{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,25]],"date-time":"2026-03-25T13:39:56Z","timestamp":1774445996988,"version":"3.50.1"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1010320","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,11,10]],"date-time":"2022-11-10T00:00:00Z","timestamp":1668038400000}}],"reference-count":74,"publisher":"Public Library of Science (PLoS)","issue":"10","license":[{"start":{"date-parts":[[2022,10,31]],"date-time":"2022-10-31T00:00:00Z","timestamp":1667174400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"crossref","award":["31680338"],"award-info":[{"award-number":["31680338"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>In general, strategies for spatial navigation could employ one of two spatial reference frames: egocentric or allocentric. Notwithstanding intuitive explanations, it remains unclear however under what circumstances one strategy is chosen over another, and how neural representations should be related to the chosen strategy. Here, we first use a deep reinforcement learning model to investigate whether a particular type of navigation strategy arises spontaneously during spatial learning without imposing a bias onto the model. We then examine the spatial representations that emerge in the network to support navigation. To this end, we study two tasks that are ethologically valid for mammals\u2014guidance, where the agent has to navigate to a goal location fixed in allocentric space, and aiming, where the agent navigates to a visible cue. We find that when both navigation strategies are available to the agent, the solutions it develops for guidance and aiming are heavily biased towards the allocentric or the egocentric strategy, respectively, as one might expect. Nevertheless, the agent can learn both tasks using either type of strategy. Furthermore, we find that place-cell-like allocentric representations emerge preferentially in guidance when using an allocentric strategy, whereas egocentric vector representations emerge when using an egocentric strategy in aiming. We thus find that alongside the type of navigational strategy, the nature of the task plays a pivotal role in the type of spatial representations that emerge.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1010320","type":"journal-article","created":{"date-parts":[[2022,10,31]],"date-time":"2022-10-31T13:55:44Z","timestamp":1667224544000},"page":"e1010320","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":21,"title":["Navigation task and action space drive the emergence of egocentric and allocentric spatial representations"],"prefix":"10.1371","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9011-2721","authenticated-orcid":true,"given":"Sandhiya","family":"Vijayabaskaran","sequence":"first","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6719-8029","authenticated-orcid":true,"given":"Sen","family":"Cheng","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2022,10,31]]},"reference":[{"key":"pcbi.1010320.ref001","first-page":"1","volume-title":"Spatial Cognition","author":"RL Klatzky","year":"1998"},{"key":"pcbi.1010320.ref002","doi-asserted-by":"crossref","first-page":"605","DOI":"10.1016\/j.neubiorev.2017.07.012","article-title":"Egocentric and Allocentric Spatial Reference Frames in Aging: A Systematic Review","volume":"80","author":"D Colombo","year":"2017","journal-title":"Neuroscience & Biobehavioral Reviews"},{"issue":"5","key":"pcbi.1010320.ref003","doi-asserted-by":"crossref","first-page":"761","DOI":"10.1037\/neu0000084","article-title":"Allocentric but Not Egocentric Orientation Is Impaired during Normal Aging: An ERP Study","volume":"28","author":"S Lithfous","year":"2014","journal-title":"Neuropsychology"},{"key":"pcbi.1010320.ref004","doi-asserted-by":"crossref","first-page":"29","DOI":"10.3389\/fnagi.2012.00029","article-title":"Aging Specifically Impairs Switching to an Allocentric Navigational Strategy","volume":"4","author":"MA Harris","year":"2012","journal-title":"Frontiers in Aging Neuroscience"},{"issue":"1","key":"pcbi.1010320.ref005","doi-asserted-by":"crossref","first-page":"202.e15","DOI":"10.1016\/j.neurobiolaging.2010.07.021","article-title":"Effects of Age on Navigation Strategy","volume":"33","author":"MK Rodgers","year":"2012","journal-title":"Neurobiology