{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T10:56:13Z","timestamp":1781175373711,"version":"3.54.1"},"reference-count":118,"publisher":"Springer Science and Business Media LLC","license":[{"start":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T00:00:00Z","timestamp":1781136000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T00:00:00Z","timestamp":1781136000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"funder":[{"DOI":"10.13039\/501100000156","name":"Fonds de Recherche du Qu\u00e9bec - Sant\u00e9","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000156","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000038","name":"Gouvernement du Canada | Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100021130","name":"Ludmer Centre for Neuroinformatics and Mental Health","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100021130","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Neuropsychopharmacol."],"DOI":"10.1038\/s41386-026-02460-9","type":"journal-article","created":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T09:57:06Z","timestamp":1781171826000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Nucleus accumbens medial shell D1 and D2 medium spiny neurons signal the valence of conditioned stimuli in a sex-dependent manner"],"prefix":"10.1038","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-9194-7862","authenticated-orcid":false,"given":"Gabrielle","family":"Siemonsmeier","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5408-5500","authenticated-orcid":false,"given":"Eshaan S.","family":"Iyer","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8462-5491","authenticated-orcid":false,"given":"Heike","family":"Schuler","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ariel M.","family":"Weinbaum","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kristine","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0221-6588","authenticated-orcid":false,"given":"Jo\u00eblle","family":"Lopez","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-5512-5294","authenticated-orcid":false,"given":"Rosemary C.","family":"Bagot","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,6,11]]},"reference":[{"key":"2460_CR1","doi-asserted-by":"publisher","first-page":"345","DOI":"10.1037\/0022-3514.37.3.345","volume":"37","author":"JA Russell","year":"1979","unstructured":"Russell JA. Affective space is bipolar. J Personal Soc Psychol. 1979;37:345\u201356. https:\/\/doi.org\/10.1037\/0022-3514.37.3.345.","journal-title":"J Personal Soc Psychol"},{"key":"2460_CR2","doi-asserted-by":"publisher","first-page":"1161","DOI":"10.1037\/h0077714","volume":"39","author":"JA Russell","year":"1980","unstructured":"Russell JA. A circumplex model of affect. J Personal Soc Psychol. 1980;39:1161\u201378. https:\/\/doi.org\/10.1037\/h0077714.","journal-title":"J Personal Soc Psychol"},{"key":"2460_CR3","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1016\/s0014-2999(99)00372-6","volume":"375","author":"G Di Chiara","year":"1999","unstructured":"Di Chiara G. Drug addiction as dopamine-dependent associative learning disorder. Eur J Pharm. 1999;375:13\u201330. https:\/\/doi.org\/10.1016\/s0014-2999(99)00372-6.","journal-title":"Eur J Pharm"},{"key":"2460_CR4","doi-asserted-by":"publisher","first-page":"748","DOI":"10.1176\/appi.ajp.2010.09091379","volume":"167","author":"T Insel","year":"2010","unstructured":"Insel T, Cuthbert B, Garvey M, Heinssen R, Pine DS, Quinn K, et al. Research domain criteria (RDoC): toward a new classification framework for research on mental disorders. Am J Psychiatry. 2010;167:748\u201351. https:\/\/doi.org\/10.1176\/appi.ajp.2010.09091379.","journal-title":"Am J Psychiatry"},{"key":"2460_CR5","doi-asserted-by":"publisher","first-page":"114","DOI":"10.1016\/j.copsyc.2014.12.011","volume":"4","author":"R Admon","year":"2015","unstructured":"Admon R, Pizzagalli DA. Dysfunctional Reward Processing in Depression. Curr Opin Psychol. 2015;4:114\u20138. https:\/\/doi.org\/10.1016\/j.copsyc.2014.12.011.","journal-title":"Curr Opin Psychol"},{"key":"2460_CR6","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1002\/da.22353","volume":"32","author":"P Duits","year":"2015","unstructured":"Duits P, Cath DC, Lissek S, Hox JJ, Hamm AO, Engelhard IM, et al. Updated meta-analysis of classical fear conditioning in the anxiety disorders. Depress Anxiety. 2015;32:239\u201353. https:\/\/doi.org\/10.1002\/da.22353.","journal-title":"Depress Anxiety"},{"key":"2460_CR7","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1007\/7854_2015_376","volume":"27","author":"DM Barch","year":"2016","unstructured":"Barch DM, Pagliaccio D, Luking K. Mechanisms Underlying Motivational Deficits in Psychopathology: Similarities and Differences in Depression and Schizophrenia. Curr Top Behav Neurosci. 2016;27:411\u201349. https:\/\/doi.org\/10.1007\/7854_2015_376.","journal-title":"Curr Top Behav Neurosci"},{"key":"2460_CR8","doi-asserted-by":"publisher","first-page":"105983","DOI":"10.1016\/j.neubiorev.2024.105983","volume":"169","author":"FM Kausche","year":"2025","unstructured":"Kausche FM, Carsten HP, Sobania KM, Riesel A. Fear and safety learning in anxiety- and stress-related disorders: An updated meta-analysis. Neurosci Biobehav Rev. 2025;169:105983 https:\/\/doi.org\/10.1016\/j.neubiorev.2024.105983.","journal-title":"Neurosci Biobehav Rev"},{"key":"2460_CR9","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1037\/a0013503","volume":"64","author":"RM Nesse","year":"2009","unstructured":"Nesse RM, Ellsworth PC. Evolution, emotions, and emotional disorders. Am Psychol. 2009;64:129\u201339. https:\/\/doi.org\/10.1037\/a0013503.","journal-title":"Am Psychol"},{"key":"2460_CR10","doi-asserted-by":"publisher","first-page":"501","DOI":"10.3390\/bs3030501","volume":"3","author":"P Trimmer","year":"2013","unstructured":"Trimmer P, Paul E, Mendl M, McNamara J, Houston A. On the evolution and optimality of mood States. Behav Sci. 2013;3:501\u201321. https:\/\/doi.org\/10.3390\/bs3030501. Sep.","journal-title":"Behav Sci"},{"key":"2460_CR11","doi-asserted-by":"publisher","first-page":"725","DOI":"10.1001\/archneurpsyc.1937.02260220069003","volume":"38","author":"JW Papez","year":"1937","unstructured":"Papez JW. A proposed mechanism of emotion. Arch Neurol Psychiatry. 1937;38:725\u201343. https:\/\/doi.org\/10.1001\/archneurpsyc.1937.02260220069003.","journal-title":"Arch Neurol Psychiatry"},{"key":"2460_CR12","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1146\/annurev-neuro-111020-103314","volume":"46","author":"M Malezieux","year":"2023","unstructured":"Malezieux M, Klein AS, Gogolla N. Neural Circuits for Emotion. Annu Rev Neurosci. 