{"id":"https://openalex.org/W7223146162","doi":"https://doi.org/10.64898/2026.10.15.759424","title":"Opposing reward-epoch population coding in anterior insular cortex dissociates sign- and goal-tracking despite equivalent conditioned responding","display_name":"Opposing reward-epoch population coding in anterior insular cortex dissociates sign- and goal-tracking despite equivalent conditioned responding","publication_year":2026,"publication_date":"2026-10-21","ids":{"openalex":"https://openalex.org/W7223146162","doi":"https://doi.org/10.64898/2026.10.15.759424"},"language":"en","primary_location":{"id":"doi:10.64898/2026.10.15.759424","doi":"https://doi.org/10.64898/2026.10.15.759424","is_oa":true,"landing_page_url":"https://doi.org/10.64898/2026.10.15.759424","pdf_url":"https://www.biorxiv.org/content/biorxiv/early/2026/10/21/2026.10.15.759424.full.pdf","source":{"id":"https://openalex.org/S4306402567","display_name":"bioRxiv (Cold Spring Harbor Laboratory)","issn_l":null,"issn":null,"is_oa":false,"is_in_doaj":false,"is_core":false,"listed_in":[],"host_organization":"https://openalex.org/I2750212522","host_organization_name":"Cold Spring Harbor Laboratory","host_organization_lineage":["https://openalex.org/I2750212522"],"host_organization_lineage_names":[],"type":"repository"},"license":"cc-by-nc-nd","license_id":"https://openalex.org/licenses/cc-by-nc-nd","version":"acceptedVersion","is_accepted":true,"is_published":false,"raw_source_name":"bioRxiv","raw_type":"posted-content"},"volume":null,"type":"preprint","is_traditional":true,"indexed_in":["crossref"],"open_access":{"is_oa":true,"oa_status":"green","oa_url":"https://www.biorxiv.org/content/biorxiv/early/2026/10/21/2026.10.15.759424.full.pdf","any_repository_has_fulltext":true},"authorships":[{"author_position":"first","author":{"id":"https://openalex.org/A5014977834","display_name":"Sara E. Keefer","orcid":"https://orcid.org/0000-0002-4905-9973","observed_orcids":["https://orcid.org/0000-0002-4905-9973"]},"institutions":[{"id":"https://openalex.org/I126744593","display_name":"University of Maryland, Baltimore","ror":"https://ror.org/04rq5mt64","country_code":"US","type":"education","lineage":["https://openalex.org/I126744593"]},{"id":"https://openalex.org/I176113532","display_name":"Gettysburg College","ror":"https://ror.org/045e26x92","country_code":"US","type":"education","lineage":["https://openalex.org/I176113532"]}],"countries":["US"],"is_corresponding":false,"raw_author_name":"Sara E. Keefer","raw_affiliation_strings":["Department of Neurobiology, University of Maryland School of Medicine, Baltimore, MD 21201","Gettysburg College, Department of Psychology, Gettysburg, PA 17325"],"raw_orcid":"https://orcid.org/0000-0002-4905-9973","affiliations":[{"raw_affiliation_string":"Department of Neurobiology, University of Maryland School of Medicine, Baltimore, MD 21201","institution_ids":["https://openalex.org/I126744593"]},{"raw_affiliation_string":"Gettysburg College, Department of Psychology, Gettysburg, PA 17325","institution_ids":["https://openalex.org/I176113532"]}]},{"author_position":"middle","author":{"id":"https://openalex.org/A5017789225","display_name":"David Alexander Martin","orcid":"https://orcid.org/0000-0001-5783-7141","observed_orcids":["https://orcid.org/0000-0001-5783-7141"]},"institutions":[{"id":"https://openalex.org/I126744593","display_name":"University of Maryland, Baltimore","ror":"https://ror.org/04rq5mt64","country_code":"US","type":"education","lineage":["https://openalex.org/I126744593"]}],"countries":["US"],"is_corresponding":false,"raw_author_name":"David A. Martin","raw_affiliation_strings":["Department of Neurobiology, University of Maryland School of Medicine, Baltimore, MD 21201"],"raw_orcid":null,"affiliations":[{"raw_affiliation_string":"Department of Neurobiology, University of Maryland School of Medicine, Baltimore, MD 