{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,6]],"date-time":"2026-10-06T02:36:11Z","timestamp":1791254171020,"version":"4.1.0"},"reference-count":203,"publisher":"The Royal Society","issue":"1954","license":[{"start":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T00:00:00Z","timestamp":1783555200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T00:00:00Z","timestamp":1783555200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000025","name":"National Institute of Mental Health","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000025","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000006","name":"Office of Naval Research","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000006","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,7,9]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Planning of future trajectories through the environment may rely on our capacity to encode and recall episodic memories that include spatial representations. Here, we describe potential mechanisms for behaviour based on episodic memory that would use neural trajectory scanning to explore and select trajectories towards remembered goal locations. This model is consistent with rodent neurophysiological data recorded in the medial entorhinal cortex that demonstrated alternating left\u2013right scans of space and data recorded in the hippocampus of place cell firing that showed alternating scans, or scans directed towards a goal location. In a related model, sequential activation of different directions along trajectories would allow encoding and retrieval of complex sequences of vectors that correspond to episodic memories of events at a specific time and place. Trajectory scanning may also detect and avoid potential collisions with representations of visible or remembered objects and barriers in allocentric or egocentric reference frameworks to plan trajectories. Moreover, the same trajectory scanning in egocentric space could generate egocentric neural responses such as egocentric boundary cells and egocentric bearing cells consistent with recent human imaging experiments. Together, these theoretical mechanisms provide a predictive coding framework linking allocentric navigation, episodic memory retrieval, and egocentric obstacle representations.<\/jats:p>\n                  <jats:p>This article is part of the theme issue \u2018The role of hippocampal predictions in cognition: bridging perception and memory\u2019.<\/jats:p>","DOI":"10.1098\/rstb.2025.0251","type":"journal-article","created":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T08:09:03Z","timestamp":1783584543000},"source":"Crossref","is-referenced-by-count":2,"title":["Trajectory scanning as a predictive coding mechanism for goal-directed navigation, obstacle avoidance and episodic memory"],"prefix":"10.1098","volume":"381","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9397-432X","authenticated-orcid":false,"given":"Jennifer C.","family":"Robinson","sequence":"first","affiliation":[{"name":"Boston University, 1","place":["Boston,, MA 02215,, USA"],"department":["Center for Systems Neuroscience,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Patrick A.","family":"LaChance","sequence":"additional","affiliation":[{"name":"Boston University, 1","place":["Boston,, MA 02215,, USA"],"department":["Center for Systems Neuroscience,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Samantha","family":"Malmberg","sequence":"additional","affiliation":[{"name":"Boston University, 1","place":["Boston,, MA 02215,, USA"],"department":["Center for Systems Neuroscience,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-9100-0445","authenticated-orcid":false,"given":"Jiarui","family":"Fang","sequence":"additional","affiliation":[{"name":"Boston University, 1","place":["Boston,, MA 02215,, USA"],"department":["Center for Systems Neuroscience,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shruthi","family":"Sankaranarayanan","sequence":"additional","affiliation":[{"name":"Boston University, 1","place":["Boston,, MA 02215,, USA"],"department":["Center for Systems Neuroscience,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chantal E.","family":"Stern","sequence":"additional","affiliation":[{"name":"Boston University, 1","place":["Boston,, MA 02215,, USA"],"department":["Center for Systems Neuroscience,"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9925-6377","authenticated-orcid":false,"given":"Michael E.","family":"Hasselmo","sequence":"additional","affiliation":[{"name":"Boston University, 1","place":["Boston,, MA 02215,, USA"],"department":["Center for Systems Neuroscience,"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"175","published-online":{"date-parts":[[2026,7,9]]},"reference":[{"key":"2026070904085472300_R1","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1016\/0014-4886(76)90055-8","article-title":"Place units in the hippocampus of the freely moving rat","volume":"51","author":"O\u2019Keefe","year":"1976","journal-title":"Exp. Neurol."},{"key":"2026070904085472300_R2","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1038\/nature03721","article-title":"Microstructure of a spatial map in the entorhinal cortex","volume":"436","author":"Hafting","year":"2005","journal-title":"Nature"},{"key":"2026070904085472300_R3","doi-asserted-by":"crossref","first-page":"4602","DOI":"10.1073\/pnas.0407332102","article-title":"Coding for spatial goals in the prelimbic\/infralimbic area of the rat frontal cortex","volume":"102","author":"Hok","year":"2005","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026070904085472300_R4","doi-asserted-by":"crossref","DOI":"10.1016\/j.nlm.2020.107215","article-title":"Dorsomedial prefrontal cortex and hippocampus represent strategic context even while simultaneously changing representation throughout a task session","volume":"171","author":"Hasz","year":"2020","journal-title":"Neurobiol. Learn. Mem."},{"key":"2026070904085472300_R5","doi-asserted-by":"crossref","first-page":"9771","DOI":"10.1523\/JNEUROSCI.1319-09.2009","article-title":"Boundary vector cells in the subiculum of the hippocampal formation","volume":"29","author":"Lever","year":"2009","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R6","doi-asserted-by":"crossref","first-page":"69","DOI":"10.3389\/fnbeh.2011.00069","article-title":"Representation of non-spatial and spatial information in the lateral entorhinal cortex","volume":"5","author":"Deshmukh","year":"2011","journal-title":"Front. Behav. Neurosci."},{"key":"2026070904085472300_R7","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1002\/hipo.22101","article-title":"Influence of local objects on hippocampal representations: landmark vectors and memory","volume":"23","author":"Deshmukh","year":"2013","journal-title":"Hippocampus"},{"key":"2026070904085472300_R8","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.aaz2322","article-title":"Egocentric boundary vector tuning of the retrosplenial cortex","volume":"6","author":"Alexander","year":"2020","journal-title":"Sci. Adv."},{"key":"2026070904085472300_R9","doi-asserted-by":"crossref","DOI":"10.1126\/science.aax4192","article-title":"A sense of space in postrhinal cortex","volume":"365","author":"LaChance","year":"2019","journal-title":"Science"},{"key":"2026070904085472300_R10","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1523\/JNEUROSCI.10-02-00420.1990","article-title":"Head-direction cells recorded from the postsubiculum in freely moving rats. I. Description and quantitative analysis","volume":"10","author":"Taube","year":"1990","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R11","doi-asserted-by":"crossref","first-page":"739","DOI":"10.1038\/nn.3383","article-title":"Segregation of cortical head direction cell assemblies on alternating theta cycles","volume":"16","author":"Brandon","year":"2013","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R12","doi-asserted-by":"crossref","first-page":"995","DOI":"10.1038\/s41586-024-08527-1","article-title":"Left\u2013right-alternating theta sweeps in entorhinal\u2013hippocampal maps of space","volume":"639","author":"Vollan","year":"2025","journal-title":"Nature"},{"key":"2026070904085472300_R13","doi-asserted-by":"crossref","first-page":"408","DOI":"10.1016\/0166-2236(87)90011-7","article-title":"Hippocampal synaptic enhancement and information storage within a distributed memory system","volume":"10","author":"McNaughton","year":"1987","journal-title":"Trends Neurosci."