{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,9]],"date-time":"2026-05-09T07:09:25Z","timestamp":1778310565831,"version":"3.51.4"},"reference-count":260,"publisher":"Wiley","issue":"3","license":[{"start":{"date-parts":[[2026,3,19]],"date-time":"2026-03-19T00:00:00Z","timestamp":1773878400000},"content-version":"vor","delay-in-days":18,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,3,1]],"date-time":"2026-03-01T00:00:00Z","timestamp":1772323200000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100000861","name":"Burroughs Wellcome Fund","doi-asserted-by":"publisher","award":["Postdoctoral Enrichment Program (PDEP)"],"award-info":[{"award-number":["Postdoctoral Enrichment Program (PDEP)"]}],"id":[{"id":"10.13039\/100000861","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000025","name":"National Institute of Mental Health","doi-asserted-by":"publisher","award":["MH117628"],"award-info":[{"award-number":["MH117628"]}],"id":[{"id":"10.13039\/100000025","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000025","name":"National Institute of Mental Health","doi-asserted-by":"publisher","award":["MH129970"],"award-info":[{"award-number":["MH129970"]}],"id":[{"id":"10.13039\/100000025","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000026","name":"National Institute on Drug Abuse","doi-asserted-by":"publisher","award":["F32DA062486"],"award-info":[{"award-number":["F32DA062486"]}],"id":[{"id":"10.13039\/100000026","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000065","name":"National Institute of Neurological Disorders and Stroke","doi-asserted-by":"publisher","award":["NS088555"],"award-info":[{"award-number":["NS088555"]}],"id":[{"id":"10.13039\/100000065","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000065","name":"National Institute of Neurological Disorders and Stroke","doi-asserted-by":"publisher","award":["NS126279"],"award-info":[{"award-number":["NS126279"]}],"id":[{"id":"10.13039\/100000065","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100006458","name":"Children's Hospital of Philadelphia","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006458","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Journal of Neurochemistry"],"published-print":{"date-parts":[[2026,3]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>The hippocampal dentate gyrus (DG) has emerged as a cornerstone of engram research. While DG fear\u2010based engrams have been extensively studied, revealing principles of allocation, consolidation, retrieval, and valence switching, engrams encoding context\u2010reward associations, particularly those involving drugs of abuse, remain comparatively underexplored. This knowledge gap has critical implications for understanding addiction, depression, and other disorders involving dysfunctional reward processing. In this review, we first establish the DG's unique anatomical and functional properties that position it as an ideal model system for engram research. We then systematically examine the DG fear engram literature, documenting how decades of contextual fear conditioning studies have elucidated mechanisms of competitive allocation, molecular consolidation, competing extinction ensembles, and context\u2010dependent discrimination versus generalization. Turning to reward engrams, we synthesize emerging evidence demonstrating that drug\u2010context associations are encoded through sparse, distributed ensembles across multiple brain regions including the nucleus accumbens, prefrontal cortex, amygdala, and hippocampus. While these studies establish foundational principles of reward engram allocation and retrieval, critical mechanistic gaps remain, particularly regarding differences between drug\u2010associated and natural reward memories, the neural coding of context versus valence in hippocampal circuits, and the mechanisms underlying drug associated reward memory extinction. Evidence from valence switching studies demonstrates that the DG processes and stores both fear and reward memories within overlapping circuits, exhibiting remarkable plasticity in linking contextual representations to opposing emotional outcomes, a flexibility distinguishing it from structures with hardwired valence encoding. This encoding capacity positions the DG as a promising target for interventions aimed at modifying pathological emotional associations in addiction and trauma\u2010related disorders while preserving contextual specificity. Understanding reward engram mechanisms with the same rigor applied to fear engrams is essential for developing comprehensive frameworks of how DG circuits contribute to memory\u2010related psychopathology and for translating engram research into therapeutic applications.