21201","institution_ids":["https://openalex.org/I126744593"]}]},{"author_position":"last","author":{"id":"https://openalex.org/A5032241182","display_name":"Donna J. Calu","orcid":"https://orcid.org/0000-0003-2377-9494","observed_orcids":["https://orcid.org/0000-0003-2377-9494","https://orcid.org/0000-0002-6410-3100"]},"institutions":[{"id":"https://openalex.org/I126744593","display_name":"University of Maryland, Baltimore","ror":"https://ror.org/04rq5mt64","country_code":"US","type":"education","lineage":["https://openalex.org/I126744593"]}],"countries":["US"],"is_corresponding":true,"raw_author_name":"Donna J. Calu","raw_affiliation_strings":["Department of Neurobiology, University of Maryland School of Medicine, Baltimore, MD 21201"],"raw_orcid":"https://orcid.org/0000-0003-2377-9494","affiliations":[{"raw_affiliation_string":"Department of Neurobiology, University of Maryland School of Medicine, Baltimore, MD 21201","institution_ids":["https://openalex.org/I126744593"]}]}],"institutions":[],"countries_distinct_count":1,"institutions_distinct_count":2,"corresponding_author_ids":["https://openalex.org/A5032241182"],"corresponding_institution_ids":["https://openalex.org/I126744593"],"apc_list":null,"apc_paid":null,"fwci":null,"has_fulltext":true,"cited_by_count":0,"citation_normalized_percentile":null,"cited_by_percentile_year":null,"biblio":{"volume":null,"issue":null,"first_page":null,"last_page":null},"is_retracted":false,"is_paratext":false,"is_xpac":false,"primary_topic":{"id":"https://openalex.org/T10448","display_name":"Memory and Neural Mechanisms","score":0.6977999806404114,"subfield":{"id":"https://openalex.org/subfields/2805","display_name":"Cognitive Neuroscience"},"field":{"id":"https://openalex.org/fields/28","display_name":"Neuroscience"},"domain":{"id":"https://openalex.org/domains/1","display_name":"Life Sciences"}},"topics":[{"id":"https://openalex.org/T10448","display_name":"Memory and Neural Mechanisms","score":0.6977999806404114,"subfield":{"id":"https://openalex.org/subfields/2805","display_name":"Cognitive Neuroscience"},"field":{"id":"https://openalex.org/fields/28","display_name":"Neuroscience"},"domain":{"id":"https://openalex.org/domains/1","display_name":"Life Sciences"}},{"id":"https://openalex.org/T10056","display_name":"Neurotransmitter Receptor Influence on Behavior","score":0.20260000228881836,"subfield":{"id":"https://openalex.org/subfields/2804","display_name":"Cellular and Molecular Neuroscience"},"field":{"id":"https://openalex.org/fields/28","display_name":"Neuroscience"},"domain":{"id":"https://openalex.org/domains/1","display_name":"Life Sciences"}},{"id":"https://openalex.org/T10042","display_name":"Neural and Behavioral Psychology Studies","score":0.0681999996304512,"subfield":{"id":"https://openalex.org/subfields/2805","display_name":"Cognitive Neuroscience"},"field":{"id":"https://openalex.org/fields/28","display_name":"Neuroscience"},"domain":{"id":"https://openalex.org/domains/1","display_name":"Life Sciences"}}],"keywords":[{"id":"https://openalex.org/keywords/anterior-insula","display_name":"anterior insula","score":0.9559999704360962},{"id":"https://openalex.org/keywords/sign-tracking","display_name":"sign-tracking","score":0.9710000157356262},{"id":"https://openalex.org/keywords/goal-monitoring","display_name":"goal tracking","score":0.9990000128746033},{"id":"https://openalex.org/keywords/classical-conditioning","display_name":"classical conditioning","score":0.8040000200271606},{"id":"https://openalex.org/keywords/reward-prediction","display_name":"reward prediction","score":0.593999981880188},{"id":"https://openalex.org/keywords/population-coding","display_name":"population coding","score":0.8159999847412109},{"id":"https://openalex.org/keywords/single-unit-activity","display_name":"single-unit activity","score":0.6690000295639038},{"id":"https://openalex.org/keywords/basolateral-amygdala","display_name":"basolateral