},{"key":"2026070904085472300_R14","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1002\/(SICI)1098-1063(1996)6:3<247::AID-HIPO4>3.0.CO;2-K","article-title":"Theory of rodent navigation based on interacting representations of space","volume":"6","author":"Touretzky","year":"1996","journal-title":"Hippocampus"},{"key":"2026070904085472300_R15","doi-asserted-by":"crossref","first-page":"5900","DOI":"10.1523\/JNEUROSCI.17-15-05900.1997","article-title":"Path integration and cognitive mapping in a continuous attractor neural network model","volume":"17","author":"Samsonovich","year":"1997","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R16","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1162\/089976698300017908","article-title":"The role of the hippocampus in solving the Morris water maze","volume":"10","author":"Redish","year":"1998","journal-title":"Neural Comput."},{"key":"2026070904085472300_R17","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1007\/s004220000171","article-title":"Spatial cognition and neuro-mimetic navigation: a model of hippocampal place cell activity","volume":"83","author":"Arleo","year":"2000","journal-title":"Biol. Cybern."},{"key":"2026070904085472300_R18","doi-asserted-by":"crossref","first-page":"340","DOI":"10.1037\/0033-295X.114.2.340","article-title":"Remembering the past and imagining the future: a neural model of spatial memory and imagery","volume":"114","author":"Byrne","year":"2007","journal-title":"Psychol. Rev."},{"key":"2026070904085472300_R19","doi-asserted-by":"crossref","first-page":"540","DOI":"10.1037\/a0016170","article-title":"Is there a geometric module for spatial orientation? Insights from a rodent navigation model","volume":"116","author":"Sheynikhovich","year":"2009","journal-title":"Psychol. Rev."},{"key":"2026070904085472300_R20","doi-asserted-by":"crossref","first-page":"916","DOI":"10.1111\/j.1460-9568.2012.08015.x","article-title":"A goal-directed spatial navigation model using forward trajectory planning based on grid cells","volume":"35","author":"Erdem","year":"2012","journal-title":"Eur. J. Neurosci."},{"key":"2026070904085472300_R21","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1016\/j.jphysparis.2013.07.002","article-title":"A biologically inspired hierarchical goal directed navigation model","volume":"108","author":"Erdem","year":"2014","journal-title":"J. Physiol. Paris"},{"key":"2026070904085472300_R22","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.33752","article-title":"A neural-level model of spatial memory and imagery","volume":"7","author":"Bicanski","year":"2018","journal-title":"eLife"},{"key":"2026070904085472300_R23","doi-asserted-by":"crossref","first-page":"12176","DOI":"10.1523\/JNEUROSCI.3761-07.2007","article-title":"Neural ensembles in CA3 transiently encode paths forward of the animal at a decision point","volume":"27","author":"Johnson","year":"2007","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R24","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1038\/nrn.2015.30","article-title":"Vicarious trial and error","volume":"17","author":"Redish","year":"2016","journal-title":"Nat. Rev. Neurosci."},{"key":"2026070904085472300_R25","doi-asserted-by":"crossref","first-page":"552","DOI":"10.1016\/j.cell.2020.01.014","article-title":"Constant sub-second cycling between representations of possible futures in the hippocampus","volume":"180","author":"Kay","year":"2020","journal-title":"Cell"},{"key":"2026070904085472300_R26","doi-asserted-by":"crossref","first-page":"20","DOI":"10.3389\/fncir.2012.00020","article-title":"Linear look-ahead in conjunctive cells: an entorhinal mechanism for vector-based navigation","volume":"6","author":"Kubie","year":"2012","journal-title":"Front. Neural Circuits"},{"key":"2026070904085472300_R27","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1038\/nature11649","article-title":"The entorhinal grid map is discretized","volume":"492","author":"Stensola","year":"2012","journal-title":"Nature"},{"key":"2026070904085472300_R28","doi-asserted-by":"crossref","first-page":"6858","DOI":"10.1523\/JNEUROSCI.5684-07.2008","article-title":"What grid cells convey about rat location","volume":"28","author":"Fiete","year":"2008","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R29","doi-asserted-by":"crossref","first-page":"1330","DOI":"10.1038\/nn.2901","article-title":"Grid cells generate an analog error-correcting code for singularly precise neural computation","volume":"14","author":"Sreenivasan","year":"2011","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R30","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1038\/nature05601","article-title":"Hippocampal remapping and grid realignment in entorhinal cortex","volume":"446","author":"Fyhn","year":"2007","journal-title":"Nature"},{"key":"2026070904085472300_R31","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1038\/297681a0","article-title":"Place navigation impaired in rats with hippocampal lesions","volume":"297","author":"Morris","year":"1982","journal-title":"Nature"},{"key":"2026070904085472300_R32","doi-asserted-by":"crossref","first-page":"1129","DOI":"10.1016\/j.neunet.2009.07.012","article-title":"A phase code for memory could arise from circuit mechanisms in entorhinal cortex","volume":"22","author":"Hasselmo","year":"2009","journal-title":"Neural Netw."},{"key":"2026070904085472300_R33","doi-asserted-by":"crossref","first-page":"1157","DOI":"10.1002\/hipo.20518","article-title":"Grid cells and theta as oscillatory interference: theory and predictions","volume":"18","author":"Burgess","year":"2008","journal-title":"Hippocampus"},{"key":"2026070904085472300_R34","doi-asserted-by":"crossref","first-page":"709","DOI":"10.1016\/j.cub.2024.08.059","article-title":"A systems model of alternating theta sweeps via firing rate adaptation","volume":"35","author":"Ji","year":"2025","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R35","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2025.115475","article-title":"Spontaneous alternation of place-cell sequences in the open field through spike frequency adaptation","volume":"44","author":"Widloski","year":"2025","journal-title":"Cell Rep."},{"key":"2026070904085472300_R36","doi-asserted-by":"crossref","first-page":"682","DOI":"10.1038\/nn1905","article-title":"Experience-dependent rescaling of entorhinal grids","volume":"10","author":"Barry","year":"2007","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R37","doi-asserted-by":"crossref","first-page":"1200","DOI":"10.1002\/hipo.20504","article-title":"Progressive increase in grid scale from dorsal to ventral medial entorhinal cortex","volume":"18","author":"Brun","year":"2008","journal-title":"Hippocampus"},{"key":"2026070904085472300_R38","doi-asserted-by":"crossref","first-page":"7347","DOI":"10.1523\/JNEUROSCI.14-12-07347.1994","article-title":"Comparison of spatial firing characteristics of units in dorsal and ventral hippocampus of the rat","volume":"14","author":"Jung","year":"1994","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R39","doi-asserted-by":"crossref","first-page":"140","DOI":"10.1126\/science.1157086","article-title":"Finite scale of spatial representation in the hippocampus","volume":"321","author":"Kjelstrup","year":"2008","journal-title":"Science"},{"key":"2026070904085472300_R40","doi-asserted-by":"crossref","first-page":"8079","DOI":"10.1523\/JNEUROSCI.5458-12.2013","article-title":"Ventral hippocampal neurons are shaped by experience to represent behaviorally relevant contexts","volume":"33","author":"Komorowski","year":"2013","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R41","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1016\/j.nlm.2014.07.003","article-title":"A hierarchical model of goal directed navigation selects trajectories in a visual environment","volume":"117","author":"Erdem","year":"2015","journal-title":"Neurobiol. Learn. Mem."