<\/jats:p>\n                  <jats:p>\n                    <jats:boxed-text content-type=\"graphic\" position=\"anchor\">\n                      <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" mimetype=\"image\/png\" position=\"anchor\" specific-use=\"enlarged-web-image\" xlink:href=\"graphic\/jnc70413-gra-0001-m.png\">\n                        <jats:alt-text>image<\/jats:alt-text>\n                      <\/jats:graphic>\n                    <\/jats:boxed-text>\n                  <\/jats:p>","DOI":"10.1111\/jnc.70413","type":"journal-article","created":{"date-parts":[[2026,3,19]],"date-time":"2026-03-19T08:25:21Z","timestamp":1773908721000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Dentate Gyrus Engrams in Fear and Reward: Mechanistic Principles, Critical Gaps, and Paths to Translation"],"prefix":"10.1111","volume":"170","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9474-753X","authenticated-orcid":false,"given":"Lorianna M.","family":"Col\u00f3n","sequence":"first","affiliation":[{"name":"Division of Basic Science, Department of Anesthesiology and Critical Care Children's Hospital of Philadelphia (CHOP)  Philadelphia Pennsylvania USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Oluwatoni A.","family":"Famuyide","sequence":"additional","affiliation":[{"name":"Division of Basic Science, Department of Anesthesiology and Critical Care Children's Hospital of Philadelphia (CHOP)  Philadelphia Pennsylvania USA"},{"name":"Cornell University, College of Arts and Sciences  Ithaca New York USA"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Amelia J.","family":"Eisch","sequence":"additional","affiliation":[{"name":"Division of Basic Science, Department of Anesthesiology and Critical Care Children's Hospital of Philadelphia (CHOP)  Philadelphia Pennsylvania USA"},{"name":"Neuroscience Graduate Group, Biomedical Graduate Studies University of Pennsylvania Perelman School of Medicine (PSoM)  Philadelphia Pennsylvania USA"},{"name":"Department of Anesthesiology and Critical Care PSoM  Philadelphia Pennsylvania USA"},{"name":"Department of Neuroscience PSoM  Philadelphia Pennsylvania USA"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2026,3,19]]},"reference":[{"key":"e_1_2_13_2_1","doi-asserted-by":"publisher","DOI":"10.1101\/lm.027581.112"},{"key":"e_1_2_13_3_1","doi-asserted-by":"publisher","DOI":"10.1002\/cne.901240303"},{"key":"e_1_2_13_4_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0079-6123(07)63001-5"},{"key":"e_1_2_13_5_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.2011.07804.x"},{"key":"e_1_2_13_6_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2019.04.002"},{"key":"e_1_2_13_7_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.70030"},{"key":"e_1_2_13_8_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tins.2018.10.005"},{"key":"e_1_2_13_9_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.nlm.2025.108036"},{"key":"e_1_2_13_10_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4737-03.2004"},{"key":"e_1_2_13_11_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrn2044"},{"key":"e_1_2_13_12_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3029-16.2017"},{"key":"e_1_2_13_13_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2017.12.097"},{"key":"e_1_2_13_14_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2025.06.007"},{"key":"e_1_2_13_15_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41380-020-00888-z"},{"key":"e_1_2_13_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2022.12.006"},{"key":"e_1_2_13_17_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41583-023-00710-z"},{"key":"e_1_2_13_18_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2758"},{"key":"e_1_2_13_19_1","doi-asserted-by":"publisher","DOI":"10.1002\/alz.12071"},{"key":"e_1_2_13_20_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1496-19.2019"},{"key":"e_1_2_13_21_1","doi-asserted-by":"publisher","DOI":"10.1177\/1073858420941528"},{"key":"e_1_2_13_22_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neubiorev.2014.09.016"},{"key":"e_1_2_13_23_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41380-025-03222-7"},{"key":"e_1_2_13_24_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3091-16.2016"},{"key":"e_1_2_13_25_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.20091"},{"key":"e_1_2_13_26_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2019.04.065"},{"key":"e_1_2_13_27_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2022.102552"},{"key":"e_1_2_13_28_