of Aging"},{"issue":"1-2","key":"pcbi.1010320.ref006","doi-asserted-by":"crossref","first-page":"77","DOI":"10.1016\/S0166-4328(98)00115-6","article-title":"Disrupted Allocentric but Preserved Egocentric Spatial Learning in Transgenic Mice with Impaired Glucocorticoid Receptor Function","volume":"100","author":"T Steckler","year":"1999","journal-title":"Behavioural Brain Research"},{"issue":"2","key":"pcbi.1010320.ref007","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1006\/nlme.2001.4008","article-title":"Multiple Parallel Memory Systems in the Brain of the Rat","volume":"77","author":"NM White","year":"2002","journal-title":"Neurobiology of Learning and Memory"},{"issue":"1","key":"pcbi.1010320.ref008","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1037\/h0081022","article-title":"Spatial Memory Deficit in Senescent Rats","volume":"34","author":"CA Barnes","year":"1980","journal-title":"Canadian Journal of Psychology\/Revue canadienne de psychologie"},{"issue":"5868","key":"pcbi.1010320.ref009","doi-asserted-by":"crossref","first-page":"681","DOI":"10.1038\/297681a0","article-title":"Place Navigation Impaired in Rats with Hippocampal Lesions","volume":"297","author":"RGM Morris","year":"1982","journal-title":"Nature"},{"issue":"2","key":"pcbi.1010320.ref010","doi-asserted-by":"crossref","first-page":"310","DOI":"10.1093\/ilar\/ilu013","article-title":"Assessing Spatial Learning and Memory in Rodents","volume":"55","author":"CV Vorhees","year":"2014","journal-title":"ILAR journal"},{"issue":"4","key":"pcbi.1010320.ref011","doi-asserted-by":"crossref","first-page":"462","DOI":"10.1016\/j.nlm.2007.08.013","article-title":"Emergence of an Egocentric Cue Guiding and Allocentric Inferring Strategy That Mirrors Hippocampal Brain-Derived Neurotrophic Factor (BDNF) Expression in the Morris Water Maze","volume":"89","author":"DR Harvey","year":"2008","journal-title":"Neurobiology of Learning and Memory"},{"issue":"2","key":"pcbi.1010320.ref012","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1016\/S0304-3940(03)00473-7","article-title":"Both Dorsal and Ventral Hippocampus Contribute to Spatial Learning in Long-Evans Rats","volume":"345","author":"J Ferbinteanu","year":"2003","journal-title":"Neuroscience Letters"},{"key":"pcbi.1010320.ref013","article-title":"Spatial Learning Impairment Parallels the Magnitude of Dorsal Hippocampal Lesions, but Is Hardly Present Following Ventral Lesions","author":"E Moser","year":"1993","journal-title":"The Journal of Neuroscience: The Official Journal of the Society for Neuroscience"},{"issue":"6","key":"pcbi.1010320.ref014","doi-asserted-by":"crossref","first-page":"e67232","DOI":"10.1371\/journal.pone.0067232","article-title":"Complementary Roles of the Hippocampus and the Dorsomedial Striatum during Spatial and Sequence-Based Navigation Behavior","volume":"8","author":"C Fouquet","year":"2013","journal-title":"PLOS ONE"},{"issue":"3","key":"pcbi.1010320.ref015","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1101\/lm.037077.114","article-title":"Dorsolateral Striatal Lesions Impair Navigation Based on Landmark-Goal Vectors but Facilitate Spatial Learning Based on a \u201cCognitive Map\u201d","volume":"22","author":"Y Kosaki","year":"2015","journal-title":"Learning & Memory"},{"key":"pcbi.1010320.ref016","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/j.nlm.2016.06.002","article-title":"Do the Anterior and Lateral Thalamic Nuclei Make Distinct Contributions to Spatial Representation and Memory?","volume":"133","author":"BJ Clark","year":"2016","journal-title":"Neurobiology of Learning and Memory"},{"issue":"1","key":"pcbi.1010320.ref017","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":"J O\u2019Keefe","year":"1971","journal-title":"Brain Research"},{"issue":"16","key":"pcbi.1010320.ref018","doi-asserted-by":"crossref","first-page":"5431","DOI":"10.1523\/JNEUROSCI.0511-14.2014","article-title":"Interaction of Egocentric and World-Centered Reference Frames in the Rat Posterior Parietal Cortex","volume":"34","author":"AA Wilber","year":"2014","journal-title":"The Journal of Neuroscience: The Official Journal of the Society for Neuroscience"},{"issue":"7052","key":"pcbi.1010320.ref019","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":"T