amygdala","score":0.7960000038146973},{"id":"https://openalex.org/keywords/nucleus-accumbens","display_name":"nucleus accumbens","score":0.9980000257492065},{"id":"https://openalex.org/keywords/approach-latency","display_name":"approach latency","score":0.5529999732971191},{"id":"https://openalex.org/keywords/motivational-strategies","display_name":"motivational strategies","score":0.5410000085830688}],"concepts":[{"id":"https://openalex.org/C169760540","wikidata":"https://www.wikidata.org/wiki/Q207011","display_name":"Neuroscience","level":1,"score":0.7620000243186951},{"id":"https://openalex.org/C2780539906","wikidata":"https://www.wikidata.org/wiki/Q1422654","display_name":"Insular cortex","level":2,"score":0.683899998664856},{"id":"https://openalex.org/C2908647359","wikidata":"https://www.wikidata.org/wiki/Q2625603","display_name":"Population","level":2,"score":0.6028000116348267},{"id":"https://openalex.org/C2780451725","wikidata":"https://www.wikidata.org/wiki/Q4867818","display_name":"Basolateral amygdala","level":3,"score":0.5655999779701233},{"id":"https://openalex.org/C2779144063","wikidata":"https://www.wikidata.org/wiki/Q338924","display_name":"Amygdala","level":2,"score":0.545799970626831},{"id":"https://openalex.org/C15744967","wikidata":"https://www.wikidata.org/wiki/Q9418","display_name":"Psychology","level":0,"score":0.5388000011444092},{"id":"https://openalex.org/C77637269","wikidata":"https://www.wikidata.org/wiki/Q7002051","display_name":"Neural coding","level":2,"score":0.48030000925064087},{"id":"https://openalex.org/C2776552330","wikidata":"https://www.wikidata.org/wiki/Q1476178","display_name":"Nucleus accumbens","level":3,"score":0.4334999918937683},{"id":"https://openalex.org/C86803240","wikidata":"https://www.wikidata.org/wiki/Q420","display_name":"Biology","level":0,"score":0.4000000059604645},{"id":"https://openalex.org/C2779918689","wikidata":"https://www.wikidata.org/wiki/Q3771842","display_name":"Stimulus (psychology)","level":2,"score":0.3343999981880188},{"id":"https://openalex.org/C2778402161","wikidata":"https://www.wikidata.org/wiki/Q2121931","display_name":"Anterior cingulate cortex","level":3,"score":0.3203999996185303},{"id":"https://openalex.org/C2778950215","wikidata":"https://www.wikidata.org/wiki/Q177601","display_name":"Macaque","level":2,"score":0.2874000072479248},{"id":"https://openalex.org/C159423971","wikidata":"https://www.wikidata.org/wiki/Q177251","display_name":"Associative property","level":2,"score":0.2854999899864197},{"id":"https://openalex.org/C2910823151","wikidata":"https://www.wikidata.org/wiki/Q936818","display_name":"Parabrachial Nucleus","level":3,"score":0.28110000491142273},{"id":"https://openalex.org/C2778187257","wikidata":"https://www.wikidata.org/wiki/Q464210","display_name":"Basal ganglia","level":3,"score":0.27379998564720154},{"id":"https://openalex.org/C2779246727","wikidata":"https://www.wikidata.org/wiki/Q184215","display_name":"Thalamus","level":2,"score":0.27160000801086426},{"id":"https://openalex.org/C2781195155","wikidata":"https://www.wikidata.org/wiki/Q18680","display_name":"Prefrontal cortex","level":3,"score":0.26669999957084656},{"id":"https://openalex.org/C2983526489","wikidata":"https://www.wikidata.org/wiki/Q133500","display_name":"Associative learning","level":2,"score":0.2639999985694885},{"id":"https://openalex.org/C112592302","wikidata":"https://www.wikidata.org/wiki/Q1207387","display_name":"Excitatory postsynaptic potential","level":3,"score":0.25380000472068787}],"mesh":[],"locations_count":2,"locations":[{"id":"doi:10.64898/2026.10.15.759424","doi":"https://doi.org/10.64898/2026.10.15.759424","is_oa":true,"landing_page_url":"https://doi.org/10.64898/2026.10.15.759424","pdf_url":"https://www.biorxiv.org/content/biorxiv/early/2026/10/21/2026.10.15.759424.full.pdf","source":{"id":"https://openalex.org/S4306402567","display_name":"bioRxiv (Cold Spring Harbor