2023;46:211\u201331. https:\/\/doi.org\/10.1146\/annurev-neuro-111020-103314.","journal-title":"Annu Rev Neurosci"},{"key":"2460_CR13","doi-asserted-by":"publisher","first-page":"1552","DOI":"10.1038\/nn.3813","volume":"17","author":"J Xiu","year":"2014","unstructured":"Xiu J, Zhang Q, Zhou T, Zhou T, Chen Y, Hu H. Visualizing an emotional valence map in the limbic forebrain by TAI-FISH. Nat Neurosci. 2014;17:1552\u20139. https:\/\/doi.org\/10.1038\/nn.3813.","journal-title":"Nat Neurosci"},{"key":"2460_CR14","doi-asserted-by":"publisher","first-page":"625","DOI":"10.1016\/s0896-6273(03)00264-2","volume":"38","author":"B Setlow","year":"2003","unstructured":"Setlow B, Schoenbaum G, Gallagher M. Neural encoding in ventral striatum during olfactory discrimination learning. Neuron. 2003;38:625\u201336. https:\/\/doi.org\/10.1016\/s0896-6273(03)00264-2.","journal-title":"Neuron"},{"key":"2460_CR15","doi-asserted-by":"publisher","first-page":"587","DOI":"10.1016\/j.neuron.2004.12.055","volume":"45","author":"MF Roitman","year":"2005","unstructured":"Roitman MF, Wheeler RA, Carelli RM. Nucleus accumbens neurons are innately tuned for rewarding and aversive taste stimuli, encode their predictors, and are linked to motor output. Neuron. 2005;45:587\u201397. https:\/\/doi.org\/10.1016\/j.neuron.2004.12.055.","journal-title":"Neuron"},{"key":"2460_CR16","doi-asserted-by":"publisher","first-page":"622","DOI":"10.1038\/nature09159","volume":"466","author":"AV Kravitz","year":"2010","unstructured":"Kravitz AV, Freeze BS, Parker PR, Kay K, Thwin MT, Deisseroth K, Kreitzer AC. Regulation of parkinsonian motor behaviours by optogenetic control of basal ganglia circuitry. Nature. 2010;466:622\u20136. https:\/\/doi.org\/10.1038\/nature09159.","journal-title":"Nature"},{"key":"2460_CR17","doi-asserted-by":"publisher","first-page":"896","DOI":"10.1016\/j.neuron.2010.05.011","volume":"66","author":"T Hikida","year":"2010","unstructured":"Hikida T, Kimura K, Wada N, Funabiki K, Nakanishi S. Distinct roles of synaptic transmission in direct and indirect striatal pathways to reward and aversive behavior. Neuron. 2010;66:896\u2013907. https:\/\/doi.org\/10.1016\/j.neuron.2010.05.011.","journal-title":"Neuron"},{"key":"2460_CR18","doi-asserted-by":"publisher","first-page":"816","DOI":"10.1038\/nn.3100","volume":"15","author":"AV Kravitz","year":"2012","unstructured":"Kravitz AV, Tye LD, Kreitzer AC. Distinct roles for direct and indirect pathway striatal neurons in reinforcement. Nat Neurosci. 2012;15:816\u20138. https:\/\/doi.org\/10.1038\/nn.3100.","journal-title":"Nat Neurosci"},{"key":"2460_CR19","doi-asserted-by":"publisher","first-page":"385","DOI":"10.1126\/science.1188472","volume":"330","author":"MK Lobo","year":"2010","unstructured":"Lobo MK, Covington HE, Chaudhury D, Friedman AK, Sun H, Damez-Werno D, et al. Cell type-specific loss of BDNF signaling mimics optogenetic control of cocaine reward. Science. 2010;330:385\u201390. https:\/\/doi.org\/10.1126\/science.1188472.","journal-title":"Science"},{"key":"2460_CR20","doi-asserted-by":"publisher","first-page":"2726","DOI":"10.1073\/pnas.1521238113","volume":"113","author":"ES Calipari","year":"2016","unstructured":"Calipari ES, Bagot RC, Purushothaman I, Davidson TJ, Yorgason JT, Pe\u00f1a CJ, et al. In vivo imaging identifies temporal signature of D1 and D2 medium spiny neurons in cocaine reward. Proc Natl Acad Sci USA. 2016;113:2726\u201331. https:\/\/doi.org\/10.1073\/pnas.1521238113.","journal-title":"Proc Natl Acad Sci USA"},{"key":"2460_CR21","doi-asserted-by":"publisher","first-page":"e0207694","DOI":"10.1371\/journal.pone.0207694","volume":"13","author":"SL Cole","year":"2018","unstructured":"Cole SL, Robinson MJF, Berridge KC. Optogenetic self-stimulation in the nucleus accumbens: D1 reward versus D2 ambivalence. PLoS One. 2018;13:e0207694 https:\/\/doi.org\/10.1371\/journal.pone.0207694.","journal-title":"PLoS One"},{"key":"2460_CR22","doi-asserted-by":"publisher","first-page":"3729","DOI":"10.1016\/j.celrep.2020.02.095","volume":"30","author":"CK Lafferty","year":"2020","unstructured":"Lafferty CK, Yang AK, Mendoza JA, Britt JP. Nucleus Accumbens Cell Type- and Input-Specific Suppression of Unproductive Reward Seeking. Cell Rep. 2020;30:3729\u201342.e3. https:\/\/doi.org\/10.1016\/j.celrep.2020.02.095.","journal-title":"Cell Rep"},{"key":"2460_CR23","doi-asserted-by":"publisher","first-page":"7520","DOI":"10.1126\/science.adp7520","volume":"386","author":"Z Tian","year":"2024","unstructured":"Tian Z, Song J, Zhao X, Zhou Y, Chen X, Le Q, et al. The interhemispheric amygdala-accumbens circuit encodes negative valence in mice. Science. 2024;386:7520 https:\/\/doi.org\/10.1126\/science.adp7520.","journal-title":"Science"},{"key":"2460_CR24","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms11829","volume":"7","author":"C Soares-Cunha","year":"2016","unstructured":"Soares-Cunha C, Coimbra B, David-Pereira A, Borges S, Pinto L, Costa P, et al. Activation of D2 dopamine receptor-expressing neurons in the nucleus accumbens increases motivation. Nat Commun. 2016;7:11829. https:\/\/doi.org\/10.1038\/ncomms11829.","journal-title":"Nat Commun"},{"key":"2460_CR25","doi-asserted-by":"publisher","first-page":"370","DOI":"10.1016\/j.neubiorev.2016.05.021","volume":"68","author":"C Soares-Cunha","year":"2016","unstructured":"Soares-Cunha C, Coimbra B, Sousa N, Rodrigues AJ. Reappraising striatal D1- and D2-neurons in reward and aversion. Neurosci Biobehav Rev. 2016;68:370\u201386. https:\/\/doi.org\/10.1016\/j.neubiorev.2016.05.021.","journal-title":"Neurosci Biobehav Rev"},{"key":"2460_CR26","doi-asserted-by":"publisher","first-page":"3241","DOI":"10.1038\/s41380-019-0484-3","volume":"25","author":"C Soares-Cunha","year":"2020","unstructured":"Soares-Cunha C, de Vasconcelos NAP, Coimbra B, Domingues AV, Silva JM, Loureiro-Campos E, et al. Nucleus accumbens medium spiny neurons subtypes signal both reward and aversion. Mol Psychiatry. 2020;25:3241\u201355. https:\/\/doi.org\/10.1038\/s41380-019-0484-3.","journal-title":"Mol Psychiatry"},{"key":"2460_CR27","doi-asserted-by":"publisher","first-page":"110380","DOI":"10.1016\/j.celrep.2022.110380","volume":"38","author":"C Soares-Cunha","year":"2022","unstructured":"Soares-Cunha C, Domingues AV, Correia R, Coimbra B, Vieitas-Gaspar N, de Vasconcelos N, et al. Distinct role of nucleus accumbens D2-MSN projections to ventral pallidum in different phases of motivated behavior. Cell Rep. 2022;38:110380 https:\/\/doi.org\/10.1016\/j.celrep.2022.110380.","journal-title":"Cell Rep"},{"key":"2460_CR28","doi-asserted-by":"publisher","first-page":"312","DOI":"10.1111\/jnc.16063","volume":"168","author":"C Deseyve","year":"2024","unstructured":"Deseyve C, Domingues AV, Carvalho TTA, Armada G, Correia R, Vieitas-Gaspar N, et al. Nucleus accumbens neurons dynamically respond to appetitive and aversive associative learning. J Neurochem. 