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aas9204"},{"key":"e_1_2_13_29_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41583-024-00814-0"},{"key":"e_1_2_13_30_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1173215"},{"key":"e_1_2_13_31_1","doi-asserted-by":"publisher","DOI":"10.1037\/0735-7044.116.1.169"},{"key":"e_1_2_13_32_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0238-14.2014"},{"key":"e_1_2_13_33_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.brainres.2014.11.005"},{"key":"e_1_2_13_34_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.3002679"},{"key":"e_1_2_13_35_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2022.07.004"},{"key":"e_1_2_13_36_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.70040"},{"key":"e_1_2_13_37_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41386-022-01477-0"},{"key":"e_1_2_13_38_1","doi-asserted-by":"publisher","DOI":"10.1038\/345718a0"},{"key":"e_1_2_13_39_1","doi-asserted-by":"publisher","DOI":"10.1037\/0033-2909.96.3.518"},{"key":"e_1_2_13_40_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1395-16.2016"},{"key":"e_1_2_13_41_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jbc.2022.101866"},{"key":"e_1_2_13_42_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrn2822"},{"key":"e_1_2_13_43_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2014.05.018"},{"key":"e_1_2_13_44_1","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2017.00023"},{"key":"e_1_2_13_45_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2018.00168"},{"key":"e_1_2_13_46_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-025-09619-2"},{"key":"e_1_2_13_47_1","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2012.00113"},{"key":"e_1_2_13_48_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0340-23.2023"},{"key":"e_1_2_13_49_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.101428.3"},{"key":"e_1_2_13_50_1","doi-asserted-by":"publisher","DOI":"10.1016\/0169-328X(92)90051-C"},{"key":"e_1_2_13_51_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-neuro-062111-150500"},{"key":"e_1_2_13_52_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2599"},{"key":"e_1_2_13_53_1","doi-asserted-by":"publisher","DOI":"10.3758\/s13420-016-0218-1"},{"key":"e_1_2_13_54_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1914-12.2012"},{"key":"e_1_2_13_55_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2009.11.031"},{"key":"e_1_2_13_56_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.adv4420"},{"key":"e_1_2_13_57_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2114230119"},{"key":"e_1_2_13_58_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.23-27-09116.2003"},{"key":"e_1_2_13_59_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1505-16.2016"},{"key":"e_1_2_13_60_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1214985"},{"key":"e_1_2_13_61_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2018.06.008"},{"key":"e_1_2_13_62_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2014.00118"},{"key":"e_1_2_13_63_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2024.115022"},{"key":"e_1_2_13_64_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2025.09.005"},{"key":"e_1_2_13_65_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.expneurol.2022.114224"},{"key":"e_1_2_13_66_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1913658116"},{"key":"e_1_2_13_67_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00213-018-5024-3"},{"key":"e_1_2_13_68_1","doi-asserted-by":"publisher","DOI":"10.3389\/fpsyt.2016.00024"},{"key":"e_1_2_13_69_1","volume-title":"Engrams, Advances in Neurobiology","author":"Gr\u00e4ff J.","year":"2024"},{"key":"e_1_2_13_70_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.65764"},{"key":"e_1_2_13_71_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnmol.2024.1342622"},{"key":"e_1_2_13_72_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-32246-8"},{"key":"e_1_2_13_73_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2013.03.025"},{"key":"e_1_2_13_74_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neurobiolaging.2021.01.019"},{"key":"e_1_2_13_75_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-14498-4"},{"key":"e_1_2_13_76_1","doi-asserted-by":"publisher","DOI":"10.1038\/nm.4491"},{"key":"e_1_2_13_77_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41380-023-02137-5"},{"key":"e_1_2_13_78_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2018.05.059"},{"key":"e_1_2_13_79_1","doi-asserted-by":"publisher","DOI":"10.1038\/16046"},{"key":"e_1_2_13_80_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-018