{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,7]],"date-time":"2026-09-07T13:27:24Z","timestamp":1788787644649,"version":"build-2803163510"},"reference-count":72,"publisher":"MDPI AG","issue":"10","license":[{"start":{"date-parts":[[2021,10,3]],"date-time":"2021-10-03T00:00:00Z","timestamp":1633219200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["NIH NS104705 (SMD), NIH MH116003 (SMD), NIH NS118731(SMD, VN), NIH NS038261 (VN), NIH NS106902 (VN), NIH NS109255 (VN)"],"award-info":[{"award-number":["NIH NS104705 (SMD), NIH MH116003 (SMD), NIH NS118731(SMD, VN), NIH NS038261 (VN), NIH NS106902 (VN), NIH NS109255 (VN)"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["NSF1456818 (SMD)"],"award-info":[{"award-number":["NSF1456818 (SMD)"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>Chronic pain is a debilitating condition involving neuronal dysfunction, but the synaptic mechanisms underlying the persistence of pain are still poorly understood. We found that the synaptic organizer glutamate delta 1 receptor (GluD1) is expressed postsynaptically at parabrachio-central laterocapsular amygdala (PB-CeLC) glutamatergic synapses at axo-somatic and punctate locations on protein kinase C \u03b4 -positive (PKC\u03b4+) neurons. Deletion of GluD1 impairs excitatory neurotransmission at the PB-CeLC synapses. In inflammatory and neuropathic pain models, GluD1 and its partner cerebellin 1 (Cbln1) are downregulated while AMPA receptor is upregulated. A single infusion of recombinant Cbln1 into the central amygdala led to sustained mitigation of behavioral pain parameters and normalized hyperexcitability of central amygdala neurons. Cbln2 was ineffective under these conditions and the effect of Cbln1 was antagonized by GluD1 ligand D-serine. The behavioral effect of Cbln1 was GluD1-dependent and showed lateralization to the right central amygdala. Selective ablation of GluD1 from the central amygdala or injection of Cbln1 into the central amygdala in normal animals led to changes in averse and fear-learning behaviors. Thus, GluD1-Cbln1 signaling in the central amygdala is a teaching signal for aversive behavior but its sustained dysregulation underlies persistence of pain. Significance statement: Chronic pain is a debilitating condition which involves synaptic dysfunction, but the underlying mechanisms are not fully understood. Our studies identify a novel mechanism involving structural synaptic changes in the amygdala caused by impaired GluD1-Cbln1 signaling in inflammatory and neuropathic pain behaviors. We also identify a novel means to mitigate pain in these conditions using protein therapeutics.<\/jats:p>","DOI":"10.3390\/cells10102644","type":"journal-article","created":{"date-parts":[[2021,10,4]],"date-time":"2021-10-04T09:24:15Z","timestamp":1633339455000},"page":"2644","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":35,"title":["Dysfunction of Glutamate Delta-1 Receptor-Cerebellin 1 Trans-Synaptic Signaling in the Central Amygdala in Chronic Pain"],"prefix":"10.3390","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9793-9107","authenticated-orcid":false,"given":"Pauravi J.","family":"Gandhi","sequence":"first","affiliation":[{"name":"Department of Pharmacology and Neuroscience, Creighton University School of Medicine, Omaha, NE 68178, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dinesh Y.","family":"Gawande","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, Creighton University School of Medicine, Omaha, NE 68178, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9351-8375","authenticated-orcid":false,"given":"Gajanan P.","family":"Shelkar","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, Creighton University School of Medicine, Omaha, NE 68178, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sukanya G.","family":"Gakare","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, Creighton University School of Medicine, Omaha, NE 68178, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5297-3379","authenticated-orcid":false,"given":"Takaki","family":"Kiritoshi","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, School of Medicine, Texas Tech University Health Sciences Center, Lubbock, TX 79430, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0606-0181","authenticated-orcid":false,"given":"Guangchen","family":"Ji","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, School of Medicine, Texas Tech University Health Sciences Center, Lubbock, TX 