},{"key":"2026070904085472300_R42","doi-asserted-by":"crossref","first-page":"709","DOI":"10.1002\/(SICI)1098-1063(1996)6:6<709::AID-HIPO13>3.0.CO;2-4","article-title":"Hippocampal place cells connected by Hebbian synapses can solve spatial problems","volume":"6","author":"Muller","year":"1996","journal-title":"Hippocampus"},{"key":"2026070904085472300_R43","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1085\/jgp.107.6.663","article-title":"The hippocampus as a cognitive graph","volume":"107","author":"Muller","year":"1996","journal-title":"J. Gen. Physiol."},{"key":"2026070904085472300_R44","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/(SICI)1098-1063(2000)10:1<1::AID-HIPO1>3.0.CO;2-1","article-title":"A model of hippocampally dependent navigation, using the temporal difference learning rule","volume":"10","author":"Foster","year":"2000","journal-title":"Hippocampus"},{"key":"2026070904085472300_R45","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1016\/0006-8993(71)90358-1","article-title":"The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat","volume":"34","author":"O\u2019Keefe","year":"1971","journal-title":"Brain Res."},{"key":"2026070904085472300_R46","doi-asserted-by":"crossref","first-page":"1935","DOI":"10.1523\/JNEUROSCI.07-07-01935.1987","article-title":"Spatial firing patterns of hippocampal complex-spike cells in a fixed environment","volume":"7","author":"Muller","year":"1987","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R47","doi-asserted-by":"crossref","first-page":"11250","DOI":"10.1523\/JNEUROSCI.2862-08.2008","article-title":"Unmasking the CA1 ensemble place code by exposures to small and large environments: more place cells and multiple, irregularly arranged, and expanded place fields in the larger space","volume":"28","author":"Fenton","year":"2008","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R48","doi-asserted-by":"crossref","first-page":"2178","DOI":"10.1016\/j.cub.2021.03.003","article-title":"Dorsal CA1 hippocampal place cells form a multi-scale representation of megaspace","volume":"31","author":"Harland","year":"2021","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R49","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/j.neuron.2006.02.023","article-title":"Temporal encoding of place sequences by hippocampal cell assemblies","volume":"50","author":"Dragoi","year":"2006","journal-title":"Neuron"},{"key":"2026070904085472300_R50","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.06063","article-title":"Hippocampal place cells construct reward related sequences through unexplored space","volume":"4","author":"\u00d3lafsd\u00f3ttir","year":"2015","journal-title":"eLife"},{"key":"2026070904085472300_R51","doi-asserted-by":"crossref","first-page":"1347","DOI":"10.1002\/hipo.23246","article-title":"Neurobiological successor features for spatial navigation","volume":"30","author":"de Cothi","year":"2020","journal-title":"Hippocampus"},{"key":"2026070904085472300_R52","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1016\/S0925-2312(02)00395-8","article-title":"A model of hippocampal circuitry mediating goal-driven navigation in a familiar environment","volume":"44\u201346","author":"Gorchetchnikov","year":"2002","journal-title":"Neurocomputing"},{"key":"2026070904085472300_R53","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1080\/09540090500140925","article-title":"A biophysical implementation of a bidirectional graph search algorithm to solve multiple goal navigation tasks","volume":"17","author":"Gorchetchnikov","year":"2005","journal-title":"Connect. Sci."},{"key":"2026070904085472300_R54","doi-asserted-by":"crossref","first-page":"429","DOI":"10.1038\/s41586-018-0102-6","article-title":"Vector-based navigation using grid-like representations in artificial agents","volume":"557","author":"Banino","year":"2018","journal-title":"Nature"},{"key":"2026070904085472300_R55","doi-asserted-by":"crossref","first-page":"1951","DOI":"10.1523\/JNEUROSCI.07-07-01951.1987","article-title":"The effects of changes in the environment on the spatial firing of hippocampal complex-spike cells","volume":"7","author":"Muller","year":"1987","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R56","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1038\/416090a","article-title":"Long-term plasticity in hippocampal place-cell representation of environmental geometry","volume":"416","author":"Lever","year":"2002","journal-title":"Nature"},{"key":"2026070904085472300_R57","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1002\/(SICI)1098-1063(1997)7:1<15::AID-HIPO3>3.0.CO;2-6","article-title":"Cognitive maps beyond the hippocampus","volume":"7","author":"Redish","year":"1997","journal-title":"Hippocampus"},{"key":"2026070904085472300_R58","doi-asserted-by":"crossref","first-page":"165","DOI":"10.1002\/hipo.450020208","article-title":"Navigation by fragment fitting: a theory of hippocampal function","volume":"2","author":"Worden","year":"1992","journal-title":"Hippocampus"},{"key":"2026070904085472300_R59","doi-asserted-by":"crossref","first-page":"3531","DOI":"10.1523\/JNEUROSCI.10-11-03531.1990","article-title":"Hippocampal representation in place learning","volume":"10","author":"Eichenbaum","year":"1990","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R60","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1002\/(SICI)1098-1063(1999)9:2<118::AID-HIPO4>3.0.CO;2-8","article-title":"Delay-dependent impairment of a matching-to-place task with chronic and intrahippocampal infusion of the NMDA-antagonist D-AP5","volume":"9","author":"Steele","year":"1999","journal-title":"Hippocampus"},{"key":"2026070904085472300_R61","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1016\/S0896-6273(00)00018-0","article-title":"Trajectory encoding in the hippocampus and entorhinal cortex","volume":"27","author":"Frank","year":"2000","journal-title":"Neuron"},{"key":"2026070904085472300_R62","doi-asserted-by":"crossref","first-page":"623","DOI":"10.1016\/S0896-6273(00)00071-4","article-title":"Hippocampal neurons encode information about different types of memory episodes occurring in the same location","volume":"27","author":"Wood","year":"2000","journal-title":"Neuron"},{"key":"2026070904085472300_R63","doi-asserted-by":"crossref","first-page":"1454","DOI":"10.1126\/science.1217230","article-title":"Awake hippocampal sharp-wave ripples support spatial memory","volume":"336","author":"Jadhav","year":"2012","journal-title":"Science"},{"key":"2026070904085472300_R64","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.66227","article-title":"Multiple time-scales of decision-making in the hippocampus and prefrontal cortex","volume":"10","author":"Tang","year":"2021","journal-title":"eLife"},{"key":"2026070904085472300_R65","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1146\/annurev-neuro-111020-084824","article-title":"Multiple-timescale representations of space: linking memory to navigation","volume":"45","author":"Tang","year":"2022","journal-title":"Annu. Rev. Neurosci."},{"key":"2026070904085472300_R66","doi-asserted-by":"crossref","first-page":"1172","DOI":"10.1016\/j.neunet.2005.08.007","article-title":"Hippocampal mechanisms for the context-dependent retrieval of episodes","volume":"18","author":"Hasselmo","year":"2005","journal-title":"Neural Netw."