2024;168:312\u201327. https:\/\/doi.org\/10.1111\/jnc.16063.","journal-title":"J Neurochem"},{"key":"2460_CR29","doi-asserted-by":"publisher","first-page":"835","DOI":"10.1016\/j.neuron.2023.11.023","volume":"112","author":"JE Zachry","year":"2024","unstructured":"Zachry JE, Kutlu MG, Yoon HJ, Leonard MZ, Chev\u00e9e M, Patel DD, et al. D1 and D2 medium spiny neurons in the nucleus accumbens core have distinct and valence-independent roles in learning. Neuron. 2024;112:835\u201349.e7. https:\/\/doi.org\/10.1016\/j.neuron.2023.11.023.","journal-title":"Neuron"},{"key":"2460_CR30","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-55269-9","volume":"16","author":"AV Domingues","year":"2025","unstructured":"Domingues AV, Carvalho TTA, Martins GJ, Correia R, Coimbra B, Bastos-Gon\u00e7alves R, et al. Dynamic representation of appetitive and aversive stimuli in nucleus accumbens shell D1- and D2-medium spiny neurons. Nat Commun. 2025;16:59. https:\/\/doi.org\/10.1038\/s41467-024-55269-9.","journal-title":"Nat Commun"},{"key":"2460_CR31","doi-asserted-by":"publisher","first-page":"785","DOI":"10.1001\/archgenpsychiatry.2009.36","volume":"66","author":"S Seedat","year":"2009","unstructured":"Seedat S, Scott KM, Angermeyer MC, Berglund P, Bromet EJ, Brugha TS, et al. Cross-national associations between gender and mental disorders in the World Health Organization World Mental Health Surveys. Arch Gen Psychiatry. 2009;66:785\u201395. https:\/\/doi.org\/10.1001\/archgenpsychiatry.2009.36.","journal-title":"Arch Gen Psychiatry"},{"key":"2460_CR32","doi-asserted-by":"publisher","first-page":"106","DOI":"10.1016\/j.yhbeh.2015.04.002","volume":"76","author":"LY Maeng","year":"2015","unstructured":"Maeng LY, Milad MR. Sex differences in anxiety disorders: Interactions between fear, stress, and gonadal hormones. Horm Behav. 2015;76:106\u201317. https:\/\/doi.org\/10.1016\/j.yhbeh.2015.04.002.","journal-title":"Horm Behav"},{"key":"2460_CR33","doi-asserted-by":"publisher","first-page":"2661","DOI":"10.2174\/0929867323666161202092908","volume":"24","author":"R Agabio","year":"2017","unstructured":"Agabio R, Pisanu C, Gessa GL, Franconi F. Sex Differences in Alcohol Use Disorder. Curr Med Chem. 2017;24:2661\u201370. https:\/\/doi.org\/10.2174\/0929867323666161202092908.","journal-title":"Curr Med Chem"},{"key":"2460_CR34","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/j.cpr.2017.10.012","volume":"66","author":"RK McHugh","year":"2018","unstructured":"McHugh RK, Votaw VR, Sugarman DE, Greenfield SF. Sex and gender differences in substance use disorders. Clin Psychol Rev. 2018;66:12\u201323. https:\/\/doi.org\/10.1016\/j.cpr.2017.10.012.","journal-title":"Clin Psychol Rev"},{"key":"2460_CR35","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1007\/s11920-023-01421-z","volume":"25","author":"LV Hiscox","year":"2023","unstructured":"Hiscox LV, Sharp TH, Olff M, Seedat S, Halligan SL. Sex-Based Contributors to and Consequences of Post-traumatic Stress Disorder. Curr Psychiatry Rep. 2023;25:233\u201345. https:\/\/doi.org\/10.1007\/s11920-023-01421-z.","journal-title":"Curr Psychiatry Rep"},{"key":"2460_CR36","doi-asserted-by":"publisher","unstructured":"Schuler H, Iyer ES, Siemonsmeier G, Weinbaum AM, Vitaro P, Shen S, et al. Sex-specific exploration accounts for differences in valence learning in male and female mice. Elife. 2025;14. https:\/\/doi.org\/10.7554\/eLife.108498","DOI":"10.7554\/eLife.108498"},{"key":"2460_CR37","doi-asserted-by":"publisher","first-page":"917","DOI":"10.1038\/nature02033","volume":"425","author":"S Gong","year":"2003","unstructured":"Gong S, Zheng C, Doughty ML, Losos K, Didkovsky N, Schambra UB, et al. A gene expression atlas of the central nervous system based on bacterial artificial chromosomes. Nature. 2003;425:917\u201325. https:\/\/doi.org\/10.1038\/nature02033.","journal-title":"Nature"},{"key":"2460_CR38","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-019-56408-9","volume":"9","author":"ZT Pennington","year":"2019","unstructured":"Pennington ZT, Dong Z, Feng Y, Vetere LM, Page-Harley L, Shuman T, et al. ezTrack: An open-source video analysis pipeline for the investigation of animal behavior. Sci Rep. 2019;9:19979. https:\/\/doi.org\/10.1038\/s41598-019-56408-9.","journal-title":"Sci Rep"},{"key":"2460_CR39","doi-asserted-by":"publisher","unstructured":"Jacobs DS, Allen MC, Park J, Moghaddam B. Learning of probabilistic punishment as a model of anxiety produces changes in action but not punisher encoding in the dmPFC and VTA. Elife. 2022;11. https:\/\/doi.org\/10.7554\/eLife.78912","DOI":"10.7554\/eLife.78912"},{"key":"2460_CR40","doi-asserted-by":"publisher","first-page":"025003","DOI":"10.1117\/1.NPh.12.2.025003","volume":"12","author":"LJ Keevers","year":"2025","unstructured":"Keevers LJ, Jean-Richard-Dit-Bressel P. Obtaining artifact-corrected signals in fiber photometry via isosbestic signals, robust regression, and dF\/F calculations. Neurophotonics. 2025;12:025003 https:\/\/doi.org\/10.1117\/1.NPh.12.2.025003. Apr.","journal-title":"Neurophotonics"},{"key":"2460_CR41","doi-asserted-by":"publisher","first-page":"370","DOI":"10.1037\/h0026779","volume":"67","author":"RJ Blanchard","year":"1969","unstructured":"Blanchard RJ, Blanchard DC. Crouching as an index of fear. J Comp Physiol Psychol. 1969;67:370\u20135. https:\/\/doi.org\/10.1037\/h0026779.","journal-title":"J Comp Physiol Psychol"},{"key":"2460_CR42","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1037\/h0027676","volume":"68","author":"RJ Blanchard","year":"1969","unstructured":"Blanchard RJ, Blanchard DC. Passive and active reactions to fear-eliciting stimuli. J Comp Physiol Psychol. 1969;68:129\u201335. https:\/\/doi.org\/10.1037\/h0027676.","journal-title":"J Comp Physiol Psychol"},{"key":"2460_CR43","doi-asserted-by":"publisher","first-page":"32","DOI":"10.1037\/h0028589","volume":"77","author":"RC Bolles","year":"1970","unstructured":"Bolles RC. Species-specific defense reactions and avoidance learning. Psychol Rev. 1970;77:32\u201348. https:\/\/doi.org\/10.1037\/h0028589.","journal-title":"Psychol Rev"},{"key":"2460_CR44","doi-asserted-by":"publisher","first-page":"6","DOI":"10.3758\/BF03211975","volume":"4","author":"RC Bolles","year":"1976","unstructured":"Bolles RC, Collier AC. The effect of predictive cues on freezing in rats. Anim Learn Behav. 