79430, USA"},{"name":"Center of Excellence for Translational Neuroscience and Therapeutics, Texas Tech University Health Sciences Center, Lubbock, TX 79430, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bishal","family":"Misra","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, Creighton University School of Medicine, Omaha, NE 68178, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ratnamala","family":"Pavuluri","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, Creighton University School of Medicine, Omaha, NE 68178, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinxu","family":"Liu","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, Creighton University School of Medicine, Omaha, NE 68178, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Volker","family":"Neugebauer","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, School of Medicine, Texas Tech University Health Sciences Center, Lubbock, TX 79430, USA"},{"name":"Center of Excellence for Translational Neuroscience and Therapeutics, Texas Tech University Health Sciences Center, Lubbock, TX 79430, USA"},{"name":"Garrison Institute on Aging, Texas Tech University Health Sciences Center, Lubbock, TX 79430, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shashank M.","family":"Dravid","sequence":"additional","affiliation":[{"name":"Department of Pharmacology and Neuroscience, Creighton University School of Medicine, Omaha, NE 68178, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2021,10,3]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1113\/eph8702357","article-title":"Pain pathways and parabrachial circuits in the rat","volume":"87","author":"Gauriau","year":"2002","journal-title":"Exp. Physiol."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1016\/j.neuron.2005.08.015","article-title":"Parallel \u201cpain\u201d pathways arise from subpopulations of primary afferent noci-ceptor","volume":"47","author":"Braz","year":"2005","journal-title":"Neuron"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"261","DOI":"10.1007\/978-3-662-46450-2_13","article-title":"Amygdala pain mechanisms","volume":"227","author":"Neugebauer","year":"2015","journal-title":"Handb. Exp. Pharmacol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1177\/1073858403261077","article-title":"The Amygdala and Persistent Pain","volume":"10","author":"Neugebauer","year":"2004","journal-title":"Neuroscientist"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"2433","DOI":"10.1046\/j.1460-9568.2003.02981.x","article-title":"A quantitative and morphological study of projection neurons in lamina I of the rat lumbar spinal cord","volume":"18","author":"Spike","year":"2003","journal-title":"Eur. J. Neurosci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"8296501","DOI":"10.1155\/2017\/8296501","article-title":"Amygdala Plasticity and Pain","volume":"2017","author":"Thompson","year":"2017","journal-title":"Pain Res. Manag."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"1543","DOI":"10.1523\/JNEUROSCI.3536-06.2007","article-title":"Activation of the extracellular signal-regulated kinase in the amygdala modulates pain perception","volume":"27","author":"Carrasquillo","year":"2007","journal-title":"J. Neurosci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"14217","DOI":"10.1523\/JNEUROSCI.1473-12.2012","article-title":"Central amygdala metabotropic gluta-mate receptor 5 in the modulation of visceral pain","volume":"32","author":"Crock","year":"2012","journal-title":"J. Neurosci."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"10717","DOI":"10.1523\/JNEUROSCI.4112-05.2005","article-title":"Critical Role of Calcitonin Gene-Related Peptide 1 Receptors in the Amygdala in Synaptic Plasticity and Pain Behavior","volume":"25","author":"Han","year":"2005","journal-title":"J. Neurosci."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1016\/j.neulet.2003.12.027","article-title":"Synaptic plasticity in the amygdala in a visceral pain model in rats","volume":"361","author":"Han","year":"2004","journal-title":"Neurosci. Lett."