Role of actin cytoskeleton in the organization and function of ionotropic glutamate receptors

Current Research in Structural Biology - Tập 3 - Trang 277-289 - 2021
Priyanka Dutta1, Pratibha Bharti1, Janesh Kumar1, Sankar Maiti2
1National Centre for Cell Science, Pune, Maharashtra, 411007, India
2Indian Institute of Science Education and Research, Kolkata, 741246, India

Tài liệu tham khảo

Alberts, 2002 Andreas, 2018, Glutamatergic signaling in the central nervous system: ionotropic and metabotropic receptors in concert, Neuron, 98, 1080, 10.1016/j.neuron.2018.05.018 Arber, 1998, Regulation of actin dynamics through phosphorylation of cofilin by LIM- kinase, Nature, 393, 805, 10.1038/31729 Bach, 2000, The LIM domain: regulation by association, Mech. Dev., 91, 5, 10.1016/S0925-4773(99)00314-7 Baines, 2014, The Protein 4.1 family: hub proteins in animals for organizing membrane proteins, Biochim. Biophys. Acta, 1838, 605, 10.1016/j.bbamem.2013.05.030 Balasuriya, 2013, α-Amino-3-hydroxy-5-methyl-4-isoxazole propionic acid (AMPA) and N-methyl-D-aspartate (NMDA) receptors adopt different subunit arrangements, J. Biol. Chem., 288, 21987, 10.1074/jbc.M113.469205 Bamburg, 1999, Proteins of the ADF/cofilin family: essential regulators of actin dynamics. 1999. 15:185–230, Annu. Rev. Cell Dev. Biol., 15, 185, 10.1146/annurev.cellbio.15.1.185 Bamji, 2005, Cadherins: actin with the cytoskeleton to form synapses, Neuron, 47, 175, 10.1016/j.neuron.2005.06.024 Bárány, 2001, Exchange of the actin-bound nucleotide in intact arterial smooth muscle, J. Biol. Chem., 276, 48398, 10.1074/jbc.M106227200 Basu, 2018, The role of actin cytoskeleton in dendritic spines in the maintenance of long-term memory, Front. Mol. Neurosci., 11 Baucum, 2017, Proteomic analysis of postsynaptic protein complexes underlying neuronal plasticity, ACS Chem. Neurosci., 8, 689, 10.1021/acschemneuro.7b00008 Behuet, 2019, Developmental changes of glutamate and GABA receptor densities in wistar rats, Front. Neuroanat., 13, 10.3389/fnana.2019.00100 Béïque, 2006, Synapse-specific regulation of AMPA receptor function by PSD-95, Proc. Natl. Acad. Sci. Unit. States Am., 103, 19535, 10.1073/pnas.0608492103 Ben Zablah, 2020, The role of ADF/cofilin in synaptic physiology and alzheimer's disease, Front. Cell Dev. Biol., 8, 1 Bennett, 2011, A model of NMDA receptor control of F-actin treadmilling in synaptic spines and their growth, Bull. Math. Biol., 73, 2109, 10.1007/s11538-010-9614-4 Benson, 1998, N-cadherin redistribution during synaptogenesis in hippocampal neurons, J. Neurosci., 18, 6892, 10.1523/JNEUROSCI.18-17-06892.1998 Biederer, 2008, Signaling by synaptogenetic molecules, Curr. Opin. Neurobiol., 18, 261, 10.1016/j.conb.2008.07.014 Bisaria, 2020, Membrane-proximal F-actin restricts local membrane protrusions and directs cell migration, Science, 368, 1205, 10.1126/science.aay7794 Bisaz, 2014, The neurobiological bases of memory formation: from physiological conditions to psychopathology, Psychopathology, 47, 347, 10.1159/000363702 Bissen, 2019 Borovac, 2018, Regulation of actin dynamics during structural plasticity of dendritic spines: signaling messengers and actin-binding proteins, Mol. Cell. Neurosci., 91, 122, 10.1016/j.mcn.2018.07.001 Bosch, 2014, Structural and molecular remodeling of dendritic spine substructures during long-term potentiation, Neuron, 82, 444, 10.1016/j.neuron.2014.03.021 Bosch, 2014, Structural and molecular remodeling of dendritic spine substructures during long-term potentiation, Neuron, 82, 444, 10.1016/j.neuron.2014.03.021 Brandt, 2007, Get to grips: steering local actin dynamics with IQGAPs, EMBO Rep., 8, 1019, 10.1038/sj.embor.7401089 Breese, 1993, Glutamate receptor subtype expression in human postmortem brain, J. Mol. Neurosci., 4, 263, 10.1007/BF02821558 Bu, 2015, Myosin IIb-dependent