{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,9,5]],"date-time":"2026-09-05T05:55:18Z","timestamp":1788587718691,"version":"build-2803163510"},"posted":{"date-parts":[[2025,9,17]]},"group-title":"Neuroscience","reference-count":88,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2025,9,17]],"date-time":"2025-09-17T00:00:00Z","timestamp":1758067200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"award":["1189761 (Joshi)"],"award-info":[{"award-number":["1189761 (Joshi)"]}],"id":[{"id":"https:\/\/ror.org\/01cmst727","id-type":"ROR","asserted-by":"publisher"}]},{"award":["1283731 (Joshi)"],"award-info":[{"award-number":["1283731 (Joshi)"]}],"id":[{"id":"https:\/\/ror.org\/01cmst727","id-type":"ROR","asserted-by":"publisher"}]},{"award":["542981 (Frank)"],"award-info":[{"award-number":["542981 (Frank)"]}],"id":[{"id":"https:\/\/ror.org\/01cmst727","id-type":"ROR","asserted-by":"publisher"}]},{"award":["Joshi"],"award-info":[{"award-number":["Joshi"]}],"id":[{"id":"https:\/\/ror.org\/0195dxj21","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100023581","name":"NSF GRFP","doi-asserted-by":"crossref","award":["1650113 (Comrie)"],"award-info":[{"award-number":["1650113 (Comrie)"]}],"id":[{"id":"10.13039\/100023581","id-type":"DOI","asserted-by":"crossref"}]},{"name":"NIH","award":["F31MH124366 (Comrie)"],"award-info":[{"award-number":["F31MH124366 (Comrie)"]}]},{"name":"UCSF Jonas Cohler Discovery Fellowship","award":["Comrie"],"award-info":[{"award-number":["Comrie"]}]},{"DOI":"10.13039\/100000025","name":"National Institute of Mental Health","doi-asserted-by":"crossref","award":["F30MH126483 (Guidera)"],"award-info":[{"award-number":["F30MH126483 (Guidera)"]}],"id":[{"id":"10.13039\/100000025","id-type":"DOI","asserted-by":"crossref"}]},{"name":"UCSF Discovery 411 Fellowship","award":["Guidera"],"award-info":[{"award-number":["Guidera"]}]},{"name":"Phi Beta Kappa Graduate Award","award":["Guidera"],"award-info":[{"award-number":["Guidera"]}]},{"award":["Frank"],"award-info":[{"award-number":["Frank"]}],"id":[{"id":"https:\/\/ror.org\/006w34k90","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,7,23]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>The ability to rapidly learn and retrieve salient information about new environments is critical for survival. In mammals, the hippocampus plays a crucial role in that learning. Specialized features of hippocampal population coding, including network-level theta oscillatory activity, location-specific firing of principal cells, and reactivation of experience during immobility (replay), have been implicated in rapid storage and retrieval of spatial information. Disruptions of theta and replay jointly, or replay alone, are sufficient to impair learning; however, the specific contribution of theta-associated temporal structure during locomotion remains unknown. In this study, we manipulated hippocampal spiking activity in rats specifically during locomotion by optogenetically activating septal parvalbumin-expressing GABAergic neurons. We developed a closed-loop, theta phase-specific stimulation protocol that reliably reduced theta power shortly after stimulation onset. This manipulation preserved the place codes of individual cells but disrupted the fine temporal structure of endogenous spatio-temporal representations (i.e., theta sequences) at the pairwise and population level. Theta disruption during locomotion was also sufficient to cause pronounced deficits in learning the more cognitively challenging component of a spatial alternation task, even though disruption was applied on only \u223c66% of trials. Notably, network effects accompanying theta disruption were restricted to locomotor periods; we did not observe changes in replay rate, length, or content during immobility. Together, these results demonstrate the importance of the precise temporal microstructure of locomotion-associated spatial representations in the hippocampus for learning.