},{"key":"2026070904085472300_R67","doi-asserted-by":"crossref","first-page":"317","DOI":"10.1002\/hipo.450030307","article-title":"Phase relationship between hippocampal place units and the EEG theta rhythm","volume":"3","author":"O\u2019Keefe","year":"1993","journal-title":"Hippocampus"},{"key":"2026070904085472300_R68","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1002\/(SICI)1098-1063(1996)6:2<149::AID-HIPO6>3.0.CO;2-K","article-title":"Theta phase precession in hippocampal neuronal populations and the compression of temporal sequences","volume":"6","author":"Skaggs","year":"1996","journal-title":"Hippocampus"},{"key":"2026070904085472300_R69","doi-asserted-by":"crossref","first-page":"2166","DOI":"10.1097\/00001756-199511000-00017","article-title":"Medial septal control of theta-correlated unit firing in the entorhinal cortex of awake rats","volume":"6","author":"Jeffery","year":"1995","journal-title":"Neuroreport"},{"key":"2026070904085472300_R70","doi-asserted-by":"crossref","first-page":"994","DOI":"10.1152\/jn.01141.2009","article-title":"Theta modulation in the medial and the lateral entorhinal cortices","volume":"104","author":"Deshmukh","year":"2010","journal-title":"J. Neurophysiol."},{"key":"2026070904085472300_R71","doi-asserted-by":"crossref","first-page":"1248","DOI":"10.1038\/nature06957","article-title":"Hippocampus-independent phase precession in entorhinal grid cells","volume":"453","author":"Hafting","year":"2008","journal-title":"Nature"},{"key":"2026070904085472300_R72","doi-asserted-by":"crossref","first-page":"2526","DOI":"10.1111\/ejn.12256","article-title":"Phase coding by grid cells in unconstrained environments: two-dimensional phase precession","volume":"38","author":"Climer","year":"2013","journal-title":"Eur. J. Neurosci."},{"key":"2026070904085472300_R73","doi-asserted-by":"crossref","first-page":"4101","DOI":"10.1523\/JNEUROSCI.09-12-04101.1989","article-title":"The firing of hippocampal place cells predicts the future position of freely moving rats","volume":"9","author":"Muller","year":"1989","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R74","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1038\/s41586-023-05928-6","article-title":"Dynamic synchronization between hippocampal representations and stepping","volume":"617","author":"Joshi","year":"2023","journal-title":"Nature"},{"key":"2026070904085472300_R75","doi-asserted-by":"crossref","first-page":"2106","DOI":"10.1038\/s41593-025-02038-6","article-title":"Allothetic and idiothetic spatial cues control the multiplexed theta phase coding of place cells","volume":"28","author":"Sueoka","year":"2025","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R76","doi-asserted-by":"crossref","first-page":"793","DOI":"10.1162\/089976602317318965","article-title":"A proposed function for hippocampal theta rhythm: separate phases of encoding and retrieval enhance reversal of prior learning","volume":"14","author":"Hasselmo","year":"2002","journal-title":"Neural Comput."},{"key":"2026070904085472300_R77","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1038\/nature12112","article-title":"Hippocampal place-cell sequences depict future paths to remembered goals","volume":"497","author":"Pfeiffer","year":"2013","journal-title":"Nature"},{"key":"2026070904085472300_R78","doi-asserted-by":"crossref","first-page":"975","DOI":"10.1016\/j.neuron.2016.10.028","article-title":"Interplay between hippocampal sharp-wave-ripple events and vicarious trial and error behaviors in decision making","volume":"92","author":"Papale","year":"2016","journal-title":"Neuron"},{"key":"2026070904085472300_R79","article-title":"Goal-directed hippocampal theta sweeps during memory-guided navigation","author":"Tang","year":"2025","journal-title":"bioRxiv"},{"key":"2026070904085472300_R80","doi-asserted-by":"crossref","first-page":"1209","DOI":"10.1002\/hipo.20359","article-title":"Network dynamics of hippocampal cell-assemblies resemble multiple spatial maps within single tasks","volume":"17","author":"Jackson","year":"2007","journal-title":"Hippocampus"},{"key":"2026070904085472300_R81","doi-asserted-by":"crossref","first-page":"246","DOI":"10.1038\/nature10439","article-title":"Theta-paced flickering between place-cell maps in the hippocampus","volume":"478","author":"Jezek","year":"2011","journal-title":"Nature"},{"key":"2026070904085472300_R82","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.nlm.2015.12.011","article-title":"Coordinating different representations in the hippocampus","volume":"129","author":"Kelemen","year":"2016","journal-title":"Neurobiol. Learn. Mem."},{"key":"2026070904085472300_R83","doi-asserted-by":"crossref","first-page":"1504","DOI":"10.1038\/nn.4656","article-title":"The cognitive map in humans: spatial navigation and beyond","volume":"20","author":"Epstein","year":"2017","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R84","doi-asserted-by":"crossref","first-page":"520","DOI":"10.1016\/j.tics.2021.02.010","article-title":"The versatile wayfinder: prefrontal contributions to spatial navigation","volume":"25","author":"Patai","year":"2021","journal-title":"Trends Cogn. Sci."},{"key":"2026070904085472300_R85","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1038\/nature08704","article-title":"Evidence for grid cells in a human memory network","volume":"463","author":"Doeller","year":"2010","journal-title":"Nature"},{"key":"2026070904085472300_R86","doi-asserted-by":"crossref","first-page":"1188","DOI":"10.1038\/nn.3466","article-title":"Direct recordings of grid-like neuronal activity in human spatial navigation","volume":"16","author":"Jacobs","year":"2013","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R87","doi-asserted-by":"crossref","first-page":"1194","DOI":"10.1002\/hipo.23250","article-title":"Spatial encoding in dorsomedial prefrontal cortex and hippocampus is related during deliberation","volume":"30","author":"Hasz","year":"2020","journal-title":"Hippocampus"},{"key":"2026070904085472300_R88","doi-asserted-by":"crossref","DOI":"10.1002\/hipo.70008","article-title":"Phase precession relative to turning angle in theta-modulated head direction cells","volume":"35","author":"Ji","year":"2025","journal-title":"Hippocampus"},{"key":"2026070904085472300_R89","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1002\/(SICI)1098-1063(1998)8:2<87::AID-HIPO1>3.0.CO;2-4","article-title":"Comparisons of head direction cell activity in the postsubiculum and anterior thalamus of freely moving rats","volume":"8","author":"Taube","year":"1998","journal-title":"Hippocampus"},{"key":"2026070904085472300_R90","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1126\/science.1201652","article-title":"Reduction of theta rhythm dissociates grid cell spatial periodicity from directional tuning","volume":"332","author":"Brandon","year":"2011","journal-title":"Science"},{"key":"2026070904085472300_R91","doi-asserted-by":"crossref","first-page":"592","DOI":"10.1126\/science.1201685","article-title":"The spatial periodicity of grid cells is not sustained during reduced theta oscillations","volume":"332","author":"Koenig","year":"2011","journal-title":"Science"},{"key":"2026070904085472300_R92","doi-asserted-by":"crossref","first-page":"789","DOI":"10.1016\/j.neuron.2014.04.013","article-title":"New and distinct hippocampal place codes are generated in a new environment during septal inactivation","volume":"82","author":"Brandon","year":"2014","journal-title":"Neuron"},{"key":"2026070904085472300_R93","doi-asserted-by":"crossref","first-page":"1179","DOI":"10.1016\/j.cub.2018.02.061","article-title":"Hippocampal neural circuits respond to optogenetic pacing of theta frequencies by generating accelerated oscillation frequencies","volume":"28","author":"Zutshi","year":"2018","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R94","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2024.114590","article-title":"Optogenetic silencing of medial septal GABAergic neurons disrupts grid cell spatial and temporal coding in the medial entorhinal cortex","volume":"43","author":"Robinson","year":"2024","journal-title":"Cell Rep."},{"key":"2026070904085472300_R95","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/0006-8993(90)91465-S","article-title":"Muscimol injections in the medial septum impair spatial learning","volume":"522","author":"Brioni","year":"1990","journal-title":"Brain Res."