regulation of actin dynamics is required for N-Methyl-D-aspartate receptor trafficking during synaptic plasticity, J. Biol. Chem., 290, 25395, 10.1074/jbc.M115.644229 Burada, 2020, Emerging insights into the structure and function of ionotropic glutamate delta receptors., Br J Pharmacol., 10.1111/bph.15313 Burada, 2020, Cryo-EM structures of the ionotropic glutamate receptor GluD1 reveal a non-swapped architecture, Nat Struct Mol Biol., 27, 84, 10.1038/s41594-019-0359-y Burada, 2020, The architecture of GluD2 ionotropic delta glutamate receptor elucidated by cryo-EM, J Struct Biol., 211, 107546, 10.1016/j.jsb.2020.107546 Chen, 2000, Stargazin regulates synaptic targeting of AMPA receptors by two distinct mechanisms, Nature, 408, 936, 10.1038/35050030 Chen, 2005, The 4.1 protein coracle mediates subunit-selective anchoring of Drosophila glutamate receptors to the postsynaptic actin cytoskeleton, J. Neurosci., 25, 6667, 10.1523/JNEUROSCI.1527-05.2005 Cho, 2013, SPIN90 dephosphorylation is required for cofilin-mediated actin depolymerization in NMDA-stimulated hippocampal neurons, Cell. Mol. Life Sci., 70, 4369, 10.1007/s00018-013-1391-4 Cingolani, 2008, Actin in action: the interplay between the actin cytoskeleton and synaptic efficacy, Nat. Rev. Neurosci., 9, 344, 10.1038/nrn2373 Colledge, 2000, Targeting of PKA to glutamate receptors through a MAGUK-AKAP complex, Neuron, 27, 107, 10.1016/S0896-6273(00)00013-1 Colón-Ramos, 2009, Synapse formation in developing neural circuits, Curr. Top. Dev. Biol., 87, 53, 10.1016/S0070-2153(09)01202-2 Comery, 1997, Abnormal dendritic spines in fragile X knockout mice: maturation and pruning deficits, Proc. Natl. Acad. Sci. U.S.A., 5401, 10.1073/pnas.94.10.5401 Coombs, 2009, Transmembrane AMPA receptor regulatory proteins and AMPA receptor function in the cerebellum, Neuroscience, 162, 656, 10.1016/j.neuroscience.2009.01.004 Cooper, 2013, Mechanisms of cell migration in the nervous system, J. Cell Biol., 202, 725, 10.1083/jcb.201305021 Copits, 2013, Kainate receptor post-translational modifications differentially regulate association with 4.1N to control activity-dependent receptor endocytosis, J. Biol. Chem., 288, 8952, 10.1074/jbc.M112.440719 Correia, 2008, Motor protein–dependent transport of AMPA receptors into spines during long-term potentiation, Nat. Neurosci., 11, 457, 10.1038/nn2063 Da Silva, 2003, RhoA/ROCK regulation of neuritogenesis via profilin IIa-mediated control of actin stability, JCB (J. Cell Biol.), 162, 1267, 10.1083/jcb.200304021 Dakoji, 2003, Interaction of transmembrane AMPA receptor regulatory proteins with multiple membrane associated guanylate kinases, Neuropharmacology, 45, 849, 10.1016/S0028-3908(03)00267-3 Das, 2018, 270363 De Mendoza, 2010, Evolution of the MAGUK protein gene family in premetazoan lineages, BMC Evol. Biol., 10, 1, 10.1186/1471-2148-10-93 Dingledine, 1999, The glutamate receptor ion channels, Pharmacol. Rev., 51, 7 Dutta, 2017, Non diaphanous formin delphilin acts as a barbed end capping protein, Exp. Cell Res., 357, 10.1016/j.yexcr.2017.05.014 Ehlers, 1996, vol. 84, 745 Elegheert, 2016, Structural basis for integration of GluD receptors within synaptic organizer complexes, Science, 353, 295, 10.1126/science.aae0104 Elias, 2008, Differential trafficking of AMPA and NMDA receptors by SAP102 and PSD-95 underlies synapse development, Proc. Natl. Acad. Sci. U. S. A, 105, 20953, 10.1073/pnas.0811025106 Fiala, 1998, Synaptogenesis via dendritic filopodia in developing hippocampal area CA1, J. Neurosci., 18, 8900, 10.1523/JNEUROSCI.18-21-08900.1998 Firth, 2011, The deciphering developmental disorders (DDD) study, Dev. Med. Child Neurol., 53, 702, 10.1111/j.1469-8749.2011.04032.x Fossati, 2019, Trans-synaptic signaling through the glutamate receptor delta-1 mediates inhibitory Synapse formation in