{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T04:48:28Z","timestamp":1792039708520,"version":"4.4.0"},"reference-count":77,"publisher":"Wiley","issue":"6","license":[{"start":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T00:00:00Z","timestamp":1791936000000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T00:00:00Z","timestamp":1791936000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2026,10,14]],"date-time":"2026-10-14T00:00:00Z","timestamp":1791936000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100000049","name":"National Institute on Aging","doi-asserted-by":"publisher","award":["K01AG066847"],"award-info":[{"award-number":["K01AG066847"]}],"id":[{"id":"10.13039\/100000049","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000049","name":"National Institute on Aging","doi-asserted-by":"publisher","award":["R01AG092662"],"award-info":[{"award-number":["R01AG092662"]}],"id":[{"id":"10.13039\/100000049","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000049","name":"National Institute on Aging","doi-asserted-by":"publisher","award":["P01AG052350"],"award-info":[{"award-number":["P01AG052350"]}],"id":[{"id":"10.13039\/100000049","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000001","name":"National Science Foundation","doi-asserted-by":"publisher","award":["2121164"],"award-info":[{"award-number":["2121164"]}],"id":[{"id":"10.13039\/100000001","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Hippocampus"],"published-print":{"date-parts":[[2026,11]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>The subiculum (SUB) is the main output structure of the hippocampus, influencing diverse behaviors through its widespread cortical and subcortical connections. Our previous work creating the mouse Hippocampus Gene Expression Atlas (HGEA) identified four genetically distinct cellular layers across five columnar domains in the SUB, with gene expression boundaries corresponding to distinct connectivity patterns and brain\u2010wide networks involved in spatial navigation, social behavior, and neuroendocrine regulation. Using the Digital Brain Mouse Projectome Atlas (MPA) tool, we conducted \u2018virtual tract\u2010tracing\u2019 to assess whether connectivity patterns of single\u2010neuron 3D reconstructions aligned with HGEA\u2010defined SUB cell types. We classified 689 SUB projection neurons into 12 HGEA cell\u2010type groups based on their laminar and columnar distributions, whose spatial organization recapitulated HGEA\u2010defined 3D boundaries. Using this population sample, we performed a SUB cell\u2010type census, characterized neuronal heterogeneity and projection prevalence, identified common and rare connectivity motifs and axonal collateralization patterns, and defined distinct projection themes for each SUB cell type. Together, this analysis integrates single\u2010neuron and population\u2010level data to advance understanding of SUB cell type organization and its contributions to brain\u2010wide networks regulating diverse behaviors.<\/jats:p>","DOI":"10.1002\/hipo.70138","type":"journal-article","created":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T04:29:39Z","timestamp":1792038579000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Mapping Projectome Heterogeneity of Subiculum Neuron Cell Types"],"prefix":"10.1002","volume":"36","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-0271-8946","authenticated-orcid":false,"given":"Ann W.","family":"Saustad","sequence":"first","affiliation":[{"name":"USC Mark and Mary Stevens Neuroimaging and Informatics Institute, University of Southern California  Los Angeles California USA"},{"name":"USC Center for Integrative Connectomics, University of Southern California  Los Angeles California USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8231-8893","authenticated-orcid":false,"given":"Michael S.","family":"Bienkowski","sequence":"additional","affiliation":[{"name":"USC Mark and Mary Stevens Neuroimaging and Informatics Institute, University of Southern California  Los Angeles California USA"},{"name":"USC Center for Integrative Connectomics, University of Southern California  Los Angeles California USA"},{"name":"USC Zilkha Neurogenetic Institute, University of Southern California  Los Angeles California USA"},{"name":"USC Alfred E. Mann Department of Biomedical Engineering, USC Viterbi School of Engineering  Los Angeles California USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,10,14]]},"reference":[{"key":"e_1_2_10_2_1","first-page":"65","volume-title":"Progress in Brain