Hafting","year":"2005","journal-title":"Nature"},{"issue":"5909","key":"pcbi.1010320.ref020","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":"T Solstad","year":"2008","journal-title":"Science"},{"issue":"6417","key":"pcbi.1010320.ref021","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":"C Wang","year":"2018","journal-title":"Science"},{"key":"pcbi.1010320.ref022","doi-asserted-by":"crossref","first-page":"51","DOI":"10.1016\/j.cobeha.2017.06.003","article-title":"Remembering Goal Locations","volume":"17","author":"B Poucet","year":"2017","journal-title":"Current Opinion in Behavioral Sciences"},{"issue":"8","key":"pcbi.1010320.ref023","doi-asserted-by":"crossref","first-page":"995","DOI":"10.1038\/nn.2599","article-title":"The Reorganization and Reactivation of Hippocampal Maps Predict Spatial Memory Performance","volume":"13","author":"D Dupret","year":"2010","journal-title":"Nature Neuroscience"},{"issue":"6321","key":"pcbi.1010320.ref024","doi-asserted-by":"crossref","first-page":"176","DOI":"10.1126\/science.aak9589","article-title":"Vectorial Representation of Spatial Goals in the Hippocampus of Bats","volume":"355","author":"A Sarel","year":"2017","journal-title":"Science"},{"key":"pcbi.1010320.ref025","doi-asserted-by":"crossref","first-page":"e52466","DOI":"10.7554\/eLife.52466","article-title":"The Hippocampus Encodes Delay and Value Information during Delay-Discounting Decision Making","volume":"9","author":"A Masuda","year":"2020","journal-title":"eLife"},{"issue":"7647","key":"pcbi.1010320.ref026","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1038\/nature21692","article-title":"Mapping of a Non-Spatial Dimension by the Hippocampal\u2013Entorhinal Circuit","volume":"543","author":"D Aronov","year":"2017","journal-title":"Nature"},{"issue":"2","key":"pcbi.1010320.ref027","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":"JS Taube","year":"1990","journal-title":"Journal of Neuroscience"},{"issue":"1-2","key":"pcbi.1010320.ref028","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1016\/j.cell.2015.12.015","article-title":"Causal Influence of Visual Cues on Hippocampal Directional Selectivity","volume":"164","author":"L Acharya","year":"2016","journal-title":"Cell"},{"issue":"1","key":"pcbi.1010320.ref029","doi-asserted-by":"crossref","first-page":"2333","DOI":"10.1038\/s41467-019-10139-7","article-title":"Heading Direction with Respect to a Reference Point Modulates Place-Cell Activity","volume":"10","author":"PE Jercog","year":"2019","journal-title":"Nature Communications"},{"issue":"1-2","key":"pcbi.1010320.ref030","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1016\/S0921-8890(99)00069-X","article-title":"Biomimetic Robot Navigation","volume":"30","author":"MO Franz","year":"2000","journal-title":"Robotics and Autonomous Systems"},{"issue":"1","key":"pcbi.1010320.ref031","doi-asserted-by":"crossref","first-page":"2713","DOI":"10.1038\/s41598-021-81157-z","article-title":"Context-Dependent Extinction Learning Emerging from Raw Sensory Inputs: A Reinforcement Learning Approach","volume":"11","author":"T Walther","year":"2021","journal-title":"Scientific Reports"},{"key":"pcbi.1010320.ref032","doi-asserted-by":"crossref","DOI":"10.3389\/fncom.2020.00063","article-title":"The Neuroscience of Spatial Navigation and the Relationship to Artificial Intelligence","volume":"14","author":"E Bermudez-Contreras","year":"2020","journal-title":"Frontiers in Computational Neuroscience"},{"issue":"3-4","key":"pcbi.1010320.ref033","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1007\/BF00992698","article-title":"Q-learning","volume":"8","author":"CJCH Watkins","year":"1992","journal-title":"Machine Learning"},{"issue":"11","key":"pcbi.1010320.ref034","doi-asserted-by":"crossref","first-page":"2278","DOI":"10.1109\/5.726791","article-title":"Gradient-Based Learning Applied to Document Recognition","volume":"86","author":"Y Lecun","year":"1998","journal-title":"Proceedings of the IEEE"},{"key":"pcbi.1010320.ref035","unstructured":"Nair V, Hinton GE. Rectified Linear Units Improve Restricted Boltzmann Machines. In: Proceedings of the 27th International Conference on International Conference on Machine Learning. ICML\u201910. Madison, WI, USA: Omnipress; 2010. p. 807\u2013814."