},{"key":"2026070904085472300_R96","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1016\/0006-8993(95)01323-7","article-title":"Effects of reversible inactivations of the medial septal area on reference and working memory versions of the Morris water maze","volume":"709","author":"Rashidy-Pour","year":"1996","journal-title":"Brain Res."},{"key":"2026070904085472300_R97","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1037\/0735-7044.104.6.849","article-title":"Cholinergic and GABAergic modulation of medial septal area: effect on working memory","volume":"104","author":"Givens","year":"1990","journal-title":"Behav. Neurosci."},{"key":"2026070904085472300_R98","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1006\/nlme.1997.3769","article-title":"Intraseptal infusions of muscimol impair spontaneous alternation performance: infusions of glucose into the hippocampus, but not the medial septum, reverse the deficit","volume":"68","author":"Parent","year":"1997","journal-title":"Neurobiol. Learn. Mem."},{"key":"2026070904085472300_R99","doi-asserted-by":"crossref","DOI":"10.1016\/j.celrep.2021.108801","article-title":"Two septal-entorhinal GABAergic projections differentially control coding properties of spatially tuned neurons in the medial entorhinal cortex","volume":"34","author":"Schlesiger","year":"2021","journal-title":"Cell Rep."},{"key":"2026070904085472300_R100","article-title":"Medial septum-dependent encoding of contextual inputs by hippocampal splitter cells","volume":"44","author":"Li","year":"2025","journal-title":"Cell Rep."},{"key":"2026070904085472300_R101","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1037\/bne0000415","article-title":"Disruption of the anterior thalamic head direction signal following reduction of the hippocampal theta rhythm","volume":"134","author":"Pastor","year":"2020","journal-title":"Behav. Neurosci."},{"key":"2026070904085472300_R102","doi-asserted-by":"crossref","first-page":"2727","DOI":"10.1007\/s00429-017-1368-4","article-title":"Medial entorhinal cortex and medial septum contribute to self-motion-based linear distance estimation","volume":"222","author":"Jacob","year":"2017","journal-title":"Brain Struct. Funct."},{"key":"2026070904085472300_R103","doi-asserted-by":"crossref","first-page":"4199","DOI":"10.1016\/j.cub.2025.07.050","article-title":"An independent coding scheme for idiothetic distance in the hippocampus","volume":"35","author":"Nordlund","year":"2025","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R104","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1016\/j.nlm.2009.07.005","article-title":"A model of episodic memory: mental time travel along encoded trajectories using grid cells","volume":"92","author":"Hasselmo","year":"2009","journal-title":"Neurobiol. Learn. Mem."},{"key":"2026070904085472300_R105","volume-title":"How we remember: brain mechanisms of episodic memory","author":"Hasselmo","year":"2012"},{"key":"2026070904085472300_R106","doi-asserted-by":"crossref","first-page":"739","DOI":"10.1038\/s41586-024-08392-y","article-title":"Episodic and associative memory from spatial scaffolds in the hippocampus","volume":"638","author":"Chandra","year":"2025","journal-title":"Nature"},{"key":"2026070904085472300_R107","doi-asserted-by":"crossref","first-page":"1547","DOI":"10.1016\/j.neuron.2022.02.002","article-title":"Flexible rerouting of hippocampal replay sequences around changing barriers in the absence of global place field remapping","volume":"110","author":"Widloski","year":"2022","journal-title":"Neuron"},{"key":"2026070904085472300_R108","doi-asserted-by":"crossref","first-page":"541","DOI":"10.1126\/science.ads4760","article-title":"The time course and organization of hippocampal replay","volume":"387","author":"Mallory","year":"2025","journal-title":"Science"},{"key":"2026070904085472300_R109","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1515\/REVNEURO.2006.17.1-2.71","article-title":"The boundary vector cell model of place cell firing and spatial memory","volume":"17","author":"Barry","year":"2006","journal-title":"Rev. Neurosci."},{"key":"2026070904085472300_R110","doi-asserted-by":"crossref","first-page":"1865","DOI":"10.1126\/science.1166466","article-title":"Representation of geometric borders in the entorhinal cortex","volume":"322","author":"Solstad","year":"2008","journal-title":"Science"},{"key":"2026070904085472300_R111","doi-asserted-by":"crossref","first-page":"2045","DOI":"10.1002\/hipo.22046","article-title":"Perirhinal cortex represents nonspatial, but not spatial, information in rats foraging in the presence of objects: comparison with lateral entorhinal cortex","volume":"22","author":"Deshmukh","year":"2012","journal-title":"Hippocampus"},{"key":"2026070904085472300_R112","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1038\/s41586-019-1077-7","article-title":"Object-vector coding in the medial entorhinal cortex","volume":"568","author":"H\u00f8ydal","year":"2019","journal-title":"Nature"},{"key":"2026070904085472300_R113","doi-asserted-by":"crossref","first-page":"8025","DOI":"10.1038\/s41467-024-52315-4","article-title":"Distinct codes for environment structure and symmetry in postrhinal and retrosplenial cortices","volume":"15","author":"LaChance","year":"2024","journal-title":"Nat. Commun."},{"key":"2026070904085472300_R114","doi-asserted-by":"crossref","first-page":"1493","DOI":"10.1098\/rstb.2001.0948","article-title":"Memory for events and their spatial context: models and experiments","volume":"356","author":"Burgess","year":"2001","journal-title":"Phil. Trans. R. Soc. Lond. B"},{"key":"2026070904085472300_R115","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1016\/S0893-6080(03)00106-0","article-title":"Modeling goal-directed spatial navigation in the rat based on physiological data from the hippocampal formation","volume":"16","author":"Koene","year":"2003","journal-title":"Neural Netw."},{"key":"2026070904085472300_R116","doi-asserted-by":"crossref","first-page":"7324","DOI":"10.1523\/JNEUROSCI.1909-08.2008","article-title":"Different CA1 and CA3 representations of novel routes in a shortcut situation","volume":"28","author":"Alvernhe","year":"2008","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R117","doi-asserted-by":"crossref","first-page":"1696","DOI":"10.1111\/j.1460-9568.2011.07653.x","article-title":"Local remapping of place cell firing in the Tolman detour task","volume":"33","author":"Alvernhe","year":"2011","journal-title":"Eur. J. Neurosci."},{"key":"2026070904085472300_R118","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1007\/s10071-011-0460-z","article-title":"Rats build and update topological representations through exploration","volume":"15","author":"Alvernhe","year":"2011","journal-title":"Anim. Cogn."},{"key":"2026070904085472300_R119","doi-asserted-by":"crossref","first-page":"695","DOI":"10.1016\/j.neuron.2010.01.034","article-title":"Hippocampal replay is not a simple function of experience","volume":"65","author":"Gupta","year":"2010","journal-title":"Neuron"},{"key":"2026070904085472300_R120","doi-asserted-by":"crossref","first-page":"1331","DOI":"10.1016\/j.cub.2014.05.001","article-title":"The hippocampus and entorhinal cortex encode the path and Euclidean distances to goals during navigation","volume":"24","author":"Howard","year":"2014","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R121","doi-asserted-by":"crossref","first-page":"15442","DOI":"10.1523\/JNEUROSCI.1209-15.2015","article-title":"There and back again: hippocampus and retrosplenial cortex track homing distance during human path integration","volume":"35","author":"Chrastil","year":"2015","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R122","article-title":"How does the response of egocentric boundary cells depend upon the coordinate system of environmental features?","author":"Robinson","year":"2025","journal-title":"bioRxiv"},{"key":"2026070904085472300_R123","doi-asserted-by":"crossref","first-page":"1728","DOI":"10.1016\/j.cub.2023.03.066","article-title":"Geometric determinants of the postrhinal egocentric spatial map","volume":"33","author":"LaChance","year":"2023","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R124","doi-asserted-by":"crossref","first-page":"749","DOI":"10.1002\/(SICI)1098-1063(1996)6:6<749::AID-HIPO16>3.0.CO;2-0","article-title":"Neuronal computations underlying the firing of place cells and their role in navigation","volume":"6","author":"Burgess","year":"1996","journal-title":"Hippocampus"},{"key":"2026070904085472300_R125","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1002\/1098-1063(2000)10:4<369::AID-HIPO3>3.0.CO;2-0","article-title":"Modeling place fields in terms of the cortical inputs to the hippocampus","volume":"10","author":"Hartley","year":"2000","journal-title":"Hippocampus"},{"key":"2026070904085472300_R126","doi-asserted-by":"crossref","first-page":"150","DOI":"10.1016\/j.neuron.2022.11.006","article-title":"Rethinking retrosplenial cortex: perspectives and predictions","volume":"111","author":"Alexander","year":"2023","journal-title":"Neuron"},{"key":"2026070904085472300_R127","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1007\/BF00243212","article-title":"Head-direction cells in the rat posterior cortex","volume":"101","author":"Chen","year":"1994","journal-title":"Exp. Brain Res."