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/j.pain.2006.09.003","article-title":"NMDA receptor-independent synaptic plasticity in the central amygdala in the rat model of neuropathic pain","volume":"127","author":"Ikeda","year":"2007","journal-title":"Pain"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1007\/978-981-13-1756-9_14","article-title":"Pain-Associated Neural Plasticity in the Parabrachial to Central Amygdala Circuit: Pain Changes the Brain, and the Brain Changes the Pain","volume":"1099","author":"Kato","year":"2018","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"8203","DOI":"10.1523\/JNEUROSCI.1216-10.2010","article-title":"Activation of metabotropic glutamate receptor 5 in the amygdala modulates pain-like behavior","volume":"30","author":"Kolber","year":"2010","journal-title":"J. Neurosci."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1523\/JNEUROSCI.23-01-00052.2003","article-title":"Synaptic plasticity in the amygdala in a model of arthritic pain: Differential roles of metabotropic glutamate receptors 1 and 5","volume":"23","author":"Neugebauer","year":"2003","journal-title":"J. Neurosci."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1186\/2049-9256-1-9","article-title":"The amygdala between sensation and affect: A role in pain","volume":"1","author":"Veinante","year":"2013","journal-title":"J. Mol. Psychiatry"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1113\/jphysiol.2005.084780","article-title":"Protein kinase A-dependent enhanced NMDA receptor function in pain-related synaptic plasticity in rat amygdala neurones","volume":"564","author":"Bird","year":"2005","journal-title":"J. Physiol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"2258","DOI":"10.1523\/JNEUROSCI.5564-10.2011","article-title":"Role of Extracellular Signal-Regulated Kinase in Synaptic Transmission and Plasticity of a Nociceptive Input on Capsular Central Amygdaloid Neurons in Normal and Acid-Induced Muscle Pain Mice","volume":"31","author":"Cheng","year":"2011","journal-title":"J. Neurosci."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s13041-015-0108-z","article-title":"The lateral parabrachial nucleus is actively in-volved in the acquisition of fear memory in mice","volume":"8","author":"Sato","year":"2015","journal-title":"Mol. Brain"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2721","DOI":"10.1152\/jn.00946.2015","article-title":"Synaptic and network consequences of monosynaptic nociceptive inputs of parabrachial nucleus origin in the central amygdala","volume":"115","author":"Sugimura","year":"2016","journal-title":"J. Neurophysiol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1186\/1756-6606-6-11","article-title":"Synaptic potentiation in the nociceptive amygdala following fear learning in mice","volume":"6","author":"Watabe","year":"2013","journal-title":"Mol. Brain"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1378","DOI":"10.1523\/JNEUROSCI.2468-16.2016","article-title":"5-HT2C Receptor Knockdown in the Amygdala Inhibits Neuropathic-Pain-Related Plasticity and Behaviors","volume":"37","author":"Ji","year":"2017","journal-title":"J. Neurosci."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1186\/1744-8069-8-51","article-title":"Role of Capsaicin-Sensitive C-Fiber Afferents in Neuropathic Pain-Induced Synaptic Potentiation in the Nociceptive Amygdala","volume":"8","author":"Nakao","year":"2012","journal-title":"Mol. Pain"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1016\/S0306-4522(00)00245-1","article-title":"Calcitonin gene-related peptide-like immunoreactivity marks putative visceral sensory pathways in human brain","volume":"100","author":"Saper","year":"2000","journal-title":"Neuroscience"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1002\/cne.20618","article-title":"Calcitonin gene-related peptide-containing pathways in the rat forebrain","volume":"489","author":"Dobolyi","year":"2005","journal-title":"J. Comp. Neurol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1016\/j.cell.2015.05.057","article-title":"Elucidating an Affective Pain Circuit that Creates a Threat Memory","volume":"162","author":"Han","year":"2015","journal-title":"Cell"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1016\/0006-8993(88)90395-2","article-title":"Calcitonin gene-related peptide (CGRP) in the rat central nervous system: Patterns of immunoreactivity and receptor binding sites","volume":"463","author":"Kruger","year":"1988","journal-title":"Brain Res."