},{"issue":"7540","key":"pcbi.1010320.ref036","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1038\/nature14236","article-title":"Human-Level Control through Deep Reinforcement Learning","volume":"518","author":"V Mnih","year":"2015","journal-title":"Nature"},{"key":"pcbi.1010320.ref037","volume-title":"Advances in Neural Information Processing Systems","author":"H Hasselt","year":"2010"},{"key":"pcbi.1010320.ref038","unstructured":"Wang Z, Schaul T, Hessel M, van Hasselt H, Lanctot M, de Freitas N. Dueling Network Architectures for Deep Reinforcement Learning. arXiv:151106581 [cs]. 2016;."},{"key":"pcbi.1010320.ref039","doi-asserted-by":"crossref","DOI":"10.3389\/fnhum.2014.00803","article-title":"A Critical Review of the Allocentric Spatial Representation and Its Neural Underpinnings: Toward a Network-Based Perspective","volume":"8","author":"AD Ekstrom","year":"2014","journal-title":"Frontiers in human neuroscience"},{"issue":"11","key":"pcbi.1010320.ref040","doi-asserted-by":"crossref","first-page":"7079","DOI":"10.1523\/JNEUROSCI.15-11-07079.1995","article-title":"Interactions between Location and Task Affect the Spatial and Directional Firing of Hippocampal Neurons","volume":"15","author":"EJ Markus","year":"1995","journal-title":"Journal of Neuroscience"},{"key":"pcbi.1010320.ref041","article-title":"Are All Spatial Reference Frames Egocentric? Reinterpreting Evidence for Allocentric, Object-Centered, or World-Centered Reference Frames","author":"F Filimon","year":"2015","journal-title":"Frontiers in Human Neuroscience"},{"issue":"3","key":"pcbi.1010320.ref042","doi-asserted-by":"crossref","first-page":"518","DOI":"10.1016\/j.neuropsychologia.2010.12.031","article-title":"Egocentric and Allocentric Memory as Assessed by Virtual Reality in Individuals with Amnestic Mild Cognitive Impairment","volume":"49","author":"G Weniger","year":"2011","journal-title":"Neuropsychologia"},{"issue":"7705","key":"pcbi.1010320.ref043","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":"A Banino","year":"2018","journal-title":"Nature"},{"key":"pcbi.1010320.ref044","unstructured":"Cueva CJ, Wei XX. Emergence of Grid-like Representations by Training Recurrent Neural Networks to Perform Spatial Localization. arXiv:180307770 [cs, q-bio, stat]. 2018;."},{"issue":"4","key":"pcbi.1010320.ref045","doi-asserted-by":"crossref","first-page":"751","DOI":"10.1037\/0735-7044.118.4.751","article-title":"Context-Dependent Reorganization of Spatial and Movement Representations by Simultaneously Recorded Hippocampal and Striatal Neurons During Performance of Allocentric and Egocentric Tasks","volume":"118","author":"O Yeshenko","year":"2004","journal-title":"Behavioral Neuroscience"},{"issue":"2","key":"pcbi.1010320.ref046","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1016\/0959-4388(93)90206-E","article-title":"Coordinate Transformations in the Representation of Spatial Information","volume":"3","author":"RA Andersen","year":"1993","journal-title":"Current Opinion in Neurobiology"},{"issue":"5","key":"pcbi.1010320.ref047","doi-asserted-by":"crossref","first-page":"416","DOI":"10.1037\/bne0000260","article-title":"The Retrosplenial-Parietal Network and Reference Frame Coordination for Spatial Navigation","volume":"132","author":"BJ Clark","year":"2018","journal-title":"Behavioral Neuroscience"},{"issue":"1","key":"pcbi.1010320.ref048","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1037\/0882-7974.6.1.10","article-title":"Spatial Cognition and Behavior in Young and Elderly Adults: Implications for Learning New Environments","volume":"6","author":"KC Kirasic","year":"1991","journal-title":"Psychology and Aging"},{"issue":"1","key":"pcbi.1010320.ref049","doi-asserted-by":"crossref","first-page":"S49","DOI":"10.1093\/geronb\/54B.1.S49","article-title":"Navigation and the Mobility of Older Drivers","volume":"54","author":"PC Burns","year":"1999","journal-title":"The Journals of Gerontology Series B, Psychological Sciences and Social Sciences"},{"issue":"3","key":"pcbi.1010320.ref050","doi-asserted-by":"crossref","first-page":"164","DOI":"10.1016\/j.rehab.2015.12.004","article-title":"The Contribution of Virtual Reality to the Diagnosis of Spatial Navigation Disorders and to the Study of the Role of Navigational Aids: A Systematic Literature Review","volume":"60","author":"M Cogn\u00e9","year":"2017","journal-title":"Annals of