Research","author":"Aggleton J. P.","year":"2015"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41597\u2010022\u201001426\u2010y"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.1016\/0891\u20100618(93)90005\u2010o"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.1016\/0166\u20102236(94)90047\u20107"},{"key":"e_1_2_10_6_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2012.10.038"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1212\/WNL.0b013e31823fcdee"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41593\u2010018\u20100241\u2010y"},{"key":"e_1_2_10_9_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598\u2010021\u201081362\u2010w"},{"key":"e_1_2_10_10_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.22407"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2602"},{"key":"e_1_2_10_12_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.20504"},{"key":"e_1_2_10_13_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.pneurobio.2026.102893"},{"key":"e_1_2_10_14_1","doi-asserted-by":"publisher","DOI":"10.1002\/cne.903240204"},{"key":"e_1_2_10_15_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2018.03.031"},{"key":"e_1_2_10_16_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.37701"},{"key":"e_1_2_10_17_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2021.772016"},{"key":"e_1_2_10_18_1","doi-asserted-by":"publisher","DOI":"10.1016\/0301\u20100082(92)90015\u20107"},{"key":"e_1_2_10_19_1","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2019.00060"},{"key":"e_1_2_10_20_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2020.107648"},{"key":"e_1_2_10_21_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.10566"},{"key":"e_1_2_10_22_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010018\u20100244\u20106"},{"key":"e_1_2_10_23_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2009.11.031"},{"key":"e_1_2_10_24_1","doi-asserted-by":"crossref","first-page":"224","DOI":"10.1007\/978-1-4899-6704-6_8","volume-title":"Neurobiology of Cingulate Cortex and Limbic Thalamus: A Comprehensive Handbook","author":"Finch D. M.","year":"1993"},{"key":"e_1_2_10_25_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.adf6162"},{"key":"e_1_2_10_26_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.70087"},{"key":"e_1_2_10_27_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00429\u2010019\u201001841\u20109"},{"key":"e_1_2_10_28_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2013.02.005"},{"key":"e_1_2_10_29_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature03721"},{"key":"e_1_2_10_30_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010019\u20101716\u2010z"},{"key":"e_1_2_10_31_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnana.2020.605021"},{"key":"e_1_2_10_32_1","doi-asserted-by":"publisher","DOI":"10.1002\/cne.1195"},{"key":"e_1_2_10_33_1","doi-asserted-by":"publisher","DOI":"10.1002\/cne.903130404"},{"key":"e_1_2_10_34_1","doi-asserted-by":"publisher","DOI":"10.1523\/ENEURO.0383\u201017.2018"},{"key":"e_1_2_10_35_1","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1096\u20109861(20000403)419:2<205::AID\u2010CNE5>3.0.CO;2\u20100"},{"key":"e_1_2_10_36_1","volume-title":"Subicular Projections to the Hypothalamus and Brainstem: Some Novel Aspects Revealed in the Rat by the Anterograde Phaseotus Vulgaris Leukoagglutinin (PHA\u2010L) Tracing Method","author":"Kohler C.","year":"1990"},{"key":"e_1_2_10_37_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.70102"},{"key":"e_1_2_10_38_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature05453"},{"key":"e_1_2_10_39_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.70074"},{"key":"e_1_2_10_40_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1319\u201009.2009"},{"key":"e_1_2_10_41_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1191776"},{"key":"e_1_2_10_42_1","first-page":"113","article-title":"Studies on the Structure of the Cerebral Cortex. II. Continuation of the Study of the Ammonic System","volume":"46","author":"Lorente De N\u00f3 R.","year":"1934","journal-title":"Journal f\u00fcr Psychologie und Neurologie"},{"key":"e_1_2_10_43_1","doi-asserted-by":"publisher","DOI":"10.1002\/cne.24760"},{"key":"e_1_2_10_44_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00424\u2010024\u201002923\u20102"},{"key":"e_1_2_10_45_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2017.08.065"},{"issue":"1","key":"e_1_2_10_46_1","doi-asserted-by":"crossref","DOI":"10.1002\/hipo.70054","article-title":"Sex Differences in the Ventral Subiculum's Microcircuitry and Function","volume":"36","author":"Miller