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"416","DOI":"10.1002\/cne.902700310","article-title":"Neurons containing calcitonin gene-related peptide in the para-brachial nucleus project to the central nucleus of the amygdala","volume":"270","author":"Schwaber","year":"1988","journal-title":"J. Comp. Neurol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1097\/j.pain.0000000000001691","article-title":"Spared nerve injury differentially alters parabrachial monosynaptic excitatory inputs to molecularly specific neurons in distinct subregions of the central amygdala","volume":"161","author":"Li","year":"2019","journal-title":"Pain"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"332","DOI":"10.1016\/j.celrep.2019.09.011","article-title":"Dual and Opposing Functions of the Central Amygdala in the Modulation of Pain","volume":"29","author":"Wilson","year":"2019","journal-title":"Cell Rep."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1007\/s00429-018-01825-1","article-title":"Cell-type specific parallel circuits in the bed nucleus of the stria terminalis and the central nucleus of the amygdala of the mouse","volume":"224","author":"Ye","year":"2019","journal-title":"Brain Struct. Funct."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Adke, A.P., Khan, A., Ahn, H.-S., Becker, J.J., Wilson, T.D., Valdivia, S., Sugimura, Y.K., Gonzalez, S.M., and Carrasquillo, Y. (2021). Cell-Type Specificity of Neuronal Excitability and Morphology in the Central Amygdala. Eneuro, 8.","DOI":"10.1523\/ENEURO.0402-20.2020"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"14116","DOI":"10.1073\/pnas.0703718104","article-title":"Ionotropic glutamate-like receptor delta2 binds D-serine and glycine","volume":"104","author":"Naur","year":"2007","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.brainres.2010.12.086","article-title":"Mutations in the transmembrane domain M3 generate spontaneously open orphan glutamate delta 1 receptor","volume":"1382","author":"Yadav","year":"2011","journal-title":"Brain Res."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Clarke, R., and Eapen, V. (2014). Balance within the Neurexin Trans-Synaptic Connexus Stabilizes Behavioral Control. Front. Hum. Neurosci., 8.","DOI":"10.3389\/fnhum.2014.00052"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"4912","DOI":"10.1073\/pnas.0900892106","article-title":"Postsynaptic glutamate receptor delta family contributes to presynaptic terminal differentiation and establishment of synaptic transmission","volume":"106","author":"Kuroyanagi","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1126\/science.1185152","article-title":"Cbln1 is a ligand for an orphan glutamate receptor delta2, a bidirectional synapse organizer","volume":"328","author":"Matsuda","year":"2010","journal-title":"Science"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1016\/j.bbrc.2011.11.075","article-title":"Induction of excitatory and inhibitory presynaptic differentiation by GluD1","volume":"417","author":"Ryu","year":"2012","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1068","DOI":"10.1016\/j.cell.2010.04.035","article-title":"Trans-synaptic interaction of GluRdelta2 and Neurexin through Cbln1 mediates synapse formation in the cerebellum","volume":"141","author":"Uemura","year":"2010","journal-title":"Cell"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1111\/j.1471-4159.2011.07631.x","article-title":"Glutamate receptor delta1 induces preferentially inhibitory presynaptic differentiation of cortical neurons by interacting with neurexins through cerebellin precursor protein subtypes","volume":"121","author":"Yasumura","year":"2012","journal-title":"J. Neurochem."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"104746","DOI":"10.1016\/j.nbd.2020.104746","article-title":"Striatal glutamate delta-1 receptor regulates behavioral flexibility and thalamostriatal connectivity","volume":"137","author":"Liu","year":"2020","journal-title":"Neurobiol. Dis."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"E5373","DOI":"10.1073\/pnas.1802737115","article-title":"Postsynaptic delta1 glutamate receptor assembles and maintains hippocampal synapses via Cbln2 and neurexin","volume":"115","author":"Tao","year":"2018","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"2797","DOI":"10.1007\/s00429-014-0827-4","article-title":"Glutamate receptors of the delta family are widely expressed in the adult brain","volume":"220","author":"Hepp","year":"2014","journal-title":"Brain Struct. Funct."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"7412","DOI":"10.1523\/JNEUROSCI.0628-14.2014","article-title":"Enriched expression of GluD1 in higher brain regions and its involvement in parallel fiber-interneuron synapse formation in the cere-bellum","volume":"34","author":"Konno","year":"2014","journal-title":"J. Neurosci."