, Hell JW | title = Selectivity and promiscuity of the first and second PDZ domains of PSD-95 and synapse-associated protein 102 | journal = The Journal of Biological Chemistry | volume = 277 | issue = 24 | pages = 21697–711 | date = June 2002 | pmid = 11937501 | doi = 10.1074/jbc.M112339200 }}</ref><ref name = "pmid9581762">{{cite journal | vauthors = Niethammer M, Valtschanoff JG, Kapoor TM, Allison DW, Weinberg RJ, Craig AM, Sheng M | title = CRIPT, a novel postsynaptic protein that binds to the third PDZ domain of PSD-95/SAP90 | journal = Neuron | volume = 20 | issue = 4 | pages = 693–707 | date = April 1998 | pmid = 9581762 | doi = 10.1016/s0896-6273(00)81009-0 }}</ref><ref name = "pmid7569905">{{cite journal | vauthors = Kornau HC, Schenker LT, Kennedy MB, Seeburg PH |authorlink3=Mary B. Kennedy| title = Domain interaction between NMDA receptor subunits and the postsynaptic density protein PSD-95 | journal = Science | volume = 269 | issue = 5231 | pages = 1737–40 | date = September 1995 | pmid = 7569905 | doi = 10.1126/science.7569905 }}</ref>
* [[EXOC4]],<ref name = pmid12738960/>
* [[LIN7B]],<ref name = "pmid10341223">{{cite journal | vauthors = Jo K, Derin R, Li M, Bredt DS | title = Characterization of MALS/Velis-1, -2, and -3: a family of mammalian LIN-7 homologs enriched at brain synapses in association with the postsynaptic density-95/NMDA receptor postsynaptic complex | journal = The Journal of Neuroscience | volume = 19 | issue = 11 | pages = 4189–99 | date = June 1999 | pmid = 10341223 | doi =  }}</ref>  and
* [[RICS (gene)|RICS]].<ref name = "pmid12857875">{{cite journal | vauthors = Nakazawa T, Watabe AM, Tezuka T, Yoshida Y, Yokoyama K, Umemori H, Inoue A, Okabe S, Manabe T, Yamamoto T | title = p250GAP, a novel brain-enriched GTPase-activating protein for Rho family GTPases, is involved in the N-methyl-d-aspartate receptor signaling | journal = Molecular Biology of the Cell | volume = 14 | issue = 7 | pages = 2921–34 | date = July 2003 | pmid = 12857875 | pmc = 165687 | doi = 10.1091/mbc.E02-09-0623 }}</ref>
{{Div col end}}
 
== See also ==
* [[NMDA receptor]]
* [[NMDA receptor]]
* [[Glutamate receptor]]
* [[Glutamate receptor]]