cortical pyramidal neurons, Neuron, 104, 1081, 10.1016/j.neuron.2019.09.027 Gohla, 2005, Chronophin, a novel HAD-type serine protein phosphatase, regulates cofilin-dependent actin dynamics, Nat. Cell Biol., 7, 21, 10.1038/ncb1201 Goo, 2018, Arc/Arg3.1 has an activity-regulated interaction with PICK1 that results in altered spatial dynamics, Sci. Rep., 8, 1, 10.1038/s41598-018-32821-4 Graham, 2009, A nomenclature for ligand-gated ion channels, Pharmacology, 56, 2 Gu, 2010, ADF/Cofilin-Mediated actin dynamics regulate AMPA receptor trafficking during synaptic plasticity, Nat. Neurosci., 13, 1208, 10.1038/nn.2634 Guo, 2017, A NMDA-receptor calcium influx assay sensitive to stimulation by glutamate and glycine/D-serine, Sci. Rep., 7, 1 Gupta, 2015, Essential role of GluD1 in dendritic spine development and GluN2B to GluN2A NMDAR subunit switch in the cortex and hippocampus reveals ability of GluN2B inhibition in correcting hyperconnectivity, Neuropharmacology, 93, 274, 10.1016/j.neuropharm.2015.02.013 Haas, 2018, Pre-post synaptic alignment through neuroligin-1 tunes synaptic transmission efficiency, eLife, 7, 1, 10.7554/eLife.31755 Haglerød, 2017, Presynaptic PICK1 facilitates trafficking of AMPA-receptors between active zone and synaptic vesicle pool, Neuroscience, 344, 104, 10.1016/j.neuroscience.2016.12.042 Hall, 1998, Rho GTPases and the actin cytoskeleton, Science, 279, 509, 10.1126/science.279.5350.509 Halpain, 2000, Actin and the agile spine: how and why do dendritic spines dance?, Trends Neurosci., 23, 141, 10.1016/S0166-2236(00)01576-9 Hanley, 2008, PICK1: a multi-talented modulator of AMPA receptor trafficking, Pharmacol. Therapeut., 118, 152, 10.1016/j.pharmthera.2008.02.002 Hanley, 2014, Actin-dependent mechanisms in AMPA receptor, Front. Cell. Neurosci., 8 Hardingham, 2019, NMDA receptor C-terminal signaling in development, plasticity, and disease [version 1; peer review: 2 approved], F1000Research, 1 Hayashi, 2010, The postsynaptic density proteins Homer and Shank form a polymeric network structure, Cell, 137, 159, 10.1016/j.cell.2009.01.050 Henley JM, 2013, AMPA receptor trafficking and the mechanisms underlying synaptic plasticity and cognitive aging, Dialogues Clin. Neurosci., 15, 11, 10.31887/DCNS.2013.15.1/jhenley Hepp, 2015, Glutamate receptors of the delta family are widely expressed in the adult brain, Brain Struct Funct, 220, 2797, 10.1007/s00429-014-0827-4 Herguedas, 2016, Structure and organization of heteromeric AMPA-type glutamate receptors, Science, 352, 10.1126/science.aad3873 Hirai, 2000, Clustering of δ glutamate receptors is regulated by the actin cytoskeleton in the dendritic spines of cultured rat Purkinje cells, Eur. J. Neurosci., 12, 563, 10.1046/j.1460-9568.2000.00938.x Hirai, 1999, Interaction of the C-terminal domain of δ glutamate receptor with spectrin in the dendritic spines of cultured Purkinje cells, Neurosci. Res., 34, 281, 10.1016/S0168-0102(99)00061-9 Hirano, 2012, Glutamate-receptor-like molecule GluRδ2 involved in synapse formation at parallel fiber-Purkinje neuron synapses, Cerebellum, 11, 71, 10.1007/s12311-010-0170-0 Hodges, 2014, α-Actinin-2 mediates spine morphology and assembly of the post-synaptic density in hippocampal neurons, PLoS One, 9, 10.1371/journal.pone.0101770 Honkura, 2008, The subspine organization of actin fibers regulates the structure and plasticity of dendritic spines, Neuron, 57, 719, 10.1016/j.neuron.2008.01.013 Hotulainen, 2010, Actin in dendritic spines: connecting dynamics to function, JCB (J. Cell Biol.), 189, 619, 10.1083/jcb.201003008 Howard Chia-Hao Chang, 2005, Cytosolic tail sequences and subunit interactions are critical for synaptic localization of glutamate receptors, JCS (J. Chromatogr. Sci.), 118, 1945 Howard, 2010, The role of SAP97 in synaptic glutamate receptor dynamics, Proc. Natl. Acad. Sci. U. S. A, 