1976;4:6\u20138. https:\/\/doi.org\/10.3758\/BF03211975.","journal-title":"Anim Learn Behav"},{"key":"2460_CR45","doi-asserted-by":"publisher","first-page":"429","DOI":"10.3758\/BF03199629","volume":"8","author":"ME Bouton","year":"1980","unstructured":"Bouton ME, Bolles RC. Conditioned fear assessed by freezing and by the suppression of three different baselines. Anim Learn Behav. 1980;8:429\u201334. https:\/\/doi.org\/10.3758\/BF03199629.","journal-title":"Anim Learn Behav"},{"key":"2460_CR46","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1016\/s0031-9384(98)00012-2","volume":"65","author":"RJ Blanchard","year":"1998","unstructured":"Blanchard RJ, Hebert MA, Ferrari PF, Palanza P, Figueira R, Blanchard DC, Parmigiani S. Defensive behaviors in wild and laboratory (Swiss) mice: the mouse defense test battery. Physiol Behav. 1998;65:201\u20139. https:\/\/doi.org\/10.1016\/s0031-9384(98)00012-2.","journal-title":"Physiol Behav"},{"key":"2460_CR47","doi-asserted-by":"publisher","first-page":"243","DOI":"10.1016\/0166-4328(93)90120-f","volume":"55","author":"G Williams","year":"1993","unstructured":"Williams G, Rolls E, Leonard C, Stern C. Neuronal responses in the ventral striatum of the behaving macaque. Behav Brain Res. 1993;55:243\u201352. https:\/\/doi.org\/10.1016\/0166-4328(93)90120-f.","journal-title":"Behav Brain Res"},{"key":"2460_CR48","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-35601-x","volume":"13","author":"M Skirzewski","year":"2022","unstructured":"Skirzewski M, Princz-Lebel O, German-Castelan L, Crooks AM, Kim GK, Tarnow SH, et al. Continuous cholinergic-dopaminergic updating in the nucleus accumbens underlies approaches to reward-predicting cues. Nat Commun. 2022;13:7924. https:\/\/doi.org\/10.1038\/s41467-022-35601-x.","journal-title":"Nat Commun"},{"key":"2460_CR49","doi-asserted-by":"publisher","first-page":"113488","DOI":"10.1016\/j.celrep.2023.113488","volume":"42","author":"A Belilos","year":"2023","unstructured":"Belilos A, Gray C, Sanders C, Black D, Mays E, Richie C, et al. Nucleus accumbens local circuit for cue-dependent aversive learning. Cell Rep. 2023;42:113488 https:\/\/doi.org\/10.1016\/j.celrep.2023.113488.","journal-title":"Cell Rep"},{"key":"2460_CR50","doi-asserted-by":"publisher","first-page":"4595","DOI":"10.1523\/JNEUROSCI.12-12-04595.1992","volume":"12","author":"W Schultz","year":"1992","unstructured":"Schultz W, Apicella P, Scarnati E, Ljungberg T. Neuronal activity in monkey ventral striatum related to the expectation of reward. J Neurosci. 1992;12:4595\u2013610. https:\/\/doi.org\/10.1523\/JNEUROSCI.12-12-04595.1992.","journal-title":"J Neurosci"},{"key":"2460_CR51","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1016\/0006-8993(94)90645-9","volume":"638","author":"AM Lavoie","year":"1994","unstructured":"Lavoie AM, Mizumori SJY. Spatial, movement- and reward-sensitive discharge by medial ventral striatum neurons of rats. Brain Res. 1994;638:157\u201368. https:\/\/doi.org\/10.1016\/0006-8993(94)90645-9.","journal-title":"Brain Res"},{"key":"2460_CR52","doi-asserted-by":"publisher","first-page":"2477","DOI":"10.1152\/jn.2001.85.6.2477","volume":"85","author":"OK Hassani","year":"2001","unstructured":"Hassani OK, Cromwell HC, Schultz W. Influence of expectation of different rewards on behavior-related neuronal activity in the striatum. J Neurophysiol. 2001;85:2477\u201389. https:\/\/doi.org\/10.1152\/jn.2001.85.6.2477.","journal-title":"J Neurophysiol"},{"key":"2460_CR53","doi-asserted-by":"publisher","first-page":"2823","DOI":"10.1152\/jn.01014.2002","volume":"89","author":"HC Cromwell","year":"2003","unstructured":"Cromwell HC, Schultz W. Effects of expectations for different reward magnitudes on neuronal activity in primate striatum. J Neurophysiol. 2003;89:2823\u201338. https:\/\/doi.org\/10.1152\/jn.01014.2002.","journal-title":"J Neurophysiol"},{"key":"2460_CR54","doi-asserted-by":"publisher","first-page":"1193","DOI":"10.1523\/JNEUROSCI.3975-04.2005","volume":"25","author":"SA Taha","year":"2005","unstructured":"Taha SA, Fields HL. Encoding of palatability and appetitive behaviors by distinct neuronal populations in the nucleus accumbens. J Neurosci. 2005;25:1193\u2013202. https:\/\/doi.org\/10.1523\/JNEUROSCI.3975-04.2005.","journal-title":"J Neurosci"},{"key":"2460_CR55","doi-asserted-by":"publisher","first-page":"3531","DOI":"10.1038\/s41380-023-02131-x","volume":"28","author":"MCC Guillaumin","year":"2023","unstructured":"Guillaumin MCC, Viskaitis P, Bracey E, Burdakov D, Peleg-Raibstein D. Disentangling the role of NAc D1 and D2 cells in hedonic eating. Mol Psychiatry. 2023;28:3531\u201347. https:\/\/doi.org\/10.1038\/s41380-023-02131-x.","journal-title":"Mol Psychiatry"},{"key":"2460_CR56","doi-asserted-by":"publisher","DOI":"10.1101\/2022.09.18.508426","author":"CE Pedersen","year":"2024","unstructured":"Pedersen CE, Gowrishankar R, Piantadosi SC, Castro DC, Gray MM, Zhou ZC, et al. Medial accumbens shell spiny projection neurons encode relative reward preference. bioRxiv. 2024. https:\/\/doi.org\/10.1101\/2022.09.18.508426.","journal-title":"bioRxiv"},{"key":"2460_CR57","doi-asserted-by":"publisher","first-page":"2777","DOI":"10.1111\/j.1460-9568.2004.03747.x","volume":"20","author":"DIG Wilson","year":"2004","unstructured":"Wilson DIG, Bowman EM. Nucleus accumbens neurons in the rat exhibit differential activity to conditioned reinforcers and primary reinforcers within a second-order schedule of saccharin reinforcement. Eur J Neurosci. 2004;20:2777\u201388. https:\/\/doi.org\/10.1111\/j.1460-9568.2004.03747.x.","journal-title":"Eur J Neurosci"},{"key":"2460_CR58","doi-asserted-by":"publisher","first-page":"1866","DOI":"10.1152\/jn.00658.2003","volume":"91","author":"SM Nicola","year":"2004","unstructured":"Nicola SM, Yun IA, Wakabayashi KT, Fields HL. Firing of nucleus accumbens neurons during the consummatory phase of a discriminative stimulus task depends on previous reward predictive cues. J Neurophysiol. 2004;91:1866\u201382. https:\/\/doi.org\/10.1152\/jn.00658.2003.","journal-title":"J Neurophysiol"},{"key":"2460_CR59","doi-asserted-by":"publisher","first-page":"652","DOI":"10.1152\/jn.00068.2006","volume":"96","author":"X Wan","year":"2006","unstructured":"Wan X, Peoples LL. Firing patterns of accumbal neurons during a pavlovian-conditioned approach task. J Neurophysiol. 