==References==
== References ==
{{reflist|2}}
{{reflist}}
==Further reading==
 
== Further reading ==
{{refbegin | 2}}
{{refbegin | 2}}
{{PBB_Further_reading
* {{cite journal | vauthors = Schröder HC, Perovic S, Kavsan V, Ushijima H, Müller WE | title = Mechanisms of prionSc- and HIV-1 gp120 induced neuronal cell death | journal = Neurotoxicology | volume = 19 | issue = 4–5 | pages = 683–8 | year = 1998 | pmid = 9745929 | doi =  }}
| citations =
* {{cite journal | vauthors = Nagy J | title = The NR2B subtype of NMDA receptor: a potential target for the treatment of alcohol dependence | journal = Current Drug Targets. CNS and Neurological Disorders | volume = 3 | issue = 3 | pages = 169–79 | date = June 2004 | pmid = 15180478 | doi = 10.2174/1568007043337409 }}
*{{cite journal | author=Schröder HC, Perovic S, Kavsan V, ''et al.'' |title=Mechanisms of prionSc- and HIV-1 gp120 induced neuronal cell death. |journal=Neurotoxicology |volume=19 |issue= 4-5 |pages= 683-8 |year= 1998 |pmid= 9745929 |doi=  }}
* {{cite journal | vauthors = King JE, Eugenin EA, Buckner CM, Berman JW | title = HIV tat and neurotoxicity | journal = Microbes and Infection | volume = 8 | issue = 5 | pages = 1347–57 | date = April 2006 | pmid = 16697675 | doi = 10.1016/j.micinf.2005.11.014 }}
*{{cite journal | author=Nagy J |title=The NR2B subtype of NMDA receptor: a potential target for the treatment of alcohol dependence. |journal=Current drug targets. CNS and neurological disorders |volume=3 |issue= 3 |pages= 169-79 |year= 2004 |pmid= 15180478 |doi= }}
* {{cite journal | vauthors = Kornau HC, Schenker LT, Kennedy MB, Seeburg PH | title = Domain interaction between NMDA receptor subunits and the postsynaptic density protein PSD-95 | journal = Science | volume = 269 | issue = 5231 | pages = 1737–40 | date = September 1995 | pmid = 7569905 | doi = 10.1126/science.7569905 }}
*{{cite journal | author=King JE, Eugenin EA, Buckner CM, Berman JW |title=HIV tat and neurotoxicity. |journal=Microbes Infect. |volume=8 |issue= 5 |pages= 1347-57 |year= 2006 |pmid= 16697675 |doi= 10.1016/j.micinf.2005.11.014 }}
* {{cite journal | vauthors = Magnuson DS, Knudsen BE, Geiger JD, Brownstone RM, Nath A | title = Human immunodeficiency virus type 1 tat activates non-N-methyl-D-aspartate excitatory amino acid receptors and causes neurotoxicity | journal = Annals of Neurology | volume = 37 | issue = 3 | pages = 373–80 | date = March 1995 | pmid = 7695237 | doi = 10.1002/ana.410370314 }}
*{{cite journal | author=Monyer H, Sprengel R, Schoepfer R, ''et al.'' |title=Heteromeric NMDA receptors: molecular and functional distinction of subtypes. |journal=Science |volume=256 |issue= 5060 |pages= 1217-21 |year= 1992 |pmid= 1350383 |doi= }}
* {{cite journal | vauthors = Mandich P, Schito AM, Bellone E, Antonacci R, Finelli P, Rocchi M, Ajmar F | title = Mapping of the human NMDAR2B receptor subunit gene (GRIN2B) to chromosome 12p12 | journal = Genomics | volume = 22 | issue = 1 | pages = 216–8 | date = July 1994 | pmid = 7959773 | doi = 10.1006/geno.1994.1366 }}