107, 3805, 10.1073/pnas.0914422107 Huttlin, 2010, A tissue-specific atlas of mouse protein phosphorylation and expression, Cell, 143, 1174, 10.1016/j.cell.2010.12.001 Ichikawa, 2016, GluD2 endows parallel fiber–Purkinje cell synapses with a high regenerative capacity, J. Neurosci., 36, 4846, 10.1523/JNEUROSCI.0161-16.2016 Kakegawa, 2007, The δ2 “ionotropic” glutamate receptor functions as a non-ionotropic receptor to control cerebellar synaptic plasticity, J. Physiol., 584, 89, 10.1113/jphysiol.2007.141291 Kalinowska, 2015, Actinin-4 governs dendritic spine dynamics and promotes their remodeling by metabotropic glutamate receptors, J. Biol. Chem., 290, 15909, 10.1074/jbc.M115.640136 Kanellos, 2016, Cellular functions of the ADF/cofilin family at a glance, J. Cell Sci., 129, 3211 Kanjhan, 2016, Emerging roles of filopodia and dendritic spines in motoneuron pllasticity during development and disease, Neural Plast., 2016, 10.1155/2016/3423267 Keith, 2008, Excitation control: balancing PSD-95 function at the synapse, Front. Mol. Neurosci., 1 Kempermann, 2010, Neurogenesis in the intact adult brain, Neuroscience Kennedy, 2016, Synaptic signaling in learning and memory, Cold Spring Harb Perspect Biol, 8, 10.1101/cshperspect.a016824 Khan, 2017, Ionotropic glutamate receptors (iGluRs) of the delta family (GluD1 and GluD2) and synaptogenesis, Alex. J. Med., 53, 201 Kim, 2003, The role of synaptic GTPase-activating protein in neuronal development and synaptic plasticity, J. Neurosci., 23, 1119, 10.1523/JNEUROSCI.23-04-01119.2003 Kim, 2016, PLPP/CIN regulates bidirectional synaptic plasticity via GluN2A interaction with postsynaptic proteins, Sci. Rep., 6, 1 Kimberley, 2007, Dynamic aspects of Synapse formation, Annu. Rev. Neurosci., 30, 425, 10.1146/annurev.neuro.29.051605.112830 Klaavuniemi, 2004, The ZASP-like motif in actinin-associated LIM protein is required for interaction with the α-actinin rod and for targeting to the muscle Z-line, J. Biol. Chem., 279, 26402, 10.1074/jbc.M401871200 Kneussel, 2013, Myosin motors at neuronal synapses: drivers of membrane transport and actin dynamics, Nat. Rev. Neurosci., 14, 233, 10.1038/nrn3445 Kohda, 2016, Delta glutamate receptor (GluD1, GluD2), 978 Konno, 2014, Enriched expression of GluD1 in higher brain regions and its involvement in parallel fiber-interneuron synapse formation in the cerebellum, J. Neurosci., 34, 7412, 10.1523/JNEUROSCI.0628-14.2014 Korkotian, 1999, Bidirectional regulation of dendritic spine dimensions by glutamate receptors, Neuroreport, 10, 2875, 10.1097/00001756-199909090-00032 Korn, 1987, Actin polymerization and ATP hydrolysis, Science, 238, 638, 10.1126/science.3672117 Kornau, 1995, Domain interaction between NMDA receptor subunits and the postsynaptic density protein PSD-95, Science, 269, 1737, 10.1126/science.7569905 Korobova, 2010, Molecular architecture of synaptic actin cytoskeleton in hippocampal neurons reveals a mechanism of dendritic spine morphogenesis, Mol Biol Cell, 21, 165, 10.1091/mbc.e09-07-0596 Lamprecht, 2016, The role of actin cytoskeleton in memory formation in amygdala, Front. Mol. Neurosci., 9 Lee, 2010, Regulation of actin cytoskeleton dynamics in cells, Mol. Cell., 4, 311, 10.1007/s10059-010-0053-8 Lei 1, 2001, Regulation of NMDA receptor activity by F-actin and myosin light chain kinase, J. Neurosci., 21, 8464, 10.1523/JNEUROSCI.21-21-08464.2001 Letellier, 2018, A unique intracellular tyrosine in neuroligin-1 regulates AMPA receptor recruitment during synapse differentiation and potentiation, Nat Commun, 9, 3979, 10.1038/s41467-018-06220-2 Letort, 2015, Dynamic reorganization of the actin cytoskeleton, F1000Research, 4 Lin, 2009, Regulation of AMPA receptor extrasynaptic insertion by 4.1N, phosphorylation and palmitoylation, Nat. Neurosci., 12, 879, 10.1038/nn.2351 