2006;96:652\u201360. https:\/\/doi.org\/10.1152\/jn.00068.2006.","journal-title":"J Neurophysiol"},{"key":"2460_CR60","doi-asserted-by":"publisher","first-page":"1341","DOI":"10.1111\/j.1460-9568.2006.04654.x","volume":"23","author":"JJ Day","year":"2006","unstructured":"Day JJ, Wheeler RA, Roitman MF, Carelli RM. Nucleus accumbens neurons encode Pavlovian approach behaviors: evidence from an autoshaping paradigm. Eur J Neurosci. 2006;23:1341\u201351. https:\/\/doi.org\/10.1111\/j.1460-9568.2006.04654.x.","journal-title":"Eur J Neurosci"},{"key":"2460_CR61","doi-asserted-by":"publisher","first-page":"4746","DOI":"10.1523\/JNEUROSCI.0197-10.2010","volume":"30","author":"M Krause","year":"2010","unstructured":"Krause M, German PW, Taha SA, Fields HL. A pause in nucleus accumbens neuron firing is required to initiate and maintain feeding. J Neurosci. 2010;30:4746\u201356. https:\/\/doi.org\/10.1523\/JNEUROSCI.0197-10.2010.","journal-title":"J Neurosci"},{"key":"2460_CR62","doi-asserted-by":"publisher","first-page":"6820","DOI":"10.1523\/JNEUROSCI.6491-10.2011","volume":"31","author":"F Ambroggi","year":"2011","unstructured":"Ambroggi F, Ghazizadeh A, Nicola SM, Fields HL. Roles of nucleus accumbens core and shell in incentive-cue responding and behavioral inhibition. J Neurosci. 2011;31:6820\u201330. https:\/\/doi.org\/10.1523\/JNEUROSCI.6491-10.2011.","journal-title":"J Neurosci"},{"key":"2460_CR63","doi-asserted-by":"publisher","first-page":"1063","DOI":"10.1016\/j.neuron.2015.08.019","volume":"87","author":"R Al-Hasani","year":"2015","unstructured":"Al-Hasani R, McCall JG, Shin G, Gomez AM, Schmitz GP, Bernardi JM, et al. Distinct Subpopulations of Nucleus Accumbens Dynorphin Neurons Drive Aversion and Reward. Neuron. 2015;87:1063\u201377. https:\/\/doi.org\/10.1016\/j.neuron.2015.08.019.","journal-title":"Neuron"},{"key":"2460_CR64","doi-asserted-by":"publisher","first-page":"217","DOI":"10.1523\/JNEUROSCI.3227-05.2006","volume":"26","author":"SA Taha","year":"2006","unstructured":"Taha SA, Fields HL. Inhibitions of nucleus accumbens neurons encode a gating signal for reward-directed behavior. J Neurosci. 2006;26:217\u201322. https:\/\/doi.org\/10.1523\/JNEUROSCI.3227-05.2006.","journal-title":"J Neurosci"},{"key":"2460_CR65","doi-asserted-by":"publisher","first-page":"910","DOI":"10.1016\/j.neuron.2013.04.010","volume":"78","author":"VB McGinty","year":"2013","unstructured":"McGinty VB, Lardeux S, Taha SA, Kim JJ, Nicola SM. Invigoration of reward seeking by cue and proximity encoding in the nucleus accumbens. Neuron. 2013;78:910\u201322. https:\/\/doi.org\/10.1016\/j.neuron.2013.04.010.","journal-title":"Neuron"},{"key":"2460_CR66","doi-asserted-by":"publisher","first-page":"2549","DOI":"10.1152\/jn.00465.2017","volume":"118","author":"SE Morrison","year":"2017","unstructured":"Morrison SE, McGinty VB, du Hoffmann J, Nicola SM. Limbic-motor integration by neural excitations and inhibitions in the nucleus accumbens. J Neurophysiol. 2017;118:2549\u201367. https:\/\/doi.org\/10.1152\/jn.00465.2017.","journal-title":"J Neurophysiol"},{"key":"2460_CR67","doi-asserted-by":"publisher","first-page":"1840","DOI":"10.1152\/jn.00657.2003","volume":"91","author":"SM Nicola","year":"2004","unstructured":"Nicola SM, Yun IA, Wakabayashi KT, Fields HL. Cue-evoked firing of nucleus accumbens neurons encodes motivational significance during a discriminative stimulus task. J Neurophysiol. 2004;91:1840\u201365. https:\/\/doi.org\/10.1152\/jn.00657.2003.","journal-title":"J Neurophysiol"},{"key":"2460_CR68","doi-asserted-by":"publisher","first-page":"e64673","DOI":"10.1371\/journal.pone.0064673","volume":"8","author":"GB Bissonette","year":"2013","unstructured":"Bissonette GB, Burton AC, Gentry RN, Goldstein BL, Hearn TN, Barnett BR, et al. Separate populations of neurons in ventral striatum encode value and motivation. PLoS ONE. 2013;8:e64673 https:\/\/doi.org\/10.1371\/journal.pone.0064673.","journal-title":"PLoS ONE"},{"key":"2460_CR69","doi-asserted-by":"publisher","first-page":"817","DOI":"10.1523\/JNEUROSCI.1107-21.2021","volume":"42","author":"MH Ray","year":"2022","unstructured":"Ray MH, Moaddab M, McDannald MA. Threat and Bidirectional Valence Signaling in the Nucleus Accumbens Core. J Neurosci. 2022;42:817\u201333. https:\/\/doi.org\/10.1523\/JNEUROSCI.1107-21.2021.","journal-title":"J Neurosci"},{"key":"2460_CR70","doi-asserted-by":"publisher","first-page":"709","DOI":"10.1523\/JNEUROSCI.15-01-00709.1995","volume":"15","author":"C Beck","year":"1995","unstructured":"Beck C, Fibiger H. Conditioned fear-induced changes in behavior and in the expression of the immediate early gene c-fos: with and without diazepam pretreatment. J Neurosci. 1995;15:709\u201320. https:\/\/doi.org\/10.1523\/JNEUROSCI.15-01-00709.1995.","journal-title":"J Neurosci"},{"key":"2460_CR71","doi-asserted-by":"publisher","first-page":"1087","DOI":"10.1016\/s0306-4522(96)00632-x","volume":"78","author":"S Campeau","year":"1997","unstructured":"Campeau S, Falls WA, Cullinan WE, Helmreich DL, Davis M, Watson SJ. Elicitation and reduction of fear: behavioural and neuroendocrine indices and brain induction of the immediate-early gene c-fos. Neuroscience. 1997;78:1087\u2013104. https:\/\/doi.org\/10.1016\/s0306-4522(96)00632-x.","journal-title":"Neuroscience"},{"key":"2460_CR72","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/s0166-4328(97)00140-x","volume":"93","author":"TR Stratford","year":"1998","unstructured":"Stratford TR, Swanson CJ, Kelley A. Specific changes in food intake elicited by blockade or activation of glutamate receptors in the nucleus accumbens shell. Behav Brain Res. 1998;93:43\u201350. https:\/\/doi.org\/10.1016\/s0166-4328(97)00140-x.","journal-title":"Behav Brain Res"},{"key":"2460_CR73","doi-asserted-by":"publisher","first-page":"2737","DOI":"10.1523\/JNEUROSCI.2902-19.2020","volume":"40","author":"HM Baumgartner","year":"2020","unstructured":"Baumgartner HM, Cole SL, Olney JJ, Berridge KC. Desire or Dread from Nucleus Accumbens Inhibitions: Reversed by Same-Site Optogenetic Excitations. J Neurosci. 2020;40:2737\u201352. https:\/\/doi.org\/10.1523\/JNEUROSCI.2902-19.2020.","journal-title":"J Neurosci"},{"key":"2460_CR74","doi-asserted-by":"publisher","first-page":"122","DOI":"10.1016\/j.neuropharm.2008.06.075","volume":"56","author":"WA Carlezon","year":"2009","unstructured":"Carlezon WA, Thomas MJ. Biological substrates of reward and aversion: a nucleus accumbens activity hypothesis. Neuropharmacology. 2009;56:122\u201332. https:\/\/doi.org\/10.1016\/j.neuropharm.2008.06.075.","journal-title":"Neuropharmacology"},{"key":"2460_CR75","doi-asserted-by":"publisher","unstructured":"Rivera-Irizarry JK, H\u00e1mor PU, Rowson SA, Asfouri J, Liu D, Zallar LJ, et al. Valence and salience encoding by parallel circuits from the paraventricular thalamus to the nucleus accumbens. bioRxiv. 