<\/jats:p>","DOI":"10.1101\/2025.09.15.675587","type":"posted-content","created":{"date-parts":[[2025,9,17]],"date-time":"2025-09-17T13:10:12Z","timestamp":1758114612000},"source":"Crossref","is-referenced-by-count":7,"title":["Disruption of theta-timescale spiking impairs learning but spares hippocampal replay"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0511-3747","authenticated-orcid":false,"given":"Abhilasha","family":"Joshi","sequence":"first","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"},{"name":"Champalimaud Center for the Unknown, Lisbon, Portugal"},{"name":"National Center for Biological Sciences, Bangalore, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2251-4892","authenticated-orcid":false,"given":"Alison E.","family":"Comrie","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"},{"name":"Kavli Institute for Fundamental Neuroscience, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9696-6920","authenticated-orcid":false,"given":"Samuel","family":"Bray","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0620-259X","authenticated-orcid":false,"given":"Abhijith","family":"Mankili","sequence":"additional","affiliation":[{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4013-214X","authenticated-orcid":false,"given":"Jennifer A.","family":"Guidera","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"},{"name":"Kavli Institute for Fundamental Neuroscience, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rhino","family":"Nevers","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"},{"name":"Kavli Institute for Fundamental Neuroscience, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3076-0994","authenticated-orcid":false,"given":"Xulu","family":"Sun","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Emily","family":"Monroe","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Viktor","family":"Kharazia","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9238-0642","authenticated-orcid":false,"given":"Ryan","family":"Ly","sequence":"additional","affiliation":[{"name":"Lawrence Berkeley National Laboratory, Berkeley, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniela Astudillo","family":"Maya","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Denisse","family":"Morales-Rodriguez","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"},{"name":"Kavli Institute for Fundamental Neuroscience, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jai","family":"Yu","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anna","family":"Kiseleva","sequence":"additional","affiliation":[{"name":"Kavli Institute for Fundamental Neuroscience, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Victor","family":"Perez","sequence":"additional","affiliation":[{"name":"Kavli Institute for Fundamental Neuroscience, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1752-5677","authenticated-orcid":false,"given":"Loren M.","family":"Frank","sequence":"additional","affiliation":[{"name":"Department of Physiology, University of California, San Francisco, CA, USA"},{"name":"Howard Hughes Medical Institute, University of California, San Francisco, CA, USA"},{"name":"Kavli Institute for Fundamental Neuroscience, University of California, San Francisco, CA, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026072719400618000_2025.09.15.675587v2.1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tins.2015.07.004"},{"key":"2026072719400618000_2025.09.15.675587v2.2","doi-asserted-by":"crossref","first-page":"8521","DOI":"10.1038\/ncomms9521","article-title":"Theta oscillations regulate the speed of locomotion via a hippocampus to lateral septum pathway","volume":"6","year":"2015","journal-title":"Nature Communications"},{"key":"2026072719400618000_2025.09.15.675587v2.3","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0190-06.2006"},{"key":"2026072719400618000_2025.09.15.675587v2.4","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2014.04.013"},{"key":"2026072719400618000_2025.09.15.675587v2.5","doi-asserted-by":"publisher","DOI":"10.1016\/0031-9384(77)90035-X"},{"key":"2026072719400618000_2025.09.15.675587v2.6","doi-asserted-by":"publisher","DOI":"10.1016\/S0893-6080(05)80159-5"},{"key":"2026072719400618000_2025.09.15.675587v2.7","doi-asserted-by":"publisher","DOI":"10.1016\/0306-4522(89)90423-5"},{"key":"2026072719400618000_2025.09.15.675587v2.8","doi-asserted-by":"publisher","DOI":"10.1016\/S0896-6273(02)00586-X"},{"key":"2026072719400618000_2025.09.15.675587v2.9","doi-asserted-by":"publisher","DOI":"10.1016\/0165-0173(83)90037-1"},{"key":"2026072719400618000_2025.09.15.675587v2.10","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2021.0336"},{"key":"2026072719400618000_2025.09.15.675587v2.11","doi-asserted-by":"crossref","unstructured":"Comrie, A. E. , Monroe, E. J. , Kahn, A. E. , Denovellis, E. L. , Joshi, A. , Guidera, J. A. , Krausz, T. A. , Berke, J. D. , Daw, N. D. , and