Laboratory)","issn_l":null,"issn":null,"is_oa":false,"is_in_doaj":false,"is_core":false,"listed_in":[],"host_organization":"https://openalex.org/I2750212522","host_organization_name":"Cold Spring Harbor Laboratory","host_organization_lineage":["https://openalex.org/I2750212522"],"host_organization_lineage_names":[],"type":"repository"},"license":"cc-by-nc-nd","license_id":"https://openalex.org/licenses/cc-by-nc-nd","version":"acceptedVersion","is_accepted":true,"is_published":false,"raw_source_name":"bioRxiv","raw_type":"posted-content"},{"id":"pmh:oai:pubmedcentral.nih.gov:13738534","doi":null,"is_oa":true,"landing_page_url":"https://pmc.ncbi.nlm.nih.gov/articles/PMC13738534/","pdf_url":null,"source":{"id":"https://openalex.org/S2764455111","display_name":"PubMed Central","issn_l":null,"issn":null,"is_oa":true,"is_in_doaj":false,"is_core":false,"listed_in":[],"host_organization":"https://openalex.org/I1299303238","host_organization_name":"National Institutes of Health","host_organization_lineage":["https://openalex.org/I1299303238"],"host_organization_lineage_names":[],"type":"repository"},"license":"cc-by-nc-nd","license_id":"https://openalex.org/licenses/cc-by-nc-nd","version":"publishedVersion","is_accepted":true,"is_published":true,"raw_source_name":"bioRxiv","raw_type":"Text"}],"best_oa_location":{"id":"doi:10.64898/2026.10.15.759424","doi":"https://doi.org/10.64898/2026.10.15.759424","is_oa":true,"landing_page_url":"https://doi.org/10.64898/2026.10.15.759424","pdf_url":"https://www.biorxiv.org/content/biorxiv/early/2026/10/21/2026.10.15.759424.full.pdf","source":{"id":"https://openalex.org/S4306402567","display_name":"bioRxiv (Cold Spring Harbor Laboratory)","issn_l":null,"issn":null,"is_oa":false,"is_in_doaj":false,"is_core":false,"listed_in":[],"host_organization":"https://openalex.org/I2750212522","host_organization_name":"Cold Spring Harbor Laboratory","host_organization_lineage":["https://openalex.org/I2750212522"],"host_organization_lineage_names":[],"type":"repository"},"license":"cc-by-nc-nd","license_id":"https://openalex.org/licenses/cc-by-nc-nd","version":"acceptedVersion","is_accepted":true,"is_published":false,"raw_source_name":"bioRxiv","raw_type":"posted-content"},"sustainable_development_goals":[],"study_designs":[],"awards":[{"id":"https://openalex.org/G4640393866","display_name":"Role of basolateral amygdala projections in mediating individual differences in motivation and flexibility","funder_award_id":"R01DA043533","funder_id":"https://openalex.org/F4320337347","funder_display_name":"National Institute on Drug Abuse"},{"id":"https://openalex.org/G8496983265","display_name":"Individual differences in cortical-striatal pathway utilization regulating flexibility","funder_award_id":"F32DA053772","funder_id":"https://openalex.org/F4320337347","funder_display_name":"National Institute on Drug Abuse"}],"funders":[{"id":"https://openalex.org/F4320308324","display_name":"McKnight Foundation","ror":"https://ror.org/003ghvj67"},{"id":"https://openalex.org/F4320337347","display_name":"National Institute on Drug Abuse","ror":"https://ror.org/00fq5cm18"}],"has_content":{"grobid_xml":true,"pdf":true},"content_urls":{"pdf":"https://content.openalex.org/works/W7223146162.pdf","grobid_xml":"https://content.openalex.org/works/W7223146162.grobid-xml"},"referenced_works_count":0,"referenced_works":[],"related_works":["https://openalex.org/W4386241944","https://openalex.org/W2919358373","https://openalex.org/W2802364329","https://openalex.org/W2085311250","https://openalex.org/W2493630655","https://openalex.org/W1927661140","https://openalex.org/W2975712666","https://openalex.org/W1978211553","https://openalex.org/W4406949758","https://openalex.org/W1964019124"],"abstract_inverted_index":{"Abstract":[0],"When":[1],"Pavlovian":[2],"lever":[3],"cues":[4],"predict":[5],"reward,":[6],"rats":[7,17,29,90,319],"diverge":[8],"in":[9,78,86,198,222,227,252,276,317,335,360],"how":[10,253,321],"they":[11,254,