-0191-2"},{"key":"e_1_2_13_81_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41583-019-0260-z"},{"key":"e_1_2_13_82_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.semcdb.2021.05.033"},{"key":"e_1_2_13_83_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1139438"},{"key":"e_1_2_13_84_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1164139"},{"key":"e_1_2_13_85_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2017.03.025"},{"key":"e_1_2_13_86_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.20316"},{"key":"e_1_2_13_87_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn887"},{"key":"e_1_2_13_88_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.nlm.2008.06.008"},{"key":"e_1_2_13_89_1","doi-asserted-by":"publisher","DOI":"10.1006\/frne.1993.1006"},{"key":"e_1_2_13_90_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3327-14.2014"},{"key":"e_1_2_13_91_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2010.06.033"},{"key":"e_1_2_13_92_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-023-01557-4"},{"key":"e_1_2_13_93_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.coemr.2022.100378"},{"key":"e_1_2_13_94_1","doi-asserted-by":"publisher","DOI":"10.1002\/jnr.25025"},{"key":"e_1_2_13_95_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41380-021-01321-9"},{"key":"e_1_2_13_96_1","doi-asserted-by":"publisher","DOI":"10.1038\/srep08388"},{"key":"e_1_2_13_97_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41422-025-01157-w"},{"key":"e_1_2_13_98_1","doi-asserted-by":"publisher","DOI":"10.1139\/jpn.100015"},{"key":"e_1_2_13_99_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0056-17.2017"},{"key":"e_1_2_13_100_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrn4000"},{"key":"e_1_2_13_101_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aaw4325"},{"key":"e_1_2_13_102_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1067020"},{"key":"e_1_2_13_103_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2014.03.001"},{"key":"e_1_2_13_104_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.0912931107"},{"key":"e_1_2_13_105_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aas9875"},{"key":"e_1_2_13_106_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2012.12.038"},{"key":"e_1_2_13_107_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.3262"},{"key":"e_1_2_13_108_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.4414"},{"key":"e_1_2_13_109_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-020-15121-2"},{"key":"e_1_2_13_110_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aam6808"},{"key":"e_1_2_13_111_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.nlm.2015.10.008"},{"key":"e_1_2_13_112_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neulet.2021.136176"},{"key":"e_1_2_13_113_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-025-08993-1"},{"key":"e_1_2_13_114_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopsych.2020.11.008"},{"key":"e_1_2_13_115_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.3232"},{"key":"e_1_2_13_116_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2364"},{"key":"e_1_2_13_117_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature14622"},{"key":"e_1_2_13_118_1","doi-asserted-by":"publisher","DOI":"10.1101\/lm.698107"},{"key":"e_1_2_13_119_1","first-page":"753","article-title":"Distinct Hippocampal Engrams Control Extinction and Relapse of Fear Memory","volume":"22","author":"Lacagnina A. F.","year":"2019","journal-title":"Nature"},{"key":"e_1_2_13_120_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.nlm.2020.107164"},{"key":"e_1_2_13_121_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-11799-1"},{"key":"e_1_2_13_122_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopsych.2022.07.021"},{"key":"e_1_2_13_123_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.pneurobio.2024.102601"},{"key":"e_1_2_13_124_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2786-20.2021"},{"key":"e_1_2_13_125_1","doi-asserted-by":"publisher","DOI":"10.1038\/srep36885"},{"key":"e_1_2_13_126_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopsych.2021.04.010"},{"key":"e_1_2_13_127_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1135801"},{"key":"e_1_2_13_128_1","doi-asserted-by":"publisher","DOI":"10.4103\/NRR.NRR"},{"key":"e_1_2_13_129_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature07319"},{"key":"e_1_2_13_130_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41398-024-03068-7"},{"key":"e_1_2_13_131_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature11028"},{"key":"e_1_2_13_132_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neubiorev.2024.105574"},{"key":"e_1_2_13_133_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41386-021-01067-6"},{"key":"e_1_2_13_134_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2022.805132"},{"key":"e_1_2_13_135_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.addbeh.2023.107782"},{"key":"e_1_2_13_136_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.2007.05984.x"},{"key":"e_1_2_13_137_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2013.0161"},{"key":"e_1_2_13_138_1","article-title":"Role