Frank, L. M . (2024). Hippocampal representations of alternative possibilities are flexibly generated to meet cognitive demands. bioRxiv.","DOI":"10.1101\/2024.09.23.613567"},{"key":"2026072719400618000_2025.09.15.675587v2.12","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2009.07.027"},{"key":"2026072719400618000_2025.09.15.675587v2.13","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.64505"},{"key":"2026072719400618000_2025.09.15.675587v2.14","doi-asserted-by":"publisher","DOI":"10.1038\/nn1961"},{"key":"2026072719400618000_2025.09.15.675587v2.15","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1978-24.2025"},{"key":"2026072719400618000_2025.09.15.675587v2.16","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2006.02.023"},{"key":"2026072719400618000_2025.09.15.675587v2.17","doi-asserted-by":"publisher","DOI":"10.1126\/science.aat2952"},{"key":"2026072719400618000_2025.09.15.675587v2.18","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.20707"},{"key":"2026072719400618000_2025.09.15.675587v2.19","doi-asserted-by":"publisher","DOI":"10.1038\/nrn.2017.74"},{"issue":"1","key":"2026072719400618000_2025.09.15.675587v2.20","doi-asserted-by":"crossref","first-page":"410","DOI":"10.1038\/s41467-023-35825-5","article-title":"Optogenetic frequency scrambling of hippocampal theta oscillations dissociates working memory retrieval from hippocampal spatiotemporal codes","volume":"14","year":"2023","journal-title":"Nature Communications"},{"key":"2026072719400618000_2025.09.15.675587v2.21","doi-asserted-by":"publisher","DOI":"10.1126\/science.abh4272"},{"key":"2026072719400618000_2025.09.15.675587v2.22","doi-asserted-by":"publisher","DOI":"10.1126\/science.aax0758"},{"key":"2026072719400618000_2025.09.15.675587v2.23","doi-asserted-by":"publisher","DOI":"10.1038\/nature04587"},{"key":"2026072719400618000_2025.09.15.675587v2.24","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.20345"},{"key":"2026072719400618000_2025.09.15.675587v2.25","doi-asserted-by":"publisher","DOI":"10.1038\/336170a0"},{"key":"2026072719400618000_2025.09.15.675587v2.26","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2015.05.001"},{"key":"2026072719400618000_2025.09.15.675587v2.27","doi-asserted-by":"publisher","DOI":"10.1101\/lm.053348.120"},{"key":"2026072719400618000_2025.09.15.675587v2.28","doi-asserted-by":"crossref","first-page":"RP90944","DOI":"10.7554\/eLife.90944","article-title":"Neurofeedback training can modulate task-relevant memory replay rate in rats","volume":"12","year":"2024","journal-title":"eLife"},{"key":"2026072719400618000_2025.09.15.675587v2.29","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2021.07.029"},{"key":"2026072719400618000_2025.09.15.675587v2.30","doi-asserted-by":"publisher","DOI":"10.1038\/nn.3138"},{"key":"2026072719400618000_2025.09.15.675587v2.31","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.23676"},{"key":"2026072719400618000_2025.09.15.675587v2.32","doi-asserted-by":"publisher","DOI":"10.1162\/089976602317318965"},{"key":"2026072719400618000_2025.09.15.675587v2.33","doi-asserted-by":"publisher","DOI":"10.1038\/nature14396"},{"key":"2026072719400618000_2025.09.15.675587v2.34","doi-asserted-by":"publisher","DOI":"10.1126\/science.1217230"},{"key":"2026072719400618000_2025.09.15.675587v2.35","doi-asserted-by":"publisher","DOI":"10.1101\/lm.3.2-3.264"},{"key":"2026072719400618000_2025.09.15.675587v2.36","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3761-07.2007"},{"key":"2026072719400618000_2025.09.15.675587v2.37","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7793.1999.0867s.x"},{"key":"2026072719400618000_2025.09.15.675587v2.38","doi-asserted-by":"publisher","DOI":"10.1038\/s41583-018-0077-1"},{"key":"2026072719400618000_2025.09.15.675587v2.39","unstructured":"Joshi, A. (2017). Behaviour-Dependent Activity and Synaptic Organisation of Septo-Hippocampal GABAergic Neurons. PhD thesis, University of Oxford."