323],"express":[12],"learned":[13],"associations:":[14],"sign-tracking":[15,318],"(ST)":[16],"approach":[18,30,99,185,260],"and":[19,34,44,49,88,147,211,288,365],"engage":[20],"the":[21,31,67,102,129,137,143,177,228,261,268,282,326,329,344],"cue":[22,141,262,327],"with":[23,47,97,187,191],"greater":[24,121],"persistence,":[25],"while":[26,116,169,264],"goal-tracking":[27,336],"(GT)":[28],"reward":[32,138,178,269,299,330,368],"site":[33,239,356],"are":[35,52,225,363],"more":[36],"consistently":[37],"outcome-sensitive.":[38],"The":[39,200],"anterior":[40,295],"insular":[41,296,341,352],"cortex":[42,297,353],"(aIC)":[43],"its":[45],"interactions":[46],"amygdala":[48,210],"striatal":[50],"regions":[51],"differentially":[53],"recruited":[54],"across":[55,304],"phenotypes,":[56,154],"implicating":[57],"aIC":[58,84,106,118,205,232,236],"as":[59,237,354],"a":[60,120,133,195,238,332,355],"node":[61],"where":[62,240,357],"associative":[63,223,277],"strategy":[64,224,242,362],"diverges":[65],"at":[66,281],"circuit":[68,290],"level.":[69,284],"To":[70],"characterize":[71],"this":[72],"neural":[73,283],"divergence":[74],"independent":[75],"of":[76,123,145,202,231],"differences":[77,221,275,359],"conditioned":[79],"responding,":[80],"we":[81,292],"recorded":[82],"single-unit":[83,286],"activity":[85,316],"ST":[87,117,159,180],"GT":[89,105,170],"performing":[91],"an":[92],"odor-guided":[93],"sucrose":[94,114],"discrimination":[95,131],"task":[96],"equivalent":[98],"behavior.":[100],"At":[101],"population":[103,233],"level,":[104],"was":[107],"dominated":[108],"by":[109],"task-excited":[110],"neurons":[111,125,149,161,172,207],"that":[112,219,294],"drove":[113],"discrimination,":[115,168],"carried":[119,128],"proportion":[122],"task-inhibited":[124],"whose":[126],"suppression":[127],"dominant":[130],"signal,":[132],"dissociation":[134],"specific":[135],"to":[136,256,328,367],"epoch.":[139,179],"During":[140],"sampling,":[142],"balance":[144],"excited":[146],"inhibited":[148],"did":[150],"not":[151],"differ":[152],"between":[153],"but":[155],"temporal":[156],"structure":[157,230],"diverged:":[158],"cue-excited":[160,171],"showed":[162,346],"transient":[163],"CS-onset":[164],"responses":[165,189],"without":[166],"sustained":[167,174],"maintained":[173],"CS+/CS-discrimination":[175],"into":[176,245],"cue-period":[181,315],"firing":[182],"predicted":[183,320],"post-cue":[184],"latency,":[186],"stronger":[188],"associated":[190],"slower":[192],"goal":[193],"approach,":[194],"relationship":[196,333],"absent":[197,334],"GT.":[199],"majority":[201,345],"optogenetically":[203,339],"identified":[204,340],"projection":[206,342],"targeting":[208],"basolateral":[209],"nucleus":[212],"accumbens":[213],"were":[214],"cue-inhibited.":[215],"These":[216,271,349],"findings":[217,350],"reveal":[218],"individual":[220,274,358],"encoded":[226,364],"organizational":[229,302],"activity,":[234],"positioning":[235],"motivational":[241,361],"is":[243,312],"translated":[244],"distinct":[246],"outputs.":[247],"Significance":[248],"statement":[249],"Rats":[250],"vary":[251],"respond":[255],"reward-predictive":[257],"cues:":[258],"some":[259],"itself,":[263],"others":[265],"orient":[266],"toward":[267],"location.":[270],"strategies":[272],"reflect":[273],"learning":[278],"poorly":[279],"understood":[280],"Using":[285],"recording":[287],"optogenetic":[289],"identification,":[291],"show":[293],"encodes":[298],"through":[300],"opposing":[301],"principles":[303],"these":[305],"two":[306],"groups,":[307],"even":[308],"when":[309],"observable":[310],"behavior":[311],"identical.":[313],"Session-level":[314],"slowly":[322],"transitioned":[324],"from":[325],"location,":[331],"rats.":[337],"Among":[338],"neurons,":[343],"cue-suppressed":[347],"activity.":[348],"identify":[351],"transmitted":[366],"circuits.":[369]},"counts_by_year":[],"updated_date":"2026-11-08T12:12:28.789561","created_date":"2026-10-22T00:00:00"}