of Immediate\u2010Early Genes in Synaptic Plasticity and Neuronal Ensembles Underlying the Memory Trace","volume":"8","author":"Minatohara K.","year":"2015","journal-title":"Frontiers in Molecular Neuroscience"},{"key":"e_1_2_13_139_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2020.632019"},{"key":"e_1_2_13_140_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2018.11.022"},{"key":"e_1_2_13_141_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2025.08.024"},{"key":"e_1_2_13_142_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0675-20.2020"},{"key":"e_1_2_13_143_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2013.00034"},{"key":"e_1_2_13_144_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2023.112149"},{"key":"e_1_2_13_145_1","doi-asserted-by":"publisher","DOI":"10.1007\/s11229-021-03216-2"},{"key":"e_1_2_13_146_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2012.01.046"},{"key":"e_1_2_13_147_1","doi-asserted-by":"publisher","DOI":"10.1111\/jnc.15297"},{"key":"e_1_2_13_148_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2022.102568"},{"key":"e_1_2_13_149_1","doi-asserted-by":"publisher","DOI":"10.1038\/npp.2015.358"},{"key":"e_1_2_13_150_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature14366"},{"key":"e_1_2_13_151_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.semcdb.2017.09.006"},{"key":"e_1_2_13_152_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2572-07.2007"},{"key":"e_1_2_13_153_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4256-09.2010"},{"key":"e_1_2_13_154_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neures.2010.12.007"},{"key":"e_1_2_13_155_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2012.02.062"},{"key":"e_1_2_13_156_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2023.10.074"},{"key":"e_1_2_13_157_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-031-62983-9_15"},{"key":"e_1_2_13_158_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41386-022-01472-5"},{"key":"e_1_2_13_159_1","doi-asserted-by":"publisher","DOI":"10.1038\/npp.2016.73"},{"key":"e_1_2_13_160_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1160-24.2024"},{"key":"e_1_2_13_161_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.22756"},{"key":"e_1_2_13_162_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0684-15.2015"},{"key":"e_1_2_13_163_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2018.11.029"},{"key":"e_1_2_13_164_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2006.08.024"},{"key":"e_1_2_13_165_1","doi-asserted-by":"publisher","DOI":"10.1186\/s12915-016-0261-6"},{"key":"e_1_2_13_166_1","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.adg9921"},{"key":"e_1_2_13_167_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41386-022-01342-0"},{"key":"e_1_2_13_168_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1208049"},{"key":"e_1_2_13_169_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1239073"},{"key":"e_1_2_13_170_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature14514"},{"key":"e_1_2_13_171_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-09960-x"},{"key":"e_1_2_13_172_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnsyn.2021.661476"},{"key":"e_1_2_13_173_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2018.10.013"},{"key":"e_1_2_13_174_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aaf0594"},{"key":"e_1_2_13_175_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature13725"},{"key":"e_1_2_13_176_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1143839"},{"key":"e_1_2_13_177_1","doi-asserted-by":"publisher","DOI":"10.1016\/bs.irn.2016.02.018"},{"key":"e_1_2_13_178_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-021-00825-5"},{"key":"e_1_2_13_179_1","article-title":"The 21st Century Engram Wiley Interdisciplinary Reviews","volume":"14","author":"Robins S.","year":"2023","journal-title":"Cognitive Science"},{"key":"e_1_2_13_180_1","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1098-1063(1996)6:6<601::AID-HIPO5>3.0.CO;2-J"},{"key":"e_1_2_13_181_1","first-page":"E9972","article-title":"Silent