*{{cite journal | author=Kornau HC, Schenker LT, Kennedy MB, Seeburg PH |title=Domain interaction between NMDA receptor subunits and the postsynaptic density protein PSD-95. |journal=Science |volume=269 |issue= 5231 |pages= 1737-40 |year= 1995 |pmid= 7569905 |doi= }}
* {{cite journal | vauthors = Adams SL, Foldes RL, Kamboj RK | title = Human N-methyl-D-aspartate receptor modulatory subunit hNR3: cloning and sequencing of the cDNA and primary structure of the protein | journal = Biochimica et Biophysica Acta | volume = 1260 | issue = 1 | pages = 105–8 | date = January 1995 | pmid = 7999784 | doi = 10.1016/0167-4781(94)00189-a }}
*{{cite journal | author=Magnuson DS, Knudsen BE, Geiger JD, ''et al.'' |title=Human immunodeficiency virus type 1 tat activates non-N-methyl-D-aspartate excitatory amino acid receptors and causes neurotoxicity. |journal=Ann. Neurol. |volume=37 |issue= 3 |pages= 373-80 |year= 1995 |pmid= 7695237 |doi= 10.1002/ana.410370314 }}
* {{cite journal | vauthors = Sheng M, Cummings J, Roldan LA, Jan YN, Jan LY | title = Changing subunit composition of heteromeric NMDA receptors during development of rat cortex | journal = Nature | volume = 368 | issue = 6467 | pages = 144–7 | date = March 1994 | pmid = 8139656 | doi = 10.1038/368144a0 }}
*{{cite journal | author=Mandich P, Schito AM, Bellone E, ''et al.'' |title=Mapping of the human NMDAR2B receptor subunit gene (GRIN2B) to chromosome 12p12. |journal=Genomics |volume=22 |issue= 1 |pages= 216-8 |year= 1994 |pmid= 7959773 |doi= 10.1006/geno.1994.1366 }}
* {{cite journal | vauthors = Roche KW, Raymond LA, Blackstone C, Huganir RL | title = Transmembrane topology of the glutamate receptor subunit GluR6 | journal = The Journal of Biological Chemistry | volume = 269 | issue = 16 | pages = 11679–82 | date = April 1994 | pmid = 8163463 | doi = }}
*{{cite journal | author=Adams SL, Foldes RL, Kamboj RK |title=Human N-methyl-D-aspartate receptor modulatory subunit hNR3: cloning and sequencing of the cDNA and primary structure of the protein. |journal=Biochim. Biophys. Acta |volume=1260 |issue= 1 |pages= 105-8 |year= 1995 |pmid= 7999784 |doi= }}
* {{cite journal | vauthors = Lannuzel A, Lledo PM, Lamghitnia HO, Vincent JD, Tardieu M | title = HIV-1 envelope proteins gp120 and gp160 potentiate NMDA-induced [Ca2+]i increase, alter [Ca2+]i homeostasis and induce neurotoxicity in human embryonic neurons | journal = The European Journal of Neuroscience | volume = 7 | issue = 11 | pages = 2285–93 | date = November 1995 | pmid = 8563977 | doi = 10.1111/j.1460-9568.1995.tb00649.x }}
*{{cite journal | author=Sheng M, Cummings J, Roldan LA, ''et al.'' |title=Changing subunit composition of heteromeric NMDA receptors during development of rat cortex. |journal=Nature |volume=368 |issue= 6467 |pages= 144-7 |year= 1994 |pmid= 8139656 |doi= 10.1038/368144a0 }}
* {{cite journal | vauthors = Corasaniti MT, Melino G, Navarra M, Garaci E, Finazzi-Agrò A, Nisticò G | title = Death of cultured human neuroblastoma cells induced by HIV-1 gp120 is prevented by NMDA receptor antagonists and inhibitors of nitric oxide and cyclooxygenase | journal = Neurodegeneration | volume = 4 | issue = 3 | pages = 315–21 | date = September 1995 | pmid = 8581564 | doi = 10.1016/1055-8330(95)90021-7 }}