Lodish, 2000, Chapter 18: secction 18.2: the dynamics of actin assembly Lu, 2014, Trafficking of α-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor (AMPA) receptor subunit GluA2 from the endoplasmic reticulum is stimulated by a complex containing Ca2+/calmodulin-activated kinase II (CaMKII) and PICK1 protein and by release o, J. Biol. Chem., 289, 19218, 10.1074/jbc.M113.511246 Lue, 1994, Cloning and characterization of hdlg: the human homologue of the Drosophila discs large tumor suppressor binds to protein 4.1, Proc. Natl. Acad. Sci. U. S. A, 91, 9818, 10.1073/pnas.91.21.9818 Lundby, 2012, Quantitative maps of protein phosphorylation sites across 14 different rat organs and tissues, Nat. Commun., 3, 10.1038/ncomms1871 Lynch, 2004, Long-term potentiation and memory, Physiol. Rev., 84, 87, 10.1152/physrev.00014.2003 MacGillavry, 2013, Nanoscale scaffolding domains within the postsynaptic density concentrate synaptic ampa receptors, Neuron, 78, 615, 10.1016/j.neuron.2013.03.009 Macgillavry, 2016, Shank–cortactin interactions control actin dynamics to maintain flexibility of neuronal spines and synapses, Eur. J. Neurosci., 43, 179, 10.1111/ejn.13129 Maiti, 2013, Actin-capping and -severing proteins, 18 Mandolesi, 2009, An orphan ionotropic glutamate receptor: the δ2 subunit, Neuroscience, 158, 67, 10.1016/j.neuroscience.2008.02.050 Matt, 2018, α-Actinin anchors PSD-95 at postsynaptic sites, Neuron, 97, 1094, 10.1016/j.neuron.2018.01.036 Meng, 2002, Abnormal spine morphology and enhanced LTP in LIMK-1 knockout mice, Neuron, 35, 121, 10.1016/S0896-6273(02)00758-4 Meng, 2004, Regulation of ADF/cofilin phosphorylation and synaptic function by LIM-kinase, Neuropharmacology, 47, 746, 10.1016/j.neuropharm.2004.06.030 Menna, 2013, Eps8 controls dendritic spine density and synaptic plasticity through its actin-capping activity, EMBO J., 32, 1730, 10.1038/emboj.2013.107 Michaelsen, 2010, Fine-tuning of neuronal architecture requires two profilin isoforms, Proc. Natl. Acad. Sci. U. S. A, 107, 15780, 10.1073/pnas.1004406107 Miermans, 2017, Biophysical model of the role of actin remodelling on dendritic spine morphology, PLoS One, 12, 10.1371/journal.pone.0170113 Miyagi, 2002, Delphilin: a novel PDZ and formin homology domain-containing protein that synaptically colocalizes and interacts with glutamate receptor δ2 subunit, J. Neurosci., 22, 803, 10.1523/JNEUROSCI.22-03-00803.2002 Mizui, 2009, Drebrin E is involved in the regulation of axonal growth through actin-myosin interactions, J. Neurochem., 109, 611, 10.1111/j.1471-4159.2009.05993.x Mondin, 2011, Neurexin-neuroligin adhesions capture surface-diffusing AMPA receptors through PSD-95 scaffolds, J. Neurosci., 31, 13500, 10.1523/JNEUROSCI.6439-10.2011 Morini, 2018, Lack of the actin capping protein, Eps8, affects NMDA-type glutamate receptor function and composition, Front. Mol. Neurosci., 11, 1 Murata, 2013, Postsynaptic density scaffold SAP102 regulates cortical synapse development through EphB and PAK signaling pathway, J. Neurosci., 33, 5040, 10.1523/JNEUROSCI.2896-12.2013 Nair, 2013, Super-resolution imaging reveals that AMPA receptors inside synapses are dynamically organized in nanodomains regulated by PSD95, J. Neurosci., 33, 13204, 10.1523/JNEUROSCI.2381-12.2013 Nakamoto, 2020, Expression mapping, quantification, and complex formation of GluD1 and GluD2 glutamate receptors in adult mouse brain, J. Comp. Neurol., 528, 1003, 10.1002/cne.24792 Nash, 2010, Disruption of the interaction between myosin VI and SAP97 is associated with a reduction in the number of AMPARs at hippocampal synapses, J. Neurochem., 112, 677, 10.1111/j.1471-4159.2009.06480.x Naur, 2007, Ionotropic glutamate-like receptor δ2 binds D-serine and glycine, Proc. Natl. Acad. Sci. U. S. A, 104, 14116, 10.1073/pnas.0703718104 Nicoll, 2006, Auxiliary subunits