Memory Engrams as the Basis for Retrograde Amnesia","volume":"114","author":"Roy D. S.","year":"2017","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"key":"e_1_2_13_182_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-29384-4"},{"key":"e_1_2_13_183_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2021.01.006"},{"key":"e_1_2_13_184_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41583-021-00548-3"},{"key":"e_1_2_13_185_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aaa5542"},{"key":"e_1_2_13_186_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.3131879"},{"key":"e_1_2_13_187_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature09817"},{"key":"e_1_2_13_188_1","doi-asserted-by":"publisher","DOI":"10.1186\/1756-6606-5-32"},{"key":"e_1_2_13_189_1","doi-asserted-by":"publisher","DOI":"10.1016\/bs.apha.2020.09.006"},{"key":"e_1_2_13_190_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-025-61004-9"},{"key":"e_1_2_13_191_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.nlm.2024.107952"},{"key":"e_1_2_13_192_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2014.10.018"},{"key":"e_1_2_13_193_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2020.107864"},{"key":"e_1_2_13_194_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0094-11.2011"},{"key":"e_1_2_13_195_1","article-title":"Hilar Mossy Cells of the Dentate Gyrus: A Historical Perspective","volume":"6","author":"Scharfman H. E.","year":"2012","journal-title":"Frontiers in Neural Circuits"},{"key":"e_1_2_13_196_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.nlm.2015.09.005"},{"key":"e_1_2_13_197_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1787-07.2007"},{"key":"e_1_2_13_198_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2020.00839"},{"key":"e_1_2_13_199_1","unstructured":"Semon R. W.1921.The Mneme."},{"key":"e_1_2_13_200_1","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-022-03906-8"},{"key":"e_1_2_13_201_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.neuro.21.1.127"},{"key":"e_1_2_13_202_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1174519"},{"key":"e_1_2_13_203_1","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-018-0043-5"},{"key":"e_1_2_13_204_1","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a018937"},{"key":"e_1_2_13_205_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2022.920667"},{"key":"e_1_2_13_206_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuropharm.2025.110684"},{"key":"e_1_2_13_207_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41583-021-00479-z"},{"key":"e_1_2_13_208_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-025-01985-4"},{"key":"e_1_2_13_209_1","doi-asserted-by":"publisher","DOI":"10.1155\/2008\/269097"},{"key":"e_1_2_13_210_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2014.03.058"},{"key":"e_1_2_13_211_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2016.01.024"},{"key":"e_1_2_13_212_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.02.055"},{"key":"e_1_2_13_213_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tins.2016.02.003"},{"key":"e_1_2_13_214_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-34114-x"},{"key":"e_1_2_13_215_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2024.113850"},{"key":"e_1_2_13_216_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.21920"},{"key":"e_1_2_13_217_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2020.11.017"},{"key":"e_1_2_13_218_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.adk6742"},{"key":"e_1_2_13_219_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aat5397"},{"key":"e_1_2_13_220_1","doi-asserted-by":"publisher","DOI":"10.1177\/1179069518815942"},{"key":"e_1_2_13_221_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2022.01.019"},{"key":"e_1_2_13_222_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-39718-5"},{"key":"e_1_2_13_223_1","article-title":"Navigating Uncertainty: Reward Location Variability Induces Reorganization of Hippocampal Spatial Representations","author":"Tessereau