*{{cite journal | author=Roche KW, Raymond LA, Blackstone C, Huganir RL |title=Transmembrane topology of the glutamate receptor subunit GluR6. |journal=J. Biol. Chem. |volume=269 |issue= 16 |pages= 11679-82 |year= 1994 |pmid= 8163463 |doi= }}
* {{cite journal | vauthors = Niethammer M, Kim E, Sheng M | title = Interaction between the C terminus of NMDA receptor subunits and multiple members of the PSD-95 family of membrane-associated guanylate kinases | journal = The Journal of Neuroscience | volume = 16 | issue = 7 | pages = 2157–63 | date = April 1996 | pmid = 8601796 | doi =  }}
*{{cite journal | author=Lannuzel A, Lledo PM, Lamghitnia HO, ''et al.'' |title=HIV-1 envelope proteins gp120 and gp160 potentiate NMDA-induced [Ca2+]i increase, alter [Ca2+]i homeostasis and induce neurotoxicity in human embryonic neurons. |journal=Eur. J. Neurosci. |volume=7 |issue= 11 |pages= 2285-93 |year= 1996 |pmid= 8563977 |doi= }}
* {{cite journal | vauthors = Pittaluga A, Pattarini R, Severi P, Raiteri M | title = Human brain N-methyl-D-aspartate receptors regulating noradrenaline release are positively modulated by HIV-1 coat protein gp120 | journal = AIDS | volume = 10 | issue = 5 | pages = 463–8 | date = May 1996 | pmid = 8724036 | doi = 10.1097/00002030-199605000-00003 }}
*{{cite journal | author=Corasaniti MT, Melino G, Navarra M, ''et al.'' |title=Death of cultured human neuroblastoma cells induced by HIV-1 gp120 is prevented by NMDA receptor antagonists and inhibitors of nitric oxide and cyclooxygenase. |journal=Neurodegeneration : a journal for neurodegenerative disorders, neuroprotection, and neuroregeneration |volume=4 |issue= 3 |pages= 315-21 |year= 1996 |pmid= 8581564 |doi=  }}
* {{cite journal | vauthors = Hess SD, Daggett LP, Crona J, Deal C, Lu CC, Urrutia A, Chavez-Noriega L, Ellis SB, Johnson EC, Veliçelebi G | title = Cloning and functional characterization of human heteromeric N-methyl-D-aspartate receptors | journal = The Journal of Pharmacology and Experimental Therapeutics | volume = 278 | issue = 2 | pages = 808–16 | date = August 1996 | pmid = 8768735 | doi =  }}
*{{cite journal | author=Niethammer M, Kim E, Sheng M |title=Interaction between the C terminus of NMDA receptor subunits and multiple members of the PSD-95 family of membrane-associated guanylate kinases. |journal=J. Neurosci. |volume=16 |issue= 7 |pages= 2157-63 |year= 1996 |pmid= 8601796 |doi= }}
* {{cite journal | vauthors = Müller BM, Kistner U, Kindler S, Chung WJ, Kuhlendahl S, Fenster SD, Lau LF, Veh RW, Huganir RL, Gundelfinger ED, Garner CC | title = SAP102, a novel postsynaptic protein that interacts with NMDA receptor complexes in vivo | journal = Neuron | volume = 17 | issue = 2 | pages = 255–65 | date = August 1996 | pmid = 8780649 | doi = 10.1016/S0896-6273(00)80157-9 }}