C. N.","year":"2026","journal-title":"Hippocampus"},{"key":"e_1_2_10_47_1","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1098\u20101063(1998)8:6<608::AID\u2010HIPO3>3.0.CO;2\u20107"},{"key":"e_1_2_10_48_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010024\u201045098\u20101"},{"issue":"3","key":"e_1_2_10_49_1","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1002\/(SICI)1096-9861(19980413)393:3<284::AID-CNE2>3.0.CO;2-Y","article-title":"Subicular Efferents Are Organized Mostly as Parallel Projections: A Double\u2010Labeling, Retrograde\u2010Tracing Study in the Rat","volume":"393","author":"Naber P. A.","year":"1998","journal-title":"Journal of Comparative Neurology"},{"key":"e_1_2_10_50_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.70110"},{"key":"e_1_2_10_51_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0301\u20100082(00)00054\u2010X"},{"key":"e_1_2_10_52_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010025\u201066613\u2010y"},{"key":"e_1_2_10_53_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010017\u201001908\u20103"},{"key":"e_1_2_10_54_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.70086"},{"key":"e_1_2_10_55_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.adj9198"},{"key":"e_1_2_10_56_1","doi-asserted-by":"crossref","DOI":"10.5962\/bhl.title.48637","volume-title":"Histologie du Syst\u00e8me Nerveux de l'homme & des Vert\u00e9br\u00e9s","author":"Ram\u00f3n y Cajal S.","year":"1909"},{"key":"e_1_2_10_57_1","doi-asserted-by":"publisher","DOI":"10.1002\/cne.25604"},{"key":"e_1_2_10_58_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrn3469"},{"key":"e_1_2_10_59_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00429\u2010022\u201002578\u20108"},{"key":"e_1_2_10_60_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrn3785"},{"key":"e_1_2_10_61_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41593\u2010019\u20100496\u2010y"},{"key":"e_1_2_10_62_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010024\u201007139\u2010z"},{"key":"e_1_2_10_63_1","doi-asserted-by":"publisher","DOI":"10.1177\/29.1A_SUPPL.7288150"},{"key":"e_1_2_10_64_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.49928"},{"key":"e_1_2_10_65_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neures.2021.01.006"},{"key":"e_1_2_10_66_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.1054"},{"key":"e_1_2_10_67_1","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.70078"},{"key":"e_1_2_10_68_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.04.007"},{"key":"e_1_2_10_69_1","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms7756"},{"key":"e_1_2_10_70_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2019.07.042"},{"key":"e_1_2_10_71_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.bbr.2006.06.022"},{"key":"e_1_2_10_72_1","doi-asserted-by":"publisher","DOI":"10.1016\/s0079\u20106123(08)61240\u20106"},{"key":"e_1_2_10_73_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.abp9186"},{"key":"e_1_2_10_74_1","doi-asserted-by":"publisher","DOI":"10.1002\/cne.24024"},{"key":"e_1_2_10_75_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586\u2010023\u201006812\u2010z"},{"key":"e_1_2_10_76_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2022.06.031"},{"key":"e_1_2_10_77_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41380\u2010025\u201003087\u2010w"},{"key":"e_1_2_10_78_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2022.1010321"}],"container-title":["Hippocampus"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/hipo.70138","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/hipo.70138","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/hipo.70138","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/hipo.70138","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T04:29:41Z","timestamp":1792038581000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/hipo.70138"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10,14]]},"references-count":77,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2026,11]]}},"alternative-id":["10.1002\/hipo.70138"],"URL":"https:\/\/doi.org\/10.1002\/hipo.70138","archive":["Portico"],"relation":{},"ISSN":["1050-9631","1098-1063"],"issn-type":[{"value":"1050-9631","type":"print"},{"value":"1098-1063","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10,14]]},"assertion":[{"value":"2026-03-30","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-09-01","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-14","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70138"}}