assist AMPA-type glutamate receptors, Science, 311, 1253, 10.1126/science.1123339 Niethammer, 1996, Interaction between the C terminus of NMDA receptor subunits and multiple members of the PSD-95 family of membrane-associated guanylate kinases, J. Neurosci., 16, 2157, 10.1523/JNEUROSCI.16-07-02157.1996 Niwa, 2002, Control of actin reorganization by slingshot, a family of phosphatases that dephosphorylate ADF/cofilin, Cell, 108, 233, 10.1016/S0092-8674(01)00638-9 Osterweil, 2005, A role for myosin VI in postsynaptic structure and glutamate receptor endocytosis, JCB (J. Cell Biol.), 168, 329, 10.1083/jcb.200410091 Oswald, 2017, New insights into the mechanism of Ca2+-dependent inactivation of NMDA receptors, Biophys. J., 113, 2131, 10.1016/j.bpj.2017.10.008 O'Kane, 2003, Activation of Rho GTPases by synaptic transmission in the hippocampus, J. Neurochem., 87, 1309, 10.1046/j.1471-4159.2003.02102.x Pandey, 2015, Development- and age-related alterations in the expression of AMPA receptor subunit GluR2 and its trafficking proteins in the hippocampus of male mouse brain, Biogerontology, 16, 317, 10.1007/s10522-014-9548-6 Park, 2016, CaMKII phosphorylation of TARPγ-8 is a mediator of LTP and learning and memory, Neuron, 92, 75, 10.1016/j.neuron.2016.09.002 Payne, 2008, The role of transmembrane AMPA receptor regulatory proteins (TARPs) in neurotransmission and receptor trafficking, J. Mol. Membr. Biol., 25 Pelucchi, 2020, Dendritic spines in alzheimer's disease: how the actin cytoskeleton contributes to synaptic failure, Int. J. Mol. Sci., 908, 10.3390/ijms21030908 Platt, 2007, The role of glutamate in central nervous system health and disease--a review, Vet. J., 173, 278, 10.1016/j.tvjl.2005.11.007 Pollard, 2016, 1 Pollard, 2003, Cellular motility driven by assembly and disassembly of actin filaments, Cell, 112, 453, 10.1016/S0092-8674(03)00120-X Qualmann, 2004, Linkage of the actin cytoskeleton to the postsynaptic density via direct interactions of Abp1 with the ProSAP/shank family, J. Neurosci., 24, 2481, 10.1523/JNEUROSCI.5479-03.2004 Racz, 2006, Spatial organization of cofilin in dendritic spines, Neuroscience, 138, 447, 10.1016/j.neuroscience.2005.11.025 Reiner, 2018, Glutamatergic signaling in the central nervous system: ionotropic and metabotropic receptors in concert, Neuron, 98, 1080, 10.1016/j.neuron.2018.05.018 Reshetniak, 2019, Interrogating synaptic architecture: approaches for labeling organelles and cytoskeleton components, Front. Synaptic Neurosci., 11 Rocca, 2015, PICK1 links AMPA receptor stimulation to Cdc42, Neurosci. Lett., 585, 155, 10.1016/j.neulet.2014.11.046 Rocca, 2008, Inhibition of Arp2/3-mediated actin polymerization by PICK1 regulates neuronal morphology and AMPA receptor endocytosis, Nat. Cell Biol., 10, 259, 10.1038/ncb1688 Rocca, 2013, The small GTPase Arf1 modulates arp2/3-mediated actin polymerization via PICK1 to regulate synaptic plasticity, Neuron, 79, 293, 10.1016/j.neuron.2013.05.003 Romorini, 2004, A functional role of postsynaptic density-95-guanylate kinase-associated protein complex in regulating shank assembly and stability to synapses, J. Neurosci., 24, 9391, 10.1523/JNEUROSCI.3314-04.2004 Rossmann, 2011, Subunit-selective N-terminal domain associations organize the formation of AMPA receptor heteromers, EMBO J., 30, 959, 10.1038/emboj.2011.16 Rudolf, 2011, Gerdes, Hans-Hermann. The role of myosin V in exocytosis and synaptic plasticity, J. Neurochem, 116, 177, 10.1111/j.1471-4159.2010.07110.x Rumbaugh, 2003, Synapse-associated protein-97 isoform-specific regulation of surface AMPA receptors and synaptic function in cultured neurons, J. Neurosci., 23, 4567, 10.1523/JNEUROSCI.23-11-04567.2003 Rycroft, 2004, Regulation of single NMDA receptor channel activity by alpha-actinin and calmodulin in rat hippocampal granule cells, J. Physiol., 