2023:2023.07.03.547570. https:\/\/doi.org\/10.1101\/2023.07.03.547570","DOI":"10.1101\/2023.07.03.547570"},{"key":"2460_CR76","doi-asserted-by":"publisher","first-page":"1966","DOI":"10.1038\/s41593-024-01748-7","volume":"27","author":"J Muir","year":"2024","unstructured":"Muir J, Iyer ES, Tse YC, Sorensen J, Wu S, Eid RS, et al. Sex-biased neural encoding of threat discrimination in nucleus accumbens afferents drives suppression of reward behavior. Nat Neurosci. 2024;27:1966\u201376. https:\/\/doi.org\/10.1038\/s41593-024-01748-7.","journal-title":"Nat Neurosci"},{"key":"2460_CR77","doi-asserted-by":"publisher","first-page":"e0100242024","DOI":"10.1523\/JNEUROSCI.0100-24.2024","volume":"44","author":"AE Copenhaver","year":"2024","unstructured":"Copenhaver AE, LeGates TA. Sex-Specific Mechanisms Underlie Long-Term Potentiation at Hippocampus\u2192Medium Spiny Neuron Synapses in the Medial Shell of the Nucleus Accumbens. J Neurosci. 2024;44:e0100242024 https:\/\/doi.org\/10.1523\/JNEUROSCI.0100-24.2024.","journal-title":"J Neurosci"},{"key":"2460_CR78","doi-asserted-by":"publisher","first-page":"111335","DOI":"10.1016\/j.pnpbp.2025.111335","volume":"138","author":"M Gauthier","year":"2025","unstructured":"Gauthier M, Hebert LP, Dugast E, Lardeux V, Letort K, Thiriet N, et al. Sex-dependent effects of stress on aIC-NAc circuit neuroplasticity: Role of the endocannabinoid system. Prog Neuropsychopharmacol Biol Psychiatry. 2025;138:111335 https:\/\/doi.org\/10.1016\/j.pnpbp.2025.111335.","journal-title":"Prog Neuropsychopharmacol Biol Psychiatry"},{"key":"2460_CR79","doi-asserted-by":"publisher","DOI":"10.1101\/2025.02.09.637318","author":"KA McLaurin","year":"2025","unstructured":"McLaurin KA, Illenberger JM, Li H, Booze RM, Mactutus CF. Sex-dependent modulation of behavioral allocation via ventral tegmental area-nucleus accumbens shell circuitry. bioRxiv. 2025. https:\/\/doi.org\/10.1101\/2025.02.09.637318.","journal-title":"bioRxiv"},{"key":"2460_CR80","doi-asserted-by":"publisher","DOI":"10.1016\/j.biopsych.2025.10.001","author":"AM Pessoni","year":"2025","unstructured":"Pessoni AM, Blanc-Arabe L, Pancotti L, Mansouri S, D'Angelo M, Huot K, et al. Transcriptional profiling of the cortico-accumbal pathway reveals sex-specific alterations underlying stress susceptibility. Biol Psychiatry. 2025. https:\/\/doi.org\/10.1016\/j.biopsych.2025.10.001.","journal-title":"Biol Psychiatry"},{"key":"2460_CR81","doi-asserted-by":"publisher","first-page":"e1239252025","DOI":"10.1523\/JNEUROSCI.1239-25.2025","volume":"46","author":"L Taniguchi","year":"2026","unstructured":"Taniguchi L, Goodpaster CM, Chen Y, de Carvalho GB, Birnie MT, Paul A, et al. Sex and Stress Govern the Function and Innervation of a Basolateral Amygdala to Nucleus Accumbens Corticotropin-Releasing Hormone\/GABA-Expressing Projection. J Neurosci. 2026;46:e1239252025 https:\/\/doi.org\/10.1523\/JNEUROSCI.1239-25.2025.","journal-title":"J Neurosci"},{"key":"2460_CR82","doi-asserted-by":"publisher","first-page":"565","DOI":"10.1016\/j.neubiorev.2010.07.002","volume":"35","author":"AK Beery","year":"2011","unstructured":"Beery AK, Zucker I. Sex bias in neuroscience and biomedical research. Neurosci Biobehav Rev. 2011;35:565\u201372. https:\/\/doi.org\/10.1016\/j.neubiorev.2010.07.002.","journal-title":"Neurosci Biobehav Rev"},{"key":"2460_CR83","doi-asserted-by":"publisher","first-page":"11817","DOI":"10.1523\/JNEUROSCI.1390-16.2016","volume":"36","author":"RM Shansky","year":"2016","unstructured":"Shansky RM, Woolley CS. Considering Sex as a Biological Variable Will Be Valuable for Neuroscience Research. J Neurosci. 2016;36:11817\u201322. https:\/\/doi.org\/10.1523\/JNEUROSCI.1390-16.2016.","journal-title":"J Neurosci"},{"key":"2460_CR84","doi-asserted-by":"publisher","first-page":"ENEURO.0278-17.","DOI":"10.1523\/ENEURO.0278-17.2017","volume":"4","author":"TR Will","year":"2017","unstructured":"Will TR, Proa\u00f1o SB, Thomas AM, Kunz LM, Thompson KC, Ginnari LA, et al. Problems and Progress regarding Sex Bias and Omission in Neuroscience Research. eNeuro. 2017;4:ENEURO.0278-17.2017 https:\/\/doi.org\/10.1523\/ENEURO.0278-17.2017.","journal-title":"eNeuro"},{"key":"2460_CR85","doi-asserted-by":"publisher","first-page":"457","DOI":"10.1038\/s41593-021-00806-8","volume":"24","author":"RM Shansky","year":"2021","unstructured":"Shansky RM, Murphy AZ. Considering sex as a biological variable will require a global shift in science culture. Nat Neurosci. 2021;24:457\u201364. https:\/\/doi.org\/10.1038\/s41593-021-00806-8.","journal-title":"Nat Neurosci"},{"key":"2460_CR86","doi-asserted-by":"publisher","first-page":"233","DOI":"10.1155\/NP.2002.233","volume":"9","author":"RCB Silva","year":"2002","unstructured":"Silva RCB, Cruz APM, Avanzi V, Landeira-Fernandez J, Brand\u00e3o ML. Distinct contributions of median raphe nucleus to contextual fear conditioning and fear-potentiated startle. Neural Plast. 2002;9:233\u201347. https:\/\/doi.org\/10.1155\/NP.2002.233.","journal-title":"Neural Plast"},{"key":"2460_CR87","doi-asserted-by":"publisher","first-page":"96","DOI":"10.1038\/nature21047","volume":"542","author":"JP Fadok","year":"2017","unstructured":"Fadok JP, Krabbe S, Markovic M, Courtin J, Xu C, Massi L, et al. A competitive inhibitory circuit for selection of active and passive fear responses. Nature. 2017;542:96\u2013100. https:\/\/doi.org\/10.1038\/nature21047.","journal-title":"Nature"},{"key":"2460_CR88","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1038\/s42003-023-04429-6","volume":"6","author":"EB Lind","year":"2023","unstructured":"Lind EB, Sweis BM, Asp AJ, Esguerra M, Silvis KA, David Redish A, et al. A quadruple dissociation of reward-related behaviour in mice across excitatory inputs to the nucleus accumbens shell. Commun Biol. 2023;6:119 https:\/\/doi.org\/10.1038\/s42003-023-04429-6.","journal-title":"Commun Biol"},{"key":"2460_CR89","doi-asserted-by":"publisher","unstructured":"Gruene TM, Flick K, Stefano A, Shea SD, Shansky RM. Sexually divergent expression of active and passive conditioned fear responses in rats. Elife. 2015;4. https:\/\/doi.org\/10.7554\/eLife.11352","DOI":"10.7554\/eLife.11352"},{"key":"2460_CR90","doi-asserted-by":"publisher","first-page":"8079","DOI":"10.1523\/JNEUROSCI.0256-23.2023","volume":"43","author":"MA McDannald","year":"2023","unstructured":"McDannald MA. Pavlovian Fear Conditioning Is More than You Think It Is. J Neurosci. 