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1111\/j.1460-9568.2006.04950.x","article-title":"Distinct expression of Cbln family mRNAs in developing and adult mouse brains","volume":"24","author":"Miura","year":"2006","journal-title":"Eur. J. Neurosci."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"11801","DOI":"10.1523\/JNEUROSCI.0322-16.2016","article-title":"Roles of Cbln1 in Non-Motor Functions of Mice","volume":"36","author":"Otsuka","year":"2016","journal-title":"J. Neurosci."},{"key":"ref_46","doi-asserted-by":"crossref","unstructured":"Yadav, R., Hillman, B.G., Gupta, S.C., Suryavanshi, P., Bhatt, J.M., Pavuluri, R., Stairs, D.J., and Dravid, S.M. (2013). Deletion of Glutamate Delta-1 Receptor in Mouse Leads to Enhanced Working Memory and Deficit in Fear Conditioning. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0060785"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"4500","DOI":"10.1128\/MCB.02051-06","article-title":"Orphan glutamate receptor delta1 subunit required for high-frequency hearing","volume":"27","author":"Gao","year":"2007","journal-title":"Mol. Cell Biol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1124\/mol.119.116780","article-title":"Modulation of Burst Firing of Neurons in Nucleus Reticularis of the Thalamus by GluN2C-Containing NMDA Receptors","volume":"96","author":"Liu","year":"2019","journal-title":"Mol. Pharmacol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"824","DOI":"10.1097\/j.pain.0000000000001458","article-title":"Kappa opioid signaling in the central nucleus of the amygdala promotes disinhibition and aversiveness of chronic neuropathic pain","volume":"160","author":"Navratilova","year":"2019","journal-title":"Pain"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"219","DOI":"10.1016\/j.neuropharm.2018.06.015","article-title":"Small conductance calcium activated potassium (SK) channel dependent and independent effects of riluzole on neuropathic pain-related amygdala activity and behaviors in rats","volume":"138","author":"Thompson","year":"2018","journal-title":"Neuropharmacology"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"101891","DOI":"10.1016\/j.pneurobio.2020.101891","article-title":"Left and right hemispheric lateralization of the amygdala in pain","volume":"196","author":"Allen","year":"2020","journal-title":"Prog. Neurobiol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"1744","DOI":"10.1186\/1744-8069-4-24","article-title":"Hemispheric lateralization of a molecular signal for pain modulation in the amygdala","volume":"4","author":"Carrasquillo","year":"2008","journal-title":"Mol. Pain"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"2253","DOI":"10.1152\/jn.00166.2009","article-title":"Hemispheric Lateralization of Pain Processing by Amygdala Neurons","volume":"102","author":"Ji","year":"2009","journal-title":"J. Neurophysiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1016\/0304-3959(92)90041-9","article-title":"An experimental model for peripheral neuropathy produced by segmental spinal nerve ligation in the rat","volume":"50","author":"Kim","year":"1992","journal-title":"Pain"},{"key":"ref_55","doi-asserted-by":"crossref","unstructured":"Ji, G., Yakhnitsa, V., Kiritoshi, T., Presto, P., and Neugebauer, V. (2018). Fear extinction learning ability predicts neuropathic pain behaviors and amygdala activity in male rats. Mol. Pain, 14.","DOI":"10.1177\/1744806918804441"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1016\/S0149-7634(05)80090-9","article-title":"Staircase bioassay: The up-and-down method","volume":"15","author":"Dixon","year":"1991","journal-title":"Neurosci. Biobehav. Rev."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"168","DOI":"10.1038\/nature05453","article-title":"Genome-wide atlas of gene expression in the adult mouse brain","volume":"445","author":"Lein","year":"2007","journal-title":"Nature"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"268","DOI":"10.1002\/cne.21495","article-title":"Topography of thalamic and parabrachial calcitonin gene-related peptide (CGRP) immunoreactive neurons projecting to subnuclei of the amygdala and extended amygdala","volume":"505","author":"Linke","year":"2007","journal-title":"J. Comp. Neurol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"747","DOI":"10.1097\/j.pain.0000000000000830","article-title":"Divergent functions of the left and right central amygdala in visceral nociception","volume":"158","author":"Sadler","year":"2017","journal-title":"Pain"},{"key":"ref_60","doi-asserted-by":"crossref","unstructured":"Yan, Y., Zhang, L., Zhu, T., Deng, S., Ma, B., Lv, H., Shan, X., Cheng, H., Jiang, K., and Zhang, T. (2020). Reconsolidation of a post-ingestive nutrient memory requires mTOR in the central amygdala. Mol. Psychiatry, 1\u201317.","DOI":"10.1038\/s41380-020-00874-5"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.conb.2017.02.002","article-title":"The C1q complement family of synaptic organizers: Not just complementary","volume":"45","author":"Yuzaki","year":"2017","journal-title":"Curr. Opin. Neurobiol."