C.","year":"2025","journal-title":"bioRxiv"},{"key":"e_1_2_13_224_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopsych.2024.06.008"},{"key":"e_1_2_13_225_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-023-01551-w"},{"key":"e_1_2_13_226_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2015.08.002"},{"key":"e_1_2_13_227_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41583-018-0031-2"},{"key":"e_1_2_13_228_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2015.07.009"},{"key":"e_1_2_13_229_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2023.03.010"},{"key":"e_1_2_13_230_1","doi-asserted-by":"publisher","DOI":"10.3390\/cells14020143"},{"key":"e_1_2_13_231_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.4250"},{"key":"e_1_2_13_232_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-018-0459-6"},{"key":"e_1_2_13_233_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.ado8316"},{"key":"e_1_2_13_234_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1304-25.2025"},{"key":"e_1_2_13_235_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-031-62983-9_4"},{"key":"e_1_2_13_236_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-019-0389-0"},{"key":"e_1_2_13_237_1","doi-asserted-by":"publisher","DOI":"10.1111\/acer.15412"},{"key":"e_1_2_13_238_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.isci.2024.109408"},{"key":"e_1_2_13_239_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0918-19.2019"},{"key":"e_1_2_13_240_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0140-16.2016"},{"key":"e_1_2_13_241_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2018.01.026"},{"key":"e_1_2_13_242_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopsych.2015.08.006"},{"key":"e_1_2_13_243_1","doi-asserted-by":"publisher","DOI":"10.1101\/lm.046698.117"},{"key":"e_1_2_13_244_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3761-16.2017"},{"key":"e_1_2_13_245_1","doi-asserted-by":"publisher","DOI":"10.3758\/s13415-022-01013-z"},{"key":"e_1_2_13_246_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-024-08170-w"},{"key":"e_1_2_13_247_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2010.06.068"},{"key":"e_1_2_13_248_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2017.05.047"},{"key":"e_1_2_13_249_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroscience.2012.07.067"},{"key":"e_1_2_13_250_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopsych.2017.04.017"},{"key":"e_1_2_13_251_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-981-15-8763-4_28"},{"key":"e_1_2_13_252_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2018.10.013"},{"key":"e_1_2_13_253_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2014.07.017"},{"key":"e_1_2_13_254_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2016.00218"},{"key":"e_1_2_13_255_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2023.1151877"},{"issue":"1","key":"e_1_2_13_256_1","first-page":"9","article-title":"Memory Engram Stability and Flexibility","volume":"50","author":"Zaki Y.","year":"2024","journal-title":"Neuropsychopharmacology"},{"key":"e_1_2_13_257_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-022-04912-w"},{"key":"e_1_2_13_258_1","doi-asserted-by":"publisher","DOI":"10.1002\/ctm2.1196"},{"key":"e_1_2_13_259_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2405"},{"key":"e_1_2_13_260_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-019-0524-y"},{"key":"e_1_2_13_261_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature16954"}],"container-title":["Journal of Neurochemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/jnc.70413","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/jnc.70413","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/jnc.70413","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,5,9]],"date-time":"2026-05-09T06:12:46Z","timestamp":1778307166000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/jnc.70413"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3]]},"references-count":260,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2026,3]]}},"alternative-id":["10.1111\/jnc.70413"],"URL":"https:\/\/doi.org\/10.1111\/jnc.70413","archive":["Portico"],"relation":{"has-review":[{"id-type":"doi","id":"10.1111\/JNC.70413\/v2\/review2","asserted-by":"object"},{"id-type":"doi","id":"10.1111\/JNC.70413\/v1\/decision1","asserted-by":"object"},{"id-type":"doi","id":"10.1111\/JNC.70413\/v1\/review1","asserted-by":"object"},{"id-type":"doi","id":"10.1111\/JNC.70413\/v1\/review2","asserted-by":"object"},{"id-type":"doi","id":"10.1111\/JNC.70413\/v2\/review1","asserted-by":"object"},{"id-type":"doi","id":"10.1111\/JNC.70413\/v2\/decision1","asserted-by":"object"},{"id-type":"doi","id":"10.1111\/JNC.70413\/v2\/response1","asserted-by":"object"}]},"ISSN":["0022-3042","1471-4159"],"issn-type":[{"value":"0022-3042","type":"print"},{"value":"1471-4159","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3]]},"assertion":[{"value":"2025-11-25","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-03-04","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-03-19","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70413"}}