*{{cite journal | author=Pittaluga A, Pattarini R, Severi P, Raiteri M |title=Human brain N-methyl-D-aspartate receptors regulating noradrenaline release are positively modulated by HIV-1 coat protein gp120. |journal=AIDS |volume=10 |issue= 5 |pages= 463-8 |year= 1996 |pmid= 8724036 |doi=  }}
* {{cite journal | vauthors = Wu P, Price P, Du B, Hatch WC, Terwilliger EF | title = Direct cytotoxicity of HIV-1 envelope protein gp120 on human NT neurons | journal = NeuroReport | volume = 7 | issue = 5 | pages = 1045–9 | date = April 1996 | pmid = 8804048 | doi = 10.1097/00001756-199604100-00018 }}
*{{cite journal | author=Hess SD, Daggett LP, Crona J, ''et al.'' |title=Cloning and functional characterization of human heteromeric N-methyl-D-aspartate receptors. |journal=J. Pharmacol. Exp. Ther. |volume=278 |issue= 2 |pages= 808-16 |year= 1996 |pmid= 8768735 |doi= }}
* {{cite journal | vauthors = Bennett BA, Rusyniak DE, Hollingsworth CK | title = HIV-1 gp120-induced neurotoxicity to midbrain dopamine cultures | journal = Brain Research | volume = 705 | issue = 1–2 | pages = 168–76 | date = December 1995 | pmid = 8821747 | doi = 10.1016/0006-8993(95)01166-8 }}
*{{cite journal | author=Müller BM, Kistner U, Kindler S, ''et al.'' |title=SAP102, a novel postsynaptic protein that interacts with NMDA receptor complexes in vivo. |journal=Neuron |volume=17 |issue= 2 |pages= 255-65 |year= 1996 |pmid= 8780649 |doi= }}
* {{cite journal | vauthors = Toggas SM, Masliah E, Mucke L | title = Prevention of HIV-1 gp120-induced neuronal damage in the central nervous system of transgenic mice by the NMDA receptor antagonist memantine | journal = Brain Research | volume = 706 | issue = 2 | pages = 303–7 | date = January 1996 | pmid = 8822372 | doi = 10.1016/0006-8993(95)01197-8 }}
*{{cite journal | author=Wu P, Price P, Du B, ''et al.'' |title=Direct cytotoxicity of HIV-1 envelope protein gp120 on human NT neurons. |journal=Neuroreport |volume=7 |issue= 5 |pages= 1045-9 |year= 1996 |pmid= 8804048 |doi= }}
* {{cite journal | vauthors = Dreyer EB, Lipton SA | title = The coat protein gp120 of HIV-1 inhibits astrocyte uptake of excitatory amino acids via macrophage arachidonic acid | journal = The European Journal of Neuroscience | volume = 7 | issue = 12 | pages = 2502–7 | date = December 1995 | pmid = 8845955 | doi = 10.1111/j.1460-9568.1995.tb01048.x }}
*{{cite journal | author=Bennett BA, Rusyniak DE, Hollingsworth CK |title=HIV-1 gp120-induced neurotoxicity to midbrain dopamine cultures. |journal=Brain Res. |volume=705 |issue= 1-2 |pages= 168-76 |year= 1996 |pmid= 8821747 |doi= }}
*{{cite journal | author=Toggas SM, Masliah E, Mucke L |title=Prevention of HIV-1 gp120-induced neuronal damage in the central nervous system of transgenic mice by the NMDA receptor antagonist memantine. |journal=Brain Res. |volume=706 |issue= 2 |pages= 303-7 |year= 1996 |pmid= 8822372 |doi= }}
*{{cite journal  | author=Dreyer EB, Lipton SA |title=The coat protein gp120 of HIV-1 inhibits astrocyte uptake of excitatory amino acids via macrophage arachidonic acid. |journal=Eur. J. Neurosci. |volume=7 |issue= 12 |pages= 2502-7 |year= 1996 |pmid= 8845955 |doi=  }}
}}
{{refend}}
{{refend}}