},{"key":"2026072719400618000_2025.09.15.675587v2.40","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-023-05928-6"},{"key":"2026072719400618000_2025.09.15.675587v2.41","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2017.10.033"},{"key":"2026072719400618000_2025.09.15.675587v2.42","doi-asserted-by":"publisher","DOI":"10.1038\/nn.2344"},{"issue":"1","key":"2026072719400618000_2025.09.15.675587v2.43","doi-asserted-by":"crossref","first-page":"17903","DOI":"10.1038\/s41598-025-01695-8","article-title":"Pedunculopontine-stimulation obstructs hippocampal theta rhythm and halts movement","volume":"15","year":"2025","journal-title":"Scientific Reports"},{"key":"2026072719400618000_2025.09.15.675587v2.44","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2020.01.014"},{"key":"2026072719400618000_2025.09.15.675587v2.45","doi-asserted-by":"publisher","DOI":"10.1038\/nature17144"},{"key":"2026072719400618000_2025.09.15.675587v2.46","doi-asserted-by":"publisher","DOI":"10.1126\/science.1585183"},{"key":"2026072719400618000_2025.09.15.675587v2.47","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0005494"},{"issue":"1","key":"2026072719400618000_2025.09.15.675587v2.48","doi-asserted-by":"crossref","first-page":"6159","DOI":"10.1038\/s41467-023-41746-0","article-title":"The medial septum controls hippocampal supra-theta oscillations","volume":"14","year":"2023","journal-title":"Nature Communications"},{"issue":"6","key":"2026072719400618000_2025.09.15.675587v2.49","doi-asserted-by":"crossref","first-page":"1535","DOI":"10.1016\/j.brs.2020.08.011","article-title":"Disruption of hippocampal rhythms via optogenetic stimulation during the critical period for memory development impairs spatial cognition","volume":"13","year":"2020","journal-title":"Brain Stimulation"},{"key":"2026072719400618000_2025.09.15.675587v2.50","doi-asserted-by":"publisher","DOI":"10.1126\/science.1201685"},{"key":"2026072719400618000_2025.09.15.675587v2.51","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0027575"},{"key":"2026072719400618000_2025.09.15.675587v2.52","doi-asserted-by":"crossref","unstructured":"Lee, K. H. , Denovellis, E. L. , Ly, R. , Magland, J. , Soules, J. , Comrie, A. E. , Gramling, D. P. , Guidera, J. A. , Nevers, R. , Adenekan, P. , Brozdowski, C. , Bray, S. R. , Monroe, E. , Bak, J. H. , Coulter, M. E. , Sun, X. , Broyles, E. , Shin, D. , Chiang, S. , Holobetz, C. , Tritt, A. , R\u00fcbel, O. , Nguyen, T. , Yatsenko, D. , Chu, J. , Kemere, C. , Garcia, S. , Buccino, A. , and Frank, L. M . (2024). Spyglass: A framework for reproducible and shareable neuroscience research. bioRxiv preprint.","DOI":"10.1101\/2024.01.25.577295"},{"key":"2026072719400618000_2025.09.15.675587v2.53","doi-asserted-by":"publisher","DOI":"10.1126\/sciadv.abd5684"},{"key":"2026072719400618000_2025.09.15.675587v2.54","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2013.03.007"},{"key":"2026072719400618000_2025.09.15.675587v2.55","doi-asserted-by":"publisher","DOI":"10.1126\/science.adi8237"},{"key":"2026072719400618000_2025.09.15.675587v2.56","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2019.02.014"},{"key":"2026072719400618000_2025.09.15.675587v2.57","doi-asserted-by":"publisher","DOI":"10.1016\/0006-8993(90)90188-H"},{"key":"2026072719400618000_2025.09.15.675587v2.58","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2009.08.037"},{"key":"2026072719400618000_2025.09.15.675587v2.59","doi-asserted-by":"publisher","DOI":"10.1038\/297681a0"},{"issue":"11","key":"2026072719400618000_2025.09.15.675587v2.60","doi-asserted-by":"crossref","first-page":"1167","DOI":"10.1002\/hipo.23248","article-title":"Optogenetic \u201clow-theta\u201d pacing of the septohippocampal circuit is sufficient for spatial goal finding and is influenced by behavioral state and cognitive