},{"key":"2026070904085472300_R128","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1038\/nn.4465","article-title":"An independent, landmark-dominated head-direction signal in dysgranular retrosplenial cortex","volume":"20","author":"Jacob","year":"2017","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R129","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1037\/0735-7044.115.1.3","article-title":"Head direction, place, and movement correlates for cells in the rat retrosplenial cortex","volume":"115","author":"Cho","year":"2001","journal-title":"Behav. Neurosci."},{"key":"2026070904085472300_R130","doi-asserted-by":"crossref","first-page":"1121","DOI":"10.1002\/hipo.20958","article-title":"Complimentary roles of the hippocampus and retrosplenial cortex in behavioral context discrimination","volume":"22","author":"Smith","year":"2011","journal-title":"Hippocampus"},{"key":"2026070904085472300_R131","first-page":"3713","article-title":"Retrosplenial cortical neurons encode navigational cues, trajectories and reward locations during goal directed navigation","volume":"27","author":"Vedder","year":"2016","journal-title":"Cereb. Cortex"},{"key":"2026070904085472300_R132","doi-asserted-by":"crossref","first-page":"2083","DOI":"10.1016\/j.cub.2019.05.034","article-title":"Retrosplenial cortical representations of space and future goal locations develop with learning","volume":"29","author":"Miller","year":"2019","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R133","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1038\/nn.4058","article-title":"Retrosplenial cortex maps the conjunction of internal and external spaces","volume":"18","author":"Alexander","year":"2015","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R134","doi-asserted-by":"crossref","first-page":"1551","DOI":"10.1016\/j.cub.2017.04.036","article-title":"Spatially periodic activation patterns of retrosplenial cortex encode route sub-spaces and distance traveled","volume":"27","author":"Alexander","year":"2017","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R135","doi-asserted-by":"crossref","first-page":"101","DOI":"10.1037\/bne0000500","article-title":"The anatomy and function of the postrhinal cortex","volume":"136","author":"Estela-Pro","year":"2022","journal-title":"Behav. Neurosci."},{"key":"2026070904085472300_R136","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.bbr.2013.07.005","article-title":"Hippocampal and subicular efferents and afferents of the perirhinal, postrhinal, and entorhinal cortices of the rat","volume":"254","author":"Agster","year":"2013","journal-title":"Behav. Brain Res."},{"key":"2026070904085472300_R137","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.59816","article-title":"Entorhinal-retrosplenial circuits for allocentric-egocentric transformation of boundary coding","volume":"9","author":"van Wijngaarden","year":"2020","journal-title":"eLife"},{"key":"2026070904085472300_R138","doi-asserted-by":"crossref","first-page":"2772","DOI":"10.1038\/s41467-019-10722-y","article-title":"Neuronal representation of environmental boundaries in egocentric coordinates","volume":"10","author":"Hinman","year":"2019","journal-title":"Nat. Commun."},{"key":"2026070904085472300_R139","doi-asserted-by":"crossref","first-page":"2751","DOI":"10.1016\/j.cub.2019.07.007","article-title":"Dissociation between postrhinal cortex and downstream parahippocampal regions in the representation of egocentric boundaries","volume":"29","author":"Gofman","year":"2019","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R140","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1126\/science.aau4940","article-title":"Egocentric coding of external items in the lateral entorhinal cortex","volume":"362","author":"Wang","year":"2018","journal-title":"Science"},{"key":"2026070904085472300_R141","doi-asserted-by":"crossref","first-page":"12","DOI":"10.1016\/j.conb.2019.11.005","article-title":"Egocentric and allocentric representations of space in the rodent brain","volume":"60","author":"Wang","year":"2020","journal-title":"Curr. Opin. Neurobiol."},{"key":"2026070904085472300_R142","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1002\/hipo.23513","article-title":"Gated transformations from egocentric to allocentric reference frames involving retrosplenial cortex, entorhinal cortex, and hippocampus","volume":"33","author":"Alexander","year":"2023","journal-title":"Hippocampus"},{"key":"2026070904085472300_R143","doi-asserted-by":"crossref","first-page":"5846","DOI":"10.1523\/JNEUROSCI.0835-08.2008","article-title":"The h channel mediates location dependence and plasticity of intrinsic phase response in rat hippocampal neurons","volume":"28","author":"Narayanan","year":"2008","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R144","doi-asserted-by":"crossref","first-page":"488","DOI":"10.1002\/hipo.23510","article-title":"A model for transforming egocentric views into goal-directed behavior","volume":"33","author":"LaChance","year":"2023","journal-title":"Hippocampus"},{"key":"2026070904085472300_R145","article-title":"Distinct cortical spatial representations learned along disparate visual pathways","author":"Lian","year":"2025","journal-title":"bioRxiv"},{"key":"2026070904085472300_R146","article-title":"Population tuning for egocentric information in human cortex.","author":"Dunne","year":"2026"},{"key":"2026070904085472300_R147","doi-asserted-by":"crossref","DOI":"10.1177\/2398212820972871","article-title":"Neurophysiological coding of space and time in the hippocampus, entorhinal cortex, and retrosplenial cortex","volume":"4","author":"Alexander","year":"2020","journal-title":"Brain Neurosci. Adv."},{"key":"2026070904085472300_R148","doi-asserted-by":"crossref","DOI":"10.1016\/j.isci.2021.103377","article-title":"Neural responses in retrosplenial cortex associated with environmental alterations","volume":"24","author":"Carstensen","year":"2021","journal-title":"iScience"},{"key":"2026070904085472300_R149","doi-asserted-by":"crossref","first-page":"2531","DOI":"10.1523\/JNEUROSCI.17-07-02531.1997","article-title":"Failure of centrally placed objects to control the firing fields of hippocampal place cells","volume":"17","author":"Cressant","year":"1997","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R150","doi-asserted-by":"crossref","first-page":"1088","DOI":"10.1016\/j.cub.2022.01.023","article-title":"Flexible encoding of objects and space in single cells of the dentate gyrus","volume":"32","author":"GoodSmith","year":"2022","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R151","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1016\/0301-0082(79)90005-4","article-title":"A review of the hippocampal place cells","volume":"13","author":"O\u2019Keefe","year":"1979","journal-title":"Prog. Neurobiol."