557, 795, 10.1113/jphysiol.2003.059212 Saglietti, 2007, Extracellular interactions between GluR2 and N-cadherin in spine regulation, Neuron, 54, 461, 10.1016/j.neuron.2007.04.012 Salinas, 2005, 73 Salinas, 2006, Actinfilin is a Cul3 substrate adaptor, linking GluR6 kainate receptor subunits to the ubiquitin-proteasome pathway, J. Biol. Chem., 281, 40164, 10.1074/jbc.M608194200 Sans, 2000, A developmental change in NMDA receptor-associated proteins at hippocampal synapses, J. Neurosci., 20, 1260, 10.1523/JNEUROSCI.20-03-01260.2000 Sans, 2003, NMDA receptor trafficking through an interaction between PDZ proteins and the exocyst complex, Nat. Cell Biol., 5, 520, 10.1038/ncb990 Schapitz, 2010, Neuroligin 1 is dynamically exchanged at postsynaptic sites, J. Neurosci., 30, 12733, 10.1523/JNEUROSCI.0896-10.2010 Schnell, 2002, Direct interactions between PSD-95 and stargazin control synaptic AMPA receptor number, Proc. Natl. Acad. Sci. U. S. A, 99, 13902, 10.1073/pnas.172511199 Schubert, 2006, Localized recruitment and activation of RhoA underlies dendritic spine morphology in a glutamate receptor-dependent manner, JCB (J. Cell Biol.), 172, 453, 10.1083/jcb.200506136 Schulz, 2004, Actin/alpha-actinin-dependent transport of AMPA receptors in dendritic spines: role of the PDZ-LIM protein RIL, J. Neurosci., 24, 8584, 10.1523/JNEUROSCI.2100-04.2004 Sekino, 2017, Role of drebrin in synaptic plasticity, Adv. Exp. Med. Biol., 1006, 183, 10.1007/978-4-431-56550-5_11 Shen, 2000, Regulation of AMPA receptor GluR1 subunit surface expression by a 4. 1N-linked actin cytoskeletal association, J. Neurosci., 20, 7932, 10.1523/JNEUROSCI.20-21-07932.2000 Sheng, 2011, The postsynaptic organization of synapses, Cold Spring Harb. Perspect. Biol., 3, 10.1101/cshperspect.a005678 Shi, 2009, Focal adhesion kinase acts downstream of EphB receptors to maintain mature dendritic spines by regulating cofilin activity, J. Neurosci., 29, 8129, 10.1523/JNEUROSCI.4681-08.2009 Shirao, 2017, The role of drebrin in neurons, J. Neurochem., 141, 819, 10.1111/jnc.13988 Silkworth, 2018, The neuron-specific formin Delphilin nucleates nonmuscle actin but does not enhance elongation, Mol. Biol. Cell, 29, 610, 10.1091/mbc.E17-06-0363 Sit, 2011, Rho GTPases and their role in organizing the actin cytoskeleton, J. Cell Sci., 124, 679, 10.1242/jcs.064964 Sonoda, 2006, Binding of glutamate receptor δ2 to its scaffold protein, Delphilin, is regulated by PKA, Biochem. Biophys. Res. Commun., 350, 748, 10.1016/j.bbrc.2006.09.109 Stamatakou, 2013, Activity-dependent spine morphogenesis: a role for the actin-capping protein Eps8, J. Neurosci., 33, 2661, 10.1523/JNEUROSCI.0998-12.2013 Stan, 2010, Essential cooperation of N-cadherin and neuroligin-1 in the transsynaptic control of vesicle accumulation, Proc. Natl. Acad. Sci. U. S. A, 107, 11116, 10.1073/pnas.0914233107 Staudinger, 1995, PICK1: a perinuclear binding protein and substrate for protein kinase C isolated by the yeast two-hybrid system, JCB (J. Cell Biol.), 128, 263, 10.1083/jcb.128.3.263 Stefen, 2016, Regulation of the postsynaptic compartment of excitatory synapses by the actin cytoskeleton in health and its disruption in disease, Neural Plast., 2371970 Stefen, 2016 Svitkina, 2018, The actin cytoskeleton and actin-based motility, Cold Spring Harb Perspect Biol, 10, 10.1101/cshperspect.a018267 Tada, 2006, Molecular mechanisms of dendritic spine morphogenesis, Curr. Opin. Neurobiol., 16, 95, 10.1016/j.conb.2005.12.001 Tadros, 2007, Glutamate-related gene expression changes with age in the mouse auditory midbrain, Brain Res., 1127, 1, 10.1016/j.brainres.2006.09.081 Takahashi, 2009, Activity of the AMPA receptor regulates drebrin stabilization in dendritic spine morphogenesis, J. Cell Sci., 122, 1211, 10.1242/jcs.043729 Takeichi, 2005, Synaptic contact dynamics controlled