Physical and Rehabilitation Medicine"},{"issue":"5","key":"pcbi.1010320.ref051","doi-asserted-by":"crossref","first-page":"851","DOI":"10.1037\/0735-7044.116.5.851","article-title":"Effects of Age on Virtual Environment Place Navigation and Allocentric Cognitive Mapping","volume":"116","author":"SD Moffat","year":"2002","journal-title":"Behavioral Neuroscience"},{"issue":"8","key":"pcbi.1010320.ref052","doi-asserted-by":"crossref","first-page":"496","DOI":"10.1038\/s41582-018-0031-x","article-title":"Spatial Navigation Deficits\u2014Overlooked Cognitive Marker for Preclinical Alzheimer Disease?","volume":"14","author":"G Coughlan","year":"2018","journal-title":"Nature Reviews Neurology"},{"issue":"1","key":"pcbi.1010320.ref053","doi-asserted-by":"crossref","first-page":"77","DOI":"10.3233\/JAD-150855","article-title":"Spatial Navigation in Preclinical Alzheimer\u2019s Disease","volume":"52","author":"SL Allison","year":"2016","journal-title":"Journal of Alzheimer\u2019s disease: JAD"},{"issue":"1","key":"pcbi.1010320.ref054","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1016\/j.schres.2008.01.011","article-title":"Allocentric Memory Impaired and Egocentric Memory Intact as Assessed by Virtual Reality in Recent-Onset Schizophrenia","volume":"101","author":"G Weniger","year":"2008","journal-title":"Schizophrenia Research"},{"issue":"4","key":"pcbi.1010320.ref055","doi-asserted-by":"crossref","first-page":"1189","DOI":"10.1037\/a0034819","article-title":"Spatial Navigation Impairments Among Intellectually High-Functioning Adults With Autism Spectrum Disorder: Exploring Relations With Theory of Mind, Episodic Memory, and Episodic Future Thinking","volume":"122","author":"SE Lind","year":"2013","journal-title":"Journal of Abnormal Psychology"},{"issue":"3","key":"pcbi.1010320.ref056","doi-asserted-by":"crossref","first-page":"649","DOI":"10.1016\/j.psychres.2015.04.051","article-title":"Allocentric but Not Egocentric Visual Memory Difficulties in Adults with ADHD May Represent Cognitive Inefficiency","volume":"228","author":"FC Brown","year":"2015","journal-title":"Psychiatry Research"},{"key":"pcbi.1010320.ref057","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/j.nlm.2016.12.007","article-title":"Search Strategy Selection in the Morris Water Maze Indicates Allocentric Map Formation during Learning That Underpins Spatial Memory Formation","volume":"139","author":"J Rogers","year":"2017","journal-title":"Neurobiology of Learning and Memory"},{"issue":"1","key":"pcbi.1010320.ref058","doi-asserted-by":"crossref","first-page":"11338","DOI":"10.1038\/s41598-020-68025-y","article-title":"Flexible Use of Allocentric and Egocentric Spatial Memories Activates Differential Neural Networks in Mice","volume":"10","author":"A Rinaldi","year":"2020","journal-title":"Scientific Reports"},{"key":"pcbi.1010320.ref059","volume-title":"The Hippocampus as a Cognitive Map","author":"J O\u2019Keefe","year":"1978"},{"key":"pcbi.1010320.ref060","doi-asserted-by":"crossref","unstructured":"Bota A, Goto A, Tsukamoto S, Schmidt A, Wolf F, Luchetti A, et al.. Shared and Unique Properties of Place Cells in Anterior Cingulate Cortex and Hippocampus; 2021.","DOI":"10.1101\/2021.03.29.437441"},{"issue":"4","key":"pcbi.1010320.ref061","doi-asserted-by":"crossref","first-page":"432","DOI":"10.1002\/(SICI)1098-1063(1999)9:4<432::AID-HIPO9>3.0.CO;2-P","article-title":"Complimentary Roles for Hippocampal versus Subicular\/Entorhinal Place Cells in Coding Place, Context, and Events","volume":"9","author":"PE Sharp","year":"1999","journal-title":"Hippocampus"},{"issue":"5688","key":"pcbi.1010320.ref062","doi-asserted-by":"crossref","first-page":"1258","DOI":"10.1126\/science.1099901","article-title":"Spatial Representation in the Entorhinal Cortex","volume":"305","author":"M Fyhn","year":"2004","journal-title":"Science"},{"issue":"3","key":"pcbi.1010320.ref063","doi-asserted-by":"crossref","first-page":"1067","DOI":"10.1523\/JNEUROSCI.5393-12.2014","article-title":"Encoding of Head Direction by Hippocampal Place Cells in Bats","volume":"34","author":"A Rubin","year":"2014","journal-title":"The Journal of Neuroscience: The Official Journal of the Society for Neuroscience"},{"issue":"21","key":"pcbi.1010320.ref064","doi-asserted-by":"crossref","first-page":"3521","DOI":"10.1016\/j.neuron.2021.09.032","article-title":"Spatial