2023;43:8079\u201387. https:\/\/doi.org\/10.1523\/JNEUROSCI.0256-23.2023.","journal-title":"J Neurosci"},{"key":"2460_CR91","doi-asserted-by":"publisher","unstructured":"Chu A, Gordon NT, DuBois AM, Michel CB, Hanrahan KE, Williams DC, et al. A fear conditioned cue orchestrates a suite of behaviors in rats. Elife. 2024;13https:\/\/doi.org\/10.7554\/eLife.82497","DOI":"10.7554\/eLife.82497"},{"key":"2460_CR92","doi-asserted-by":"publisher","first-page":"1440601","DOI":"10.3389\/fnbeh.2024.1440601","volume":"18","author":"K Chanthongdee","year":"2024","unstructured":"Chanthongdee K, Fuentealba Y, Wahlestedt T, Foulhac L, Kardash T, Coppola A, et al. Comprehensive ethological analysis of fear expression in rats using DeepLabCut and SimBA machine learning model. Front Behav Neurosci. 2024;18:1440601 https:\/\/doi.org\/10.3389\/fnbeh.2024.1440601.","journal-title":"Front Behav Neurosci"},{"key":"2460_CR93","doi-asserted-by":"publisher","first-page":"3887","DOI":"10.1016\/j.neuron.2025.11.003","volume":"113","author":"A Nair","year":"2025","unstructured":"Nair A, Vinograd A, Liu M, Mountoufaris G, Linderman S, Anderson DJ. The neural computation of affective internal states in the hypothalamus: A dynamical systems perspective. Neuron. 2025;113:3887\u2013907. https:\/\/doi.org\/10.1016\/j.neuron.2025.11.003.","journal-title":"Neuron"},{"key":"2460_CR94","doi-asserted-by":"publisher","first-page":"1395","DOI":"10.1016\/j.neuron.2017.08.025","volume":"95","author":"YK Takahashi","year":"2017","unstructured":"Takahashi YK, Batchelor HM, Liu B, Khanna A, Morales M, Schoenbaum G. Dopamine Neurons Respond to Errors in the Prediction of Sensory Features of Expected Rewards. Neuron. 2017;95:1395\u20131405.e3. https:\/\/doi.org\/10.1016\/j.neuron.2017.08.025.","journal-title":"Neuron"},{"key":"2460_CR95","doi-asserted-by":"publisher","first-page":"767","DOI":"10.1016\/s0306-4522(97)00327-8","volume":"82","author":"XY Lu","year":"1997","unstructured":"Lu XY, Behnam Ghasemzadeh M, Kalivas PW. Expression of D1 receptor, D2 receptor, substance P and enkephalin messenger RNAs in the neurons projecting from the nucleus accumbens. Neuroscience. 1997;82:767\u201380. https:\/\/doi.org\/10.1016\/s0306-4522(97)00327-8.","journal-title":"Neuroscience"},{"key":"2460_CR96","doi-asserted-by":"publisher","first-page":"783","DOI":"10.1016\/s0306-4522(03)00326-9","volume":"120","author":"L Zhou","year":"2003","unstructured":"Zhou L, Furuta T, Kaneko T. Chemical organization of projection neurons in the rat accumbens nucleus and olfactory tubercle. Neuroscience. 2003;120:783\u201398. https:\/\/doi.org\/10.1016\/s0306-4522(03)00326-9.","journal-title":"Neuroscience"},{"key":"2460_CR97","doi-asserted-by":"publisher","first-page":"1230","DOI":"10.1038\/nn.4068","volume":"18","author":"YM Kupchik","year":"2015","unstructured":"Kupchik YM, Brown RM, Heinsbroek JA, Lobo MK, Schwartz DJ, Kalivas PW. Coding the direct\/indirect pathways by D1 and D2 receptors is not valid for accumbens projections. Nat Neurosci. 2015;18:1230\u20132. https:\/\/doi.org\/10.1038\/nn.4068.","journal-title":"Nat Neurosci"},{"key":"2460_CR98","doi-asserted-by":"publisher","first-page":"1593","DOI":"10.1126\/science.275.5306.1593","volume":"275","author":"W Schultz","year":"1997","unstructured":"Schultz W, Dayan P, Montague PR. A neural substrate of prediction and reward. Science. 1997;275:1593\u20139. https:\/\/doi.org\/10.1126\/science.275.5306.1593.","journal-title":"Science"},{"key":"2460_CR99","doi-asserted-by":"publisher","first-page":"304","DOI":"10.1038\/1124","volume":"1","author":"JR Hollerman","year":"1998","unstructured":"Hollerman JR, Schultz W. Dopamine neurons report an error in the temporal prediction of reward during learning. Nat Neurosci. 1998;1:304\u20139. https:\/\/doi.org\/10.1038\/1124.","journal-title":"Nat Neurosci"},{"key":"2460_CR100","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1016\/j.neuron.2005.05.020","volume":"47","author":"HM Bayer","year":"2005","unstructured":"Bayer HM, Glimcher PW. Midbrain dopamine neurons encode a quantitative reward prediction error signal. Neuron. 2005;47:129\u201341. https:\/\/doi.org\/10.1016\/j.neuron.2005.05.020.","journal-title":"Neuron"},{"key":"2460_CR101","doi-asserted-by":"publisher","first-page":"719","DOI":"10.1016\/0301-0082(94)90025-6","volume":"42","author":"C Pennartz","year":"1994","unstructured":"Pennartz C, Groenewegen H. Lopes da Silva FH. The nucleus accumbens as a complex of functionally distinct neuronal ensembles: an integration of behavioural, electrophysiological and anatomical data. Prog Neurobiol. 1994;42:719\u201361. https:\/\/doi.org\/10.1016\/0301-0082(94)90025-6.","journal-title":"Prog Neurobiol"},{"key":"2460_CR102","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1111\/j.1749-6632.1999.tb09260.x","volume":"877","author":"HJ Groenewegen","year":"1999","unstructured":"Groenewegen HJ, Wright CI, Beijer AVJ, Voorn P. Convergence and segregation of ventral striatal inputs and outputs. Ann N Y Acad Sci. 1999;877:49\u201363. https:\/\/doi.org\/10.1111\/j.1749-6632.1999.tb09260.x.","journal-title":"Ann N Y Acad Sci"},{"key":"2460_CR103","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1111\/j.1749-6632.1999.tb09267.x","volume":"877","author":"P O'DONNELL","year":"1999","unstructured":"O'DONNELL P, Greene J, Pabello N, Lewis BL, Grace AA. Modulation of cell firing in the nucleus accumbens. Ann N Y Acad Sci. 1999;877:157\u201375. https:\/\/doi.org\/10.1111\/j.1749-6632.1999.tb09267.x.","journal-title":"Ann N Y Acad Sci"},{"key":"2460_CR104","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1038\/s41422-021-00588-5","volume":"32","author":"Z Liu","year":"2022","unstructured":"Liu Z, Le Q, Lv Y, Chen X, Cui J, Zhou Y, et al. A distinct D1-MSN subpopulation down-regulates dopamine to promote negative emotional state. Cell Res. 2022;32:139\u201356. https:\/\/doi.org\/10.1038\/s41422-021-00588-5.","journal-title":"Cell Res"},{"key":"2460_CR105","doi-asserted-by":"publisher","first-page":"1126","DOI":"10.1016\/j.celrep.2016.06.059","volume":"16","author":"O Gokce","year":"2016","unstructured":"Gokce O, Stanley GM, Treutlein B, Neff NF, Camp JG, Malenka RC, et al. Cellular Taxonomy of the Mouse Striatum as Revealed by Single-Cell RNA-Seq. Cell Rep. 2016;16:1126\u201337. https:\/\/doi.org\/10.1016\/j.celrep.2016.06.059.","journal-title":"Cell Rep"},{"key":"2460_CR106","doi-asserted-by":"publisher","first-page":"688","DOI":"10.1016\/j.neuron.2019.11.004","volume":"105","author":"G Stanley","year":"2020","unstructured":"Stanley G, Gokce O, Malenka RC, S\u00fcdhof TC, Quake SR. Continuous and Discrete Neuron Types of the Adult Murine Striatum. Neuron. 