demand","volume":"30","year":"2020","journal-title":"Hippocampus"},{"key":"2026072719400618000_2025.09.15.675587v2.61","doi-asserted-by":"publisher","DOI":"10.1016\/0014-4886(76)90055-8"},{"key":"2026072719400618000_2025.09.15.675587v2.62","doi-asserted-by":"publisher","DOI":"10.1002\/hipo.450030307"},{"key":"2026072719400618000_2025.09.15.675587v2.63","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2018.04.006"},{"key":"2026072719400618000_2025.09.15.675587v2.64","doi-asserted-by":"publisher","DOI":"10.1126\/science.1159775"},{"key":"2026072719400618000_2025.09.15.675587v2.65","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2020.05.023"},{"key":"2026072719400618000_2025.09.15.675587v2.66","doi-asserted-by":"publisher","DOI":"10.1016\/0013-4694(62)90030-5"},{"key":"2026072719400618000_2025.09.15.675587v2.67","doi-asserted-by":"publisher","DOI":"10.1038\/nature12112"},{"key":"2026072719400618000_2025.09.15.675587v2.68","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-021-00919-0"},{"key":"2026072719400618000_2025.09.15.675587v2.69","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2407-09.2009"},{"key":"2026072719400618000_2025.09.15.675587v2.70","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2141-15.2016"},{"key":"2026072719400618000_2025.09.15.675587v2.71","doi-asserted-by":"publisher","DOI":"10.1016\/j.tins.2023.06.004"},{"issue":"8","key":"2026072719400618000_2025.09.15.675587v2.72","first-page":"114529","article-title":"Optogenetic silencing of medial septal GABAergic neurons disrupts grid cell spatial and temporal coding in the medial entorhinal cortex","volume":"43","year":"2024","journal-title":"Cell Reports"},{"key":"2026072719400618000_2025.09.15.675587v2.73","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.78362"},{"key":"2026072719400618000_2025.09.15.675587v2.74","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3024-18.2019"},{"key":"2026072719400618000_2025.09.15.675587v2.75","doi-asserted-by":"publisher","DOI":"10.1136\/jnnp.20.1.11"},{"key":"2026072719400618000_2025.09.15.675587v2.76","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.03061"},{"key":"2026072719400618000_2025.09.15.675587v2.77","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1098-1063(1996)6:2<149::AID-HIPO6>3.0.CO;2-K"},{"key":"2026072719400618000_2025.09.15.675587v2.78","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2908-03.2004"},{"key":"2026072719400618000_2025.09.15.675587v2.79","doi-asserted-by":"publisher","DOI":"10.1038\/s41593-023-01306-7"},{"key":"2026072719400618000_2025.09.15.675587v2.80","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1210929109"},{"key":"2026072719400618000_2025.09.15.675587v2.81","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2023.10.001"},{"key":"2026072719400618000_2025.09.15.675587v2.82","doi-asserted-by":"publisher","DOI":"10.1038\/nn.3550"},{"key":"2026072719400618000_2025.09.15.675587v2.83","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.34395"},{"key":"2026072719400618000_2025.09.15.675587v2.84","doi-asserted-by":"publisher","DOI":"10.1038\/nn.3904"},{"key":"2026072719400618000_2025.09.15.675587v2.85","doi-asserted-by":"crossref","first-page":"e20697","DOI":"10.7554\/eLife.20697","article-title":"Synchronized excitability in a network enables generation of internal neuronal sequences","volume":"5","year":"2016","journal-title":"eLife"},{"key":"2026072719400618000_2025.09.15.675587v2.86","doi-asserted-by":"crossref","unstructured":"Yong, H. C. , Chang, H. , and Brandon, M. P. (2022). Optogenetic reduction of theta oscillations reveals that a single reliable time cell sequence is not required for working memory. bioRxiv preprint.","DOI":"10.1101\/2022.06.25.497592"},{"key":"2026072719400618000_2025.09.15.675587v2.87","doi-asserted-by":"crossref","unstructured":"Yu, J. Y. , Pettibone, J. R. , Guo, C. , Zhang, S. , Saunders, T. L. , Hughes, E. D. , Filipiak, W. E. , Zeidler, M. G. , Bender, K. J. , Hopf, F. , Smyth, C. N. , Kharazia, V. , Kiseleva, A. , Davidson, T. J. , Frank, L. M. , and Berke, J. D. (2018). Knock-in rats expressing Cre and Flp recombinases at the parvalbumin locus. bioRxiv preprint.","DOI":"10.1101\/386474"},{"key":"2026072719400618000_2025.09.15.675587v2.88","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2018.02.061"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1101\/2025.09.15.675587","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,28]],"date-time":"2026-07-28T02:40:20Z","timestamp":1785206420000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.1101\/2025.09.15.675587"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,9,17]]},"references-count":88,"URL":"https:\/\/doi.org\/10.1101\/2025.09.15.675587","relation":{"has-version":[{"id-type":"doi","id":"10.7554\/eLife.109375.1","asserted-by":"object"}]},"subject":[],"published":{"date-parts":[[2025,9,17]]},"subtype":"preprint"}}