by cadherin and catenins, Trends Cell Biol., 15, 216, 10.1016/j.tcb.2005.02.002 Tao, 2019, Mechanisms underlying the synaptic trafficking of the glutamate delta receptor GluD1, Mol. Psychiatr., 24, 1451, 10.1038/s41380-019-0378-4 Tejada-Simon, 2006, NMDA receptor activation induces translocation and activation of Rac in mouse hippocampal area CA1, Biochem. Biophys. Res. Commun., 343, 504, 10.1016/j.bbrc.2006.02.183 Tomita, 2005, Bidirectional synaptic plasticity regulated by phosphorylation of stargazin-like TARPs, Neuron, 45, 269, 10.1016/j.neuron.2005.01.009 Toshima, 2001, Cofilin phosphorylation and actin reorganization activities of testicular protein kinase 2 and its predominant expression in testicular sertoli cells, J. Biol. Chem., 276, 31449, 10.1074/jbc.M102988200 Traynelis, 2010, Glutamate receptor ion channels: structure, regulation, and function, Pharmacol. Rev., 62, 405, 10.1124/pr.109.002451 Vallenius, 2004, The PDZ-LIM protein RIL modulates actin stress fiber turnover and enhances the association of α-actinin with F-actin, Exp. Cell Res., 293, 117, 10.1016/j.yexcr.2003.09.004 Van Spronsen, 2010, Synapse pathology in psychiatric and neurologic disease, Curr. Neurol. Neurosci. Rep., 10, 207, 10.1007/s11910-010-0104-8 Vieira, 2020, Regulation of NMDA glutamate receptor functions by the GluN2 subunits, J. Neurochem., 154, 121, 10.1111/jnc.14970 Vyas, 2016, The role of postsynaptic density proteins in neural degeneration and regeneration, Neural Regener. Res., 11, 906, 10.4103/1673-5374.184481 Wang, 2014, Active calcium/calmodulin-dependent protein kinase II (CaMKII) regulates nmda receptor mediated postischemic long-term potentiation (i-LTP) by promoting the interaction between CaMKII and NMDA receptors in ischemia, Neural Plast., 10.1155/2014/827161 Warnet, 2020, The C-terminal domains of the NMDA receptor: how intrinsically disordered tails affect signalling, plasticity and disease, Eur. J. Neurosci. Wechsler, 1998, Brain spectrin binding to the NMDA receptor is regulated by phosphorylation, calcium and calmodulin, EMBO J., 17, 3931, 10.1093/emboj/17.14.3931 Winder, 2005, Actin-binding proteins, J. Cell Sci., 118, 651, 10.1242/jcs.01670 Wozny, 2009, The function of glutamatergic synapses is not perturbed by severe knockdown of 4.1N and 4.1G expression, J. Cell Sci., 122, 735, 10.1242/jcs.037382 Wyszynski, 1997, Competitive binding of α-actinin and calmodulin to the NMDA receptor, Nature, 385, 439, 10.1038/385439a0 Xu, 2004, Crystal structures of a Formin Homology-2 domain reveal a tethered dimer architecture, Cell, 116, 711, 10.1016/S0092-8674(04)00210-7 Yamazaki, 1992, Molecular cloning of a cDNA encoding a novel member of the mouse glutamate receptor channel family, Biochem. Biophys. Res. Commun., 183, 886, 10.1016/0006-291X(92)90566-4 Yan, 2016, Synaptic actin dysregulation, a convergent mechanism of mental disorders?, J. Neurosci., 36, 11411, 10.1523/JNEUROSCI.2360-16.2016 Yan, 2016, Synaptic actin dysregulation, a convergent mechanism of mental disorders?, J. Neurosci., 36, 11411, 10.1523/JNEUROSCI.2360-16.2016 Yang, 1998, Cofflin phosphorylation by LIM-kinase 1 and its role in Rac-mediated actin reorganization, Nature, 393, 809, 10.1038/31735 Yasuda, 2018, Drebrin isoforms critically regulate nmdar- and mglur-dependent LTD induction, Front. Cell. Neurosci., 12, 1 Yuzaki, 2003, 10 years later glutamate receptor.pdf, Neurosci. Res., 46, 11, 10.1016/S0168-0102(03)00036-1 Yuzaki, 2017, A GluD coming-of-age story, Trends Neurosci., 40, 138, 10.1016/j.tins.2016.12.004 Zhou, 2001, Contribution of cytoskeleton to the internalization of AMPA receptors, Proc. Natl. Acad. Sci. U. S. A., 98, 1261, 10.1073/pnas.98.3.1261 Zhu, 2016, Mechanistic basis of MAGUK-organized complexes in synaptic development and signalling, Nat. Rev. Neurosci., 17, 209, 10.1038/nrn.2016.18