},{"key":"2026070904085472300_R152","doi-asserted-by":"crossref","first-page":"5598","DOI":"10.1523\/JNEUROSCI.5265-11.2012","article-title":"Neural correlates of long-term object memory in the mouse anterior cingulate cortex","volume":"32","author":"Weible","year":"2012","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R153","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1016\/j.cub.2013.01.036","article-title":"Traces of experience in the lateral entorhinal cortex","volume":"23","author":"Tsao","year":"2013","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R154","doi-asserted-by":"crossref","first-page":"266","DOI":"10.1038\/s41593-020-00761-w","article-title":"Vector trace cells in the subiculum of the hippocampal formation","volume":"24","author":"Poulter","year":"2020","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R155","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1016\/S0306-4522(03)00160-X","article-title":"Positional firing properties of postrhinal cortex neurons","volume":"119","author":"Burwell","year":"2003","journal-title":"Neuroscience"},{"key":"2026070904085472300_R156","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.nlm.2006.05.007","article-title":"Hippocampal CA1 spiking during encoding and retrieval: relation to theta phase","volume":"87","author":"Manns","year":"2007","journal-title":"Neurobiol. Learn. Mem."},{"key":"2026070904085472300_R157","doi-asserted-by":"crossref","first-page":"976","DOI":"10.1016\/j.neuron.2012.10.039","article-title":"Single neuron activity and theta modulation in postrhinal cortex during visual object discrimination","volume":"76","author":"Furtak","year":"2012","journal-title":"Neuron"},{"key":"2026070904085472300_R158","doi-asserted-by":"crossref","first-page":"1978","DOI":"10.1016\/j.neuron.2022.03.026","article-title":"Signatures of rapid plasticity in hippocampal CA1 representations during novel experiences","volume":"110","author":"Priestley","year":"2022","journal-title":"Neuron"},{"key":"2026070904085472300_R159","first-page":"171","article-title":"Spatial responses to environmental manipulations in the retrosplenial cortex","volume":"49","author":"Malmberg","year":"2023","journal-title":"Soc. Neurosci. Abstr."},{"key":"2026070904085472300_R160","first-page":"190.","article-title":"A subset of egocentric boundary cells in the retrosplenial cortex show preferential tuning to circumventable barriers in an environment","volume":"50","author":"Malmberg","year":"2024","journal-title":"Soc. Neurosci. Abstr."},{"key":"2026070904085472300_R161","doi-asserted-by":"crossref","first-page":"264","DOI":"10.1038\/nn.3329","article-title":"Long-term dynamics of CA1 hippocampal place codes","volume":"16","author":"Ziv","year":"2013","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R162","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.12247","article-title":"Hippocampal ensemble dynamics timestamp events in long-term memory","volume":"4","author":"Rubin","year":"2015","journal-title":"eLife"},{"key":"2026070904085472300_R163","doi-asserted-by":"crossref","first-page":"2443","DOI":"10.1038\/s41467-020-16226-4","article-title":"Trajectory-modulated hippocampal neurons persist throughout memory-guided navigation","volume":"11","author":"Kinsky","year":"2020","journal-title":"Nat. Commun."},{"key":"2026070904085472300_R164","doi-asserted-by":"crossref","first-page":"244","DOI":"10.1002\/hipo.23272","article-title":"Hippocampal spatial memory representations in mice are heterogeneously stable","volume":"31","author":"Levy","year":"2020","journal-title":"Hippocampus"},{"key":"2026070904085472300_R165","volume-title":"The neuropsychology of anxiety: an enquiry into the functions of the septo-hippocampal system","author":"Gray","year":"1982"},{"key":"2026070904085472300_R166","first-page":"187","volume-title":"Neuroscience and connectionist theory","author":"Levy","year":"1990"},{"key":"2026070904085472300_R167","doi-asserted-by":"crossref","first-page":"8027","DOI":"10.1523\/JNEUROSCI.16-24-08027.1996","article-title":"Dynamics of mismatch correction in the hippocampal ensemble code for space: interaction between path integration and environmental cues","volume":"16","author":"Gothard","year":"1996","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R168","doi-asserted-by":"crossref","first-page":"8660","DOI":"10.1073\/pnas.93.16.8660","article-title":"The hippocampal formation participates in novel picture encoding: evidence from functional magnetic resonance imaging","volume":"93","author":"Stern","year":"1996","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026070904085472300_R169","doi-asserted-by":"crossref","first-page":"337","DOI":"10.1002\/hipo.1048","article-title":"Medial temporal and prefrontal contributions to working memory tasks with novel and familiar stimuli","volume":"11","author":"Stern","year":"2001","journal-title":"Hippocampus"},{"key":"2026070904085472300_R170","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1177\/0269881115612238","article-title":"Neurotransmitters and novelty: a systematic review","volume":"30","author":"Rangel-Gomez","year":"2015","journal-title":"J. Psychopharmacol."},{"key":"2026070904085472300_R171","doi-asserted-by":"crossref","first-page":"1377","DOI":"10.1126\/science.1962197","article-title":"A neural mechanism for working and recognition memory in inferior temporal cortex","volume":"254","author":"Miller","year":"1991","journal-title":"Science"},{"key":"2026070904085472300_R172","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1007\/BF00234113","article-title":"Neuronal activity related to visual recognition memory: long-term memory and the encoding of recency and familiarity information in the primate anterior and medial inferior temporal and rhinal cortex","volume":"96","author":"Fahy","year":"1993","journal-title":"Exp. Brain Res."},{"key":"2026070904085472300_R173","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1038\/nrn1052","article-title":"Neural mechanisms for detecting and remembering novel events","volume":"4","author":"Ranganath","year":"2003","journal-title":"Nat. Rev. Neurosci."},{"key":"2026070904085472300_R174","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1038\/s41583-023-00784-9","article-title":"Curiosity: primate neural circuits for novelty and information seeking","volume":"25","author":"Monosov","year":"2024","journal-title":"Nat. Rev. Neurosci."},{"key":"2026070904085472300_R175","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1016\/j.bbr.2005.09.005","article-title":"The role of the hippocampus in object recognition in rats: examination of the influence of task parameters and lesion size","volume":"167","author":"Ainge","year":"2006","journal-title":"Behav. Brain Res."},{"key":"2026070904085472300_R176","doi-asserted-by":"crossref","first-page":"21","DOI":"10.1016\/j.brainresbull.2011.09.008","article-title":"Cholinergic mechanisms of episodic memory: what specific behavioural tasks can tell us about specific neural mechanisms","volume":"92","author":"Easton","year":"2013","journal-title":"Brain Res. Bull."},{"key":"2026070904085472300_R177","doi-asserted-by":"crossref","first-page":"352","DOI":"10.1002\/hipo.22095","article-title":"Lateral entorhinal cortex is critical for novel object-context recognition","volume":"23","author":"Wilson","year":"2013","journal-title":"Hippocampus"},{"key":"2026070904085472300_R178","doi-asserted-by":"crossref","DOI":"10.1016\/j.nlm.2021.107544","article-title":"Functional connectivity of anterior retrosplenial cortex in object recognition memory","volume":"186","author":"de Landeta","year":"2021","journal-title":"Neurobiol. Learn. Mem."