{{membrane-protein-stub}}
{{NLM content}}
{{NLM content}}
{{Ligand-gated ion channels}}
{{Ligand-gated ion channels}}
[[Category:Ion channels]]
 
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[[Category:Ionotropic glutamate receptors]]

Latest revision as of 08:49, 1 June 2018

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Orthologs
SpeciesHumanMouse
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Glutamate [NMDA] receptor subunit epsilon-2, also known as N-methyl D-aspartate receptor subtype 2B (NMDAR2B or NR2B), is a protein that in humans is encoded by the GRIN2B gene.[1]

NMDA receptors

N-methyl-D-aspartate (NMDA) receptors are a class of ionotropic glutamate receptors. The NMDA receptor channel has been shown to be involved in long-term potentiation, an activity-dependent increase in the efficiency of synaptic transmission thought to underlie certain kinds of memory and learning. NMDA receptor channels are heterotetramers composed of two molecules of the key receptor subunit NMDAR1 (GRIN1) and two drawn from one or more of the four NMDAR2 subunits: NMDAR2A (GRIN2A), NMDAR2B (GRIN2B), NMDAR2C (GRIN2C), and NMDAR2D (GRIN2D). The NR2 subunit acts as the agonist binding site for glutamate, one of the predominant excitatory neurotransmitter receptors in the mammalian brain.[2]

Function

NR2B has been associated with age- and visual-experience-dependent plasticity in the neocortex of rats, where an increased NR2B/NR2A ratio correlates directly with the stronger excitatory LTP in young animals. This is thought to contribute to experience-dependent refinement of developing cortical circuits.[3]

Both mice and rats that were engineered to over-express GRIN2B in their brains have increased mental ability.[4][5]