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1146\/annurev-physiol-021317-121322","article-title":"Two Classes of Secreted Synaptic Organizers in the Central Nervous System","volume":"80","author":"Yuzaki","year":"2018","journal-title":"Annu. Rev. Physiol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"138","DOI":"10.1016\/j.tins.2016.12.004","article-title":"A GluD Coming-Of-Age Story","volume":"40","author":"Yuzaki","year":"2017","journal-title":"Trends Neurosci."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"1744806918766745","DOI":"10.1177\/1744806918766745","article-title":"Downregulation of neuroligin1 ameliorates postoperative pain through inhibiting neuroligin1\/postsynaptic density 95-mediated synaptic targeting of alpha-amino-3-hydroxy-5-methyl-4-isoxazole propionate receptor GluA1 subunits in rat dorsal horns","volume":"14","author":"Guo","year":"2018","journal-title":"Mol. Pain"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1097\/ALN.0000000000000809","article-title":"Neuropathic Allodynia Involves Spinal Neurexin-1beta-dependent Neuroligin-1\/Postsynaptic Density-95\/NR2B Cascade in Rats","volume":"123","author":"Lin","year":"2015","journal-title":"Anesthesiology"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.neuroscience.2018.06.047","article-title":"Activity-dependent Synaptic Recruitment of Neuroligin 1 in Spinal Dorsal Horn Contributed to Inflammatory Pain","volume":"388","author":"Zhao","year":"2018","journal-title":"Neuroscience"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"5425","DOI":"10.1523\/JNEUROSCI.4473-08.2009","article-title":"Activity-dependent repression of Cbln1 expression: Mechanism for developmental and homeo-static regulation of synapses in the cerebellum","volume":"29","author":"Iijima","year":"2009","journal-title":"J. Neurosci."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1601","DOI":"10.1016\/j.cell.2011.11.028","article-title":"SAM68 Regulates Neuronal Activity-Dependent Alternative Splicing of Neurexin-1","volume":"147","author":"Iijima","year":"2011","journal-title":"Cell"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"717","DOI":"10.1111\/j.1471-4159.2012.07648.x","article-title":"The Cbln family of proteins interact with multiple signaling pathways","volume":"121","author":"Wei","year":"2012","journal-title":"J. Neurochem."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"3286","DOI":"10.1002\/cne.24278","article-title":"Cerebellins are differentially expressed in selective subsets of neurons throughout the brain","volume":"525","author":"Seigneur","year":"2017","journal-title":"J. Comp. Neurol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Miyazawa, Y., Takahashi, Y., Watabe, A.M., and Kato, F. (2018). Predominant synaptic potentiation and activation in the right central amygdala are independent of bilateral parabrachial activation in the hemilateral trigeminal inflammatory pain model of rats. Mol. Pain, 14.","DOI":"10.1177\/1744806918807102"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"907","DOI":"10.1523\/JNEUROSCI.4081-08.2009","article-title":"Modulation of the dimer interface at ionotropic glutamate-like receptor delta2 by D-serine and extracellular calcium","volume":"29","author":"Hansen","year":"2009","journal-title":"J. Neurosci."}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/10\/10\/2644\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T07:09:11Z","timestamp":1760166551000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/10\/10\/2644"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2021,10,3]]},"references-count":72,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2021,10]]}},"alternative-id":["cells10102644"],"URL":"https:\/\/doi.org\/10.3390\/cells10102644","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,10,3]]}}}