Modulation of Hippocampal Activity in Freely Moving Macaques","volume":"109","author":"D Mao","year":"2021","journal-title":"Neuron"},{"issue":"1","key":"pcbi.1010320.ref065","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1007\/BF00237147","article-title":"The Contributions of Position, Direction, and Velocity to Single Unit Activity in the Hippocampus of Freely-Moving Rats","volume":"52","author":"BL McNaughton","year":"1983","journal-title":"Experimental Brain Research"},{"key":"pcbi.1010320.ref066","doi-asserted-by":"crossref","first-page":"598570","DOI":"10.3389\/fnbeh.2020.598570","article-title":"Place vs. Response Learning: History, Controversy, and Neurobiology","volume":"14","author":"J Goodman","year":"2021","journal-title":"Frontiers in Behavioral Neuroscience"},{"issue":"2","key":"pcbi.1010320.ref067","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1037\/h0055224","article-title":"Studies in Spatial Learning. VIII. Place Performance and the Acquisition of Place Dispositions","volume":"43","author":"BF Ritchie","year":"1950","journal-title":"Journal of Comparative and Physiological Psychology"},{"issue":"2","key":"pcbi.1010320.ref068","doi-asserted-by":"crossref","first-page":"459","DOI":"10.2466\/pr0.1964.15.2.459","article-title":"Effects of Overtraining on Acquisition and Reversal of Place and Response Learning","volume":"15","author":"LH Hicks","year":"1964","journal-title":"Psychological Reports"},{"issue":"1","key":"pcbi.1010320.ref069","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1006\/nlme.1996.0007","article-title":"Inactivation of Hippocampus or Caudate Nucleus with Lidocaine Differentially Affects Expression of Place and Response Learning","volume":"65","author":"MG Packard","year":"1996","journal-title":"Neurobiology of Learning and Memory"},{"issue":"22","key":"pcbi.1010320.ref070","doi-asserted-by":"crossref","first-page":"12881","DOI":"10.1073\/pnas.96.22.12881","article-title":"Glutamate Infused Posttraining into the Hippocampus or Caudate-Putamen Differentially Strengthens Place and Response Learning","volume":"96","author":"MG Packard","year":"1999","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"pcbi.1010320.ref071","doi-asserted-by":"crossref","DOI":"10.3389\/fnbeh.2012.00079","article-title":"Integrating Cortico-Limbic-Basal Ganglia Architectures for Learning Model-Based and Model-Free Navigation Strategies","volume":"6","author":"M Khamassi","year":"2012","journal-title":"Frontiers in Behavioral Neuroscience"},{"issue":"3","key":"pcbi.1010320.ref072","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1385\/NI:3:3:223","article-title":"A Computational Model of Parallel Navigation Systems in Rodents","volume":"3","author":"R Chavarriaga","year":"2005","journal-title":"Neuroinformatics"},{"issue":"7","key":"pcbi.1010320.ref073","doi-asserted-by":"crossref","first-page":"484","DOI":"10.1002\/hipo.22847","article-title":"The response strategy and the place strategy in a plus-maze have different sensitivities to devaluation of expected outcome","volume":"28","author":"Y Kosaki","year":"2018","journal-title":"Hippocampus"},{"issue":"3","key":"pcbi.1010320.ref074","doi-asserted-by":"crossref","first-page":"683","DOI":"10.1016\/j.neuron.2019.02.014","article-title":"Hippocampal Contributions to Model-Based Planning and Spatial Memory","volume":"102","author":"OM Vikbladh","year":"2019","journal-title":"Neuron"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1010320","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,11,10]],"date-time":"2022-11-10T00:00:00Z","timestamp":1668038400000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1010320","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,11,10]],"date-time":"2022-11-10T13:50:47Z","timestamp":1668088247000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1010320"}},"subtitle":[],"editor":[{"given":"Daniel","family":"Bush","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2022,10,31]]},"references-count":74,"journal-issue":{"issue":"10","published-online":{"date-parts":[[2022,10,31]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1010320","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2022.06.22.497126","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,10,31]]}}}