Ligands

  • Besonprodil
  • CERC-301, a selective NR2B receptor antagonist
  • Eliprodil
  • Evt 101, a selective NR2B receptor antagonist. This compound was tested as a potentially fast-acting antidepressant.[6] In 2011 it was voluntarily withdrawn from a Phase II clinical study in treatment-resistant depression due to an unsatisfactory toxicity profile.[7]
  • Felbamate, an anticonvulsant that is also a positive allosteric modulator for the GABAA receptor
  • Ro-25-6981 (also known as MI-4), a selective NR2B receptor antagonist
  • Traxoprodil, a selective NR2B receptor antagonist

Interactions

GRIN2B has been shown to interact with:

See also

References

  1. ↑ Monyer H, Sprengel R, Schoepfer R, Herb A, Higuchi M, Lomeli H, Burnashev N, Sakmann B, Seeburg PH (May 1992). "Heteromeric NMDA receptors: molecular and functional distinction of subtypes". Science. 256 (5060): 1217–21. doi:10.1126/science.256.5060.1217. PMID 1350383.
  2. ↑ "Entrez Gene: GRIN2B glutamate receptor, ionotropic, N-methyl D-aspartate 2B".
  3. ↑ Yoshimura Y, Ohmura T, Komatsu Y (July 2003). "Two forms of synaptic plasticity with distinct dependence on age, experience, and NMDA receptor subtype in rat visual cortex". The Journal of Neuroscience. 23 (16): 6557–66. PMID 12878697.
  4. ↑ Tang YP, Shimizu E, Dube GR, Rampon C, Kerchner GA, Zhuo M, Liu G, Tsien JZ (September 1999). "Genetic enhancement of learning and memory in mice". Nature. 401 (6748): 63–9. doi:10.1038/43432. PMID 10485705.
  5. ↑ Wang D, Cui Z, Zeng Q, Kuang H, Wang LP, Tsien JZ, Cao X (October 2009). "Genetic enhancement of memory and long-term potentiation but not CA1 long-term depression in NR2B transgenic rats". PLOS ONE. 4 (10): e7486. doi:10.1371/journal.pone.0007486. PMC 2759522. PMID 19838302.
  6. ↑ "The Effects of a Novel NMDA NR2B-Subtype Selective Antagonist, EVT 101, on Brain Function". NCT00526968. ClinicalTrials.gov. 2008-02-14. Retrieved 2010-08-19.
  7. ↑ "Phase II study with NR2B sub-type selective NMDA antagonist in treatment-resistant depression voluntarily terminated". evotec.com. 2011-05-18. Retrieved 2015-08-24.
  8. ↑ Wyszynski M, Lin J, Rao A, Nigh E, Beggs AH, Craig AM, Sheng M (January 1997). "Competitive binding of alpha-actinin and calmodulin to the NMDA receptor". Nature. 385 (6615): 439–42. doi:10.1038/385439a0. PMID 9009191.
  9. ↑ 9.0 9.1 9.2 Inanobe A, Fujita A, Ito M, Tomoike H, Inageda K, Kurachi Y (June 2002). "Inward rectifier K+ channel Kir2.3 is localized at the postsynaptic membrane of excitatory synapses". American Journal of Physiology. Cell Physiology. 282 (6): C1396–403. doi:10.1152/ajpcell.00615.2001. PMID 11997254.
  10. ↑ 10.0 10.1 10.2 Irie M, Hata Y, Takeuchi M, Ichtchenko K, Toyoda A, Hirao K, Takai Y, Rosahl TW, Südhof TC (September 1997). "Binding of neuroligins to PSD-95". Science. 277 (5331): 1511–5. doi:10.1126/science.277.5331.1511. PMID 9278515.
  11. ↑ 11.0 11.1 11.2 Sans N, Prybylowski K, Petralia RS, Chang K, Wang YX, Racca C, Vicini S, Wenthold RJ (June 2003). "NMDA receptor trafficking through an interaction between PDZ proteins and the exocyst complex". Nature Cell Biology. 5 (6): 520–30. doi:10.1038/ncb990. PMID 12738960.
  12. ↑ 12.0 12.1 Lim IA, Hall DD, Hell JW (June 2002). "Selectivity and promiscuity of the first and second PDZ domains of PSD-95 and synapse-associated protein 102". The Journal of Biological Chemistry. 277 (24): 21697–711. doi:10.1074/jbc.M112339200. PMID 11937501.
  13. ↑ Niethammer M, Valtschanoff JG, Kapoor TM, Allison DW, Weinberg RJ, Craig AM, Sheng M (April 1998). "CRIPT, a novel postsynaptic protein that binds to the third PDZ domain of PSD-95/SAP90". Neuron. 20 (4): 693–707. doi:10.1016/s0896-6273(00)81009-0. PMID 9581762.
  14. ↑ Kornau HC, Schenker LT, Kennedy MB, Seeburg PH (September 1995). "Domain interaction between NMDA receptor subunits and the postsynaptic density protein PSD-95". Science. 269 (5231): 1737–40. doi:10.1126/science.7569905. PMID 7569905.
  15. ↑ Jo K, Derin R, Li M, Bredt DS (June 1999). "Characterization of MALS/Velis-1, -2, and -3: a family of mammalian LIN-7 homologs enriched at brain synapses in association with the postsynaptic density-95/NMDA receptor postsynaptic complex". The Journal of Neuroscience. 19 (11): 4189–99. PMID 10341223.
  16. ↑ Nakazawa T, Watabe AM, Tezuka T, Yoshida Y, Yokoyama K, Umemori H, Inoue A, Okabe S, Manabe T, Yamamoto T (July 2003). "p250GAP, a novel brain-enriched GTPase-activating protein for Rho family GTPases, is involved in the N-methyl-d-aspartate receptor signaling". Molecular Biology of the Cell. 14 (7): 2921–34. doi:10.1091/mbc.E02-09-0623. PMC 165687. PMID 12857875.