2020;105:688\u2013699.e8. https:\/\/doi.org\/10.1016\/j.neuron.2019.11.004.","journal-title":"Neuron"},{"key":"2460_CR107","doi-asserted-by":"publisher","first-page":"1757","DOI":"10.1038\/s41593-021-00938-x","volume":"24","author":"R Chen","year":"2021","unstructured":"Chen R, Blosser TR, Djekidel MN, Hao J, Bhattacherjee A, Chen W, et al. Decoding molecular and cellular heterogeneity of mouse nucleus accumbens. Nat Neurosci. 2021;24:1757\u201371. https:\/\/doi.org\/10.1038\/s41593-021-00938-x.","journal-title":"Nat Neurosci"},{"key":"2460_CR108","doi-asserted-by":"publisher","first-page":"5473","DOI":"10.1016\/j.cub.2021.10.015","volume":"31","author":"J He","year":"2021","unstructured":"He J, Kleyman M, Chen J, Alikaya A, Rothenhoefer KM, Ozturk BE, et al. Transcriptional and anatomical diversity of medium spiny neurons in the primate striatum. Curr Biol. 2021;31:5473\u201386.e6. https:\/\/doi.org\/10.1016\/j.cub.2021.10.015.","journal-title":"Curr Biol"},{"key":"2460_CR109","doi-asserted-by":"publisher","first-page":"7308","DOI":"10.1523\/JNEUROSCI.22-16-07308.2002","volume":"22","author":"SM Reynolds","year":"2002","unstructured":"Reynolds SM, Berridge KC. Positive and negative motivation in nucleus accumbens shell: bivalent rostrocaudal gradients for GABA-elicited eating, taste \u201cliking\u201d\/\u201cdisliking\u201d reactions, place preference\/avoidance, and fear. J Neurosci. 2002;22:7308\u201320.","journal-title":"J Neurosci"},{"key":"2460_CR110","doi-asserted-by":"publisher","first-page":"3261","DOI":"10.1523\/JNEUROSCI.21-09-03261.2001","volume":"21","author":"SM Reynolds","year":"2001","unstructured":"Reynolds SM, Berridge KC. Fear and feeding in the nucleus accumbens shell: rostrocaudal segregation of GABA-elicited defensive behavior versus eating behavior. J Neurosci. 2001;21:3261\u201370. https:\/\/doi.org\/10.1523\/JNEUROSCI.21-09-03261.2001.","journal-title":"J Neurosci"},{"key":"2460_CR111","doi-asserted-by":"publisher","first-page":"2187","DOI":"10.1046\/j.1460-9568.2003.02642.x","volume":"17","author":"SM Reynolds","year":"2003","unstructured":"Reynolds SM, Berridge KC. Glutamate motivational ensembles in nucleus accumbens: rostrocaudal shell gradients of fear and feeding. Eur J Neurosci. 2003;17:2187\u2013200. https:\/\/doi.org\/10.1046\/j.1460-9568.2003.02642.x.","journal-title":"Eur J Neurosci"},{"key":"2460_CR112","doi-asserted-by":"publisher","first-page":"1260","DOI":"10.1016\/j.neuron.2018.07.051","volume":"99","author":"SJ Reed","year":"2018","unstructured":"Reed SJ, Lafferty CK, Mendoza JA, Yang AK, Davidson TJ, Grosenick L, et al. Coordinated Reductions in Excitatory Input to the Nucleus Accumbens Underlie Food Consumption. Neuron. 2018;99:1260\u201373.e4. https:\/\/doi.org\/10.1016\/j.neuron.2018.07.051.","journal-title":"Neuron"},{"key":"2460_CR113","doi-asserted-by":"publisher","first-page":"112069","DOI":"10.1016\/j.celrep.2023.112069","volume":"42","author":"G Chen","year":"2023","unstructured":"Chen G, Lai S, Bao G, Ke J, Meng X, Lu S, et al. Distinct reward processing by subregions of the nucleus accumbens. Cell Rep. 2023;42:112069 https:\/\/doi.org\/10.1016\/j.celrep.2023.112069.","journal-title":"Cell Rep"},{"key":"2460_CR114","doi-asserted-by":"publisher","first-page":"642","DOI":"10.1038\/nature17400","volume":"531","author":"KA Zalocusky","year":"2016","unstructured":"Zalocusky KA, Ramakrishnan C, Lerner TN, Davidson TJ, Knutson B, Deisseroth K. Nucleus accumbens D2R cells signal prior outcomes and control risky decision-making. Nature. 2016;531:642\u20136. https:\/\/doi.org\/10.1038\/nature17400.","journal-title":"Nature"},{"key":"2460_CR115","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-38025-3","volume":"14","author":"T Nishioka","year":"2023","unstructured":"Nishioka T, Attachaipanich S, Hamaguchi K, Lazarus M, de Kerchove d'Exaerde A, Macpherson T, et al. Error-related signaling in nucleus accumbens D2 receptor-expressing neurons guides inhibition-based choice behavior in mice. Nat Commun. 2023;14:2284. https:\/\/doi.org\/10.1038\/s41467-023-38025-3.","journal-title":"Nat Commun"},{"key":"2460_CR116","doi-asserted-by":"publisher","first-page":"16362","DOI":"10.1523\/JNEUROSCI.1392-15.2015","volume":"35","author":"GE Hodes","year":"2015","unstructured":"Hodes GE, Pfau ML, Purushothaman I, Ahn HF, Golden SA, Christoffel DJ, et al. Sex Differences in Nucleus Accumbens Transcriptome Profiles Associated with Susceptibility versus Resilience to Subchronic Variable Stress. J Neurosci. 2015;35:16362\u201376. https:\/\/doi.org\/10.1523\/JNEUROSCI.1392-15.2015.","journal-title":"J Neurosci"},{"key":"2460_CR117","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1016\/j.neuroscience.2016.08.032","volume":"335","author":"AM Henricks","year":"2016","unstructured":"Henricks AM, Berger AL, Lugo JM, Baxter-Potter LN, Bieniasz KV, Craft RM, et al. Sex differences in alcohol consumption and alterations in nucleus accumbens endocannabinoid mRNA in alcohol-dependent rats. Neuroscience. 2016;335:195\u2013206. https:\/\/doi.org\/10.1016\/j.neuroscience.2016.08.032.","journal-title":"Neuroscience"},{"key":"2460_CR118","doi-asserted-by":"publisher","DOI":"10.1101\/2023.06.28.546959","author":"AD Chapp","year":"2023","unstructured":"Chapp AD, Nwakama CA, Mermelstein PG, Thomas MJ. Fundamental Sex Differences in Cocaine-Induced Plasticity of D1- and D2-MSNs in the Mouse Nucleus Accumbens Shell. bioRxiv. 2023. https:\/\/doi.org\/10.1101\/2023.06.28.546959.","journal-title":"bioRxiv"}],"container-title":["Neuropsychopharmacology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41386-026-02460-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41386-026-02460-9","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41386-026-02460-9.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,11]],"date-time":"2026-06-11T09:57:11Z","timestamp":1781171831000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41386-026-02460-9"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,11]]},"references-count":118,"alternative-id":["2460"],"URL":"https:\/\/doi.org\/10.1038\/s41386-026-02460-9","relation":{},"ISSN":["0893-133X","1740-634X"],"issn-type":[{"value":"0893-133X","type":"print"},{"value":"1740-634X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,11]]},"assertion":[{"value":"25 November 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"27 May 2026","order":2,"name":"revised","label":"Revised","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 May 2026","order":3,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"11 June 2026","order":4,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}