},{"key":"2026070904085472300_R179","doi-asserted-by":"crossref","first-page":"6173","DOI":"10.1523\/JNEUROSCI.20-16-06173.2000","article-title":"Prefrontal-temporal circuitry for episodic encoding and subsequent memory","volume":"20","author":"Kirchhoff","year":"2000","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R180","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1126\/science.1221936","article-title":"Memory\u2019s penumbra: episodic memory decisions induce lingering mnemonic biases","volume":"337","author":"Duncan","year":"2012","journal-title":"Science"},{"key":"2026070904085472300_R181","doi-asserted-by":"crossref","first-page":"3898","DOI":"10.1523\/JNEUROSCI.14-06-03898.1994","article-title":"Laminar selectivity of the cholinergic suppression of synaptic transmission in rat hippocampal region CA1: computational modeling and brain slice physiology","volume":"14","author":"Hasselmo","year":"1994","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R182","doi-asserted-by":"crossref","first-page":"5249","DOI":"10.1523\/JNEUROSCI.15-07-05249.1995","article-title":"Dynamics of learning and recall at excitatory recurrent synapses and cholinergic modulation in rat hippocampal region CA3","volume":"15","author":"Hasselmo","year":"1995","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R183","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0166-4328(97)00048-X","article-title":"Free recall and recognition in a network model of the hippocampus: simulating effects of scopolamine on human memory function","volume":"89","author":"Hasselmo","year":"1997","journal-title":"Behav. Brain Res."},{"key":"2026070904085472300_R184","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1037\/0033-295X.110.4.611","article-title":"Modeling hippocampal and neocortical contributions to recognition memory: a complementary-learning-systems approach","volume":"110","author":"Norman","year":"2003","journal-title":"Psychol. Rev."},{"key":"2026070904085472300_R185","doi-asserted-by":"crossref","DOI":"10.1016\/j.nlm.2023.107826","article-title":"Predictive learning by a burst-dependent learning rule","volume":"205","author":"Chapman","year":"2023","journal-title":"Neurobiol. Learn. Mem."},{"key":"2026070904085472300_R186","doi-asserted-by":"crossref","first-page":"3088","DOI":"10.1073\/pnas.81.10.3088","article-title":"Neurons with graded response have collective computational properties like those of two-state neurons","volume":"81","author":"Hopfield","year":"1984","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2026070904085472300_R187","doi-asserted-by":"crossref","first-page":"710","DOI":"10.1016\/j.conb.2006.09.002","article-title":"The role of acetylcholine in learning and memory","volume":"16","author":"Hasselmo","year":"2006","journal-title":"Curr. Opin. Neurobiol."},{"key":"2026070904085472300_R188","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1038\/nn.3904","article-title":"Theta sequences are essential for internally generated hippocampal firing fields","volume":"18","author":"Wang","year":"2014","journal-title":"Nat. Neurosci."},{"key":"2026070904085472300_R189","doi-asserted-by":"crossref","first-page":"3175","DOI":"10.1523\/JNEUROSCI.21-09-03175.2001","article-title":"Gating of human theta oscillations by a working memory task","volume":"21","author":"Raghavachari","year":"2001","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R190","doi-asserted-by":"crossref","first-page":"68","DOI":"10.1016\/j.neuroimage.2012.07.064","article-title":"Expected reward modulates encoding-related theta activity before an event","volume":"64","author":"Gruber","year":"2013","journal-title":"Neuroimage"},{"key":"2026070904085472300_R191","doi-asserted-by":"crossref","DOI":"10.1016\/j.neuroimage.2020.117558","article-title":"Theta rhythm supports hippocampus-dependent integrative encoding in schematic\/semantic memory networks","volume":"226","author":"Nicol\u00e1s","year":"2021","journal-title":"Neuroimage"},{"key":"2026070904085472300_R192","doi-asserted-by":"crossref","DOI":"10.1016\/j.neuroimage.2022.119581","article-title":"Theta oscillations support active exploration in human spatial navigation","volume":"262","author":"Chrastil","year":"2022","journal-title":"Neuroimage"},{"key":"2026070904085472300_R193","doi-asserted-by":"crossref","first-page":"1705","DOI":"10.1523\/JNEUROSCI.02-12-01705.1982","article-title":"Temporal relationship between sniffing and the limbic theta rhythm during odor discrimination reversal learning","volume":"2","author":"Macrides","year":"1982","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R194","doi-asserted-by":"crossref","first-page":"7048","DOI":"10.1038\/s41467-021-27323-3","article-title":"Theta rhythmicity governs human behavior and hippocampal signals during memory-dependent tasks","volume":"12","author":"ter Wal","year":"2021","journal-title":"Nat. Commun."},{"key":"2026070904085472300_R195","doi-asserted-by":"crossref","first-page":"1352","DOI":"10.1016\/j.neuroimage.2006.01.009","article-title":"Human neocortical oscillations exhibit theta phase differences between encoding and retrieval","volume":"31","author":"Rizzuto","year":"2006","journal-title":"Neuroimage"},{"key":"2026070904085472300_R196","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1097\/00001756-199612200-00032","article-title":"Stimulus-evoked resetting of the dentate theta rhythm","volume":"8","author":"Givens","year":"1996","journal-title":"Neuroreport"},{"key":"2026070904085472300_R197","doi-asserted-by":"crossref","first-page":"3383","DOI":"10.1016\/j.cub.2018.08.065","article-title":"An optimal oscillatory phase for pattern reactivation during memory retrieval","volume":"28","author":"Kerr\u00e9n","year":"2018","journal-title":"Curr. Biol."},{"key":"2026070904085472300_R198","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1002\/hipo.20671","article-title":"Environmental novelty elicits a later theta phase of firing in CA1 but not subiculum","volume":"20","author":"Lever","year":"2009","journal-title":"Hippocampus"},{"key":"2026070904085472300_R199","doi-asserted-by":"crossref","first-page":"8689","DOI":"10.1523\/JNEUROSCI.4483-12.2013","article-title":"Evidence for encoding versus retrieval scheduling in the hippocampus by theta phase and acetylcholine","volume":"33","author":"Douchamps","year":"2013","journal-title":"J. Neurosci."},{"key":"2026070904085472300_R200","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1038\/nature08573","article-title":"Frequency of gamma oscillations routes flow of information in the hippocampus","volume":"462","author":"Colgin","year":"2009","journal-title":"Nature"},{"key":"2026070904085472300_R201","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.conb.2014.08.001","article-title":"Theta\u2013gamma coupling in the entorhinal\u2013hippocampal system","volume":"31","author":"Colgin","year":"2015","journal-title":"Curr. Opin. Neurobiol."},{"key":"2026070904085472300_R202","doi-asserted-by":"crossref","first-page":"476","DOI":"10.1152\/jn.00472.2017","article-title":"Experience-dependent trends in CA1 theta and slow gamma rhythms in freely behaving mice","volume":"119","author":"Gereke","year":"2018","journal-title":"J. Neurophysiol."},{"key":"2026070904085472300_R203","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.03061","article-title":"Enhancement of encoding and retrieval functions through theta phase-specific manipulation of hippocampus","volume":"3","author":"Siegle","year":"2014","journal-title":"eLife"}],"container-title":["Philosophical Transactions B"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/royalsocietypublishing.org\/rstb\/article-pdf\/doi\/10.1098\/rstb.2025.0251\/6151443\/rstb.2025.0251.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/royalsocietypublishing.org\/rstb\/article-pdf\/doi\/10.1098\/rstb.2025.0251\/6151443\/rstb.2025.0251.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,9]],"date-time":"2026-07-09T08:09:37Z","timestamp":1783584577000},"score":1,"resource":{"primary":{"URL":"https:\/\/royalsocietypublishing.org\/rstb\/article\/381\/1954\/20250251\/482496\/Trajectory-scanning-as-a-predictive-coding"}},"subtitle":[],"editor":[{"given":"Mariam","family":"Aly","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,7,9]]},"references-count":203,"journal-issue":{"issue":"1954","published-online":{"date-parts":[[2026,7,9]]},"published-print":{"date-parts":[[2026,7,9]]}},"URL":"https:\/\/doi.org\/10.1098\/rstb.2025.0251","relation":{},"ISSN":["0962-8436","1471-2970"],"issn-type":[{"value":"0962-8436","type":"print"},{"value":"1471-2970","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,9]]},"article-number":"20250251"}}