Further reading

  • Schröder HC, Perovic S, Kavsan V, Ushijima H, Müller WE (1998). "Mechanisms of prionSc- and HIV-1 gp120 induced neuronal cell death". Neurotoxicology. 19 (4–5): 683–8. PMID 9745929.
  • Nagy J (June 2004). "The NR2B subtype of NMDA receptor: a potential target for the treatment of alcohol dependence". Current Drug Targets. CNS and Neurological Disorders. 3 (3): 169–79. doi:10.2174/1568007043337409. PMID 15180478.
  • King JE, Eugenin EA, Buckner CM, Berman JW (April 2006). "HIV tat and neurotoxicity". Microbes and Infection. 8 (5): 1347–57. doi:10.1016/j.micinf.2005.11.014. PMID 16697675.
  • Kornau HC, Schenker LT, Kennedy MB, Seeburg PH (September 1995). "Domain interaction between NMDA receptor subunits and the postsynaptic density protein PSD-95". Science. 269 (5231): 1737–40. doi:10.1126/science.7569905. PMID 7569905.
  • Magnuson DS, Knudsen BE, Geiger JD, Brownstone RM, Nath A (March 1995). "Human immunodeficiency virus type 1 tat activates non-N-methyl-D-aspartate excitatory amino acid receptors and causes neurotoxicity". Annals of Neurology. 37 (3): 373–80. doi:10.1002/ana.410370314. PMID 7695237.
  • Mandich P, Schito AM, Bellone E, Antonacci R, Finelli P, Rocchi M, Ajmar F (July 1994). "Mapping of the human NMDAR2B receptor subunit gene (GRIN2B) to chromosome 12p12". Genomics. 22 (1): 216–8. doi:10.1006/geno.1994.1366. PMID 7959773.
  • Adams SL, Foldes RL, Kamboj RK (January 1995). "Human N-methyl-D-aspartate receptor modulatory subunit hNR3: cloning and sequencing of the cDNA and primary structure of the protein". Biochimica et Biophysica Acta. 1260 (1): 105–8. doi:10.1016/0167-4781(94)00189-a. PMID 7999784.
  • Sheng M, Cummings J, Roldan LA, Jan YN, Jan LY (March 1994). "Changing subunit composition of heteromeric NMDA receptors during development of rat cortex". Nature. 368 (6467): 144–7. doi:10.1038/368144a0. PMID 8139656.
  • Roche KW, Raymond LA, Blackstone C, Huganir RL (April 1994). "Transmembrane topology of the glutamate receptor subunit GluR6". The Journal of Biological Chemistry. 269 (16): 11679–82. PMID 8163463.
  • Lannuzel A, Lledo PM, Lamghitnia HO, Vincent JD, Tardieu M (November 1995). "HIV-1 envelope proteins gp120 and gp160 potentiate NMDA-induced [Ca2+]i increase, alter [Ca2+]i homeostasis and induce neurotoxicity in human embryonic neurons". The European Journal of Neuroscience. 7 (11): 2285–93. doi:10.1111/j.1460-9568.1995.tb00649.x. PMID 8563977.
  • Corasaniti MT, Melino G, Navarra M, Garaci E, Finazzi-Agrò A, Nisticò G (September 1995). "Death of cultured human neuroblastoma cells induced by HIV-1 gp120 is prevented by NMDA receptor antagonists and inhibitors of nitric oxide and cyclooxygenase". Neurodegeneration. 4 (3): 315–21. doi:10.1016/1055-8330(95)90021-7. PMID 8581564.
  • Niethammer M, Kim E, Sheng M (April 1996). "Interaction between the C terminus of NMDA receptor subunits and multiple members of the PSD-95 family of membrane-associated guanylate kinases". The Journal of Neuroscience. 16 (7): 2157–63. PMID 8601796.
  • Pittaluga A, Pattarini R, Severi P, Raiteri M (May 1996). "Human brain N-methyl-D-aspartate receptors regulating noradrenaline release are positively modulated by HIV-1 coat protein gp120". AIDS. 10 (5): 463–8. doi:10.1097/00002030-199605000-00003. PMID 8724036.
  • Hess SD, Daggett LP, Crona J, Deal C, Lu CC, Urrutia A, Chavez-Noriega L, Ellis SB, Johnson EC, Veliçelebi G (August 1996). "Cloning and functional characterization of human heteromeric N-methyl-D-aspartate receptors". The Journal of Pharmacology and Experimental Therapeutics. 278 (2): 808–16. PMID 8768735.
  • Müller BM, Kistner U, Kindler S, Chung WJ, Kuhlendahl S, Fenster SD, Lau LF, Veh RW, Huganir RL, Gundelfinger ED, Garner CC (August 1996). "SAP102, a novel postsynaptic protein that interacts with NMDA receptor complexes in vivo". Neuron. 17 (2): 255–65. doi:10.1016/S0896-6273(00)80157-9. PMID 8780649.
  • Wu P, Price P, Du B, Hatch WC, Terwilliger EF (April 1996). "Direct cytotoxicity of HIV-1 envelope protein gp120 on human NT neurons". NeuroReport. 7 (5): 1045–9. doi:10.1097/00001756-199604100-00018. PMID 8804048.
  • Bennett BA, Rusyniak DE, Hollingsworth CK (December 1995). "HIV-1 gp120-induced neurotoxicity to midbrain dopamine cultures". Brain Research. 705 (1–2): 168–76. doi:10.1016/0006-8993(95)01166-8. PMID 8821747.
  • Toggas SM, Masliah E, Mucke L (January 1996). "Prevention of HIV-1 gp120-induced neuronal damage in the central nervous system of transgenic mice by the NMDA receptor antagonist memantine". Brain Research. 706 (2): 303–7. doi:10.1016/0006-8993(95)01197-8. PMID 8822372.
  • Dreyer EB, Lipton SA (December 1995). "The coat protein gp120 of HIV-1 inhibits astrocyte uptake of excitatory amino acids via macrophage arachidonic acid". The European Journal of Neuroscience. 7 (12): 2502–7. doi:10.1111/j.1460-9568.1995.tb01048.x. PMID 8845955.

This article incorporates text from the United States National Library of Medicine, which is in the public domain.

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