{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,1]],"date-time":"2026-08-01T21:22:26Z","timestamp":1785619346831,"version":"3.56.0"},"reference-count":84,"publisher":"Public Library of Science (PLoS)","issue":"6","license":[{"start":{"date-parts":[[2025,6,23]],"date-time":"2025-06-23T00:00:00Z","timestamp":1750636800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100007766","name":"Max-Planck-Institut f\u00fcr Kognitions- und Neurowissenschaften","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100007766","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100013296","name":"Max-Planck-Institut f\u00fcr Mathematik in den Naturwissenschaften","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100013296","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100018668","name":"Max Planck School of Cognition","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100018668","id-type":"DOI","asserted-by":"publisher"}]},{"name":"ScaDS.AI Leipzig"},{"DOI":"10.13039\/100010269","name":"Wellcome Trust","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100010269","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>In spatial cognition, the Successor Representation (SR) from reinforcement learning provides a compelling candidate of how predictive representations are used to encode space. In particular, hippocampal place cells are hypothesized to encode the SR. Here, we investigate how varying the temporal symmetry in learning rules influences those representations. To this end, we use a simple local learning rule which can be made insensitive to the temporal order. We analytically find that a symmetric learning rule results in a successor representation under a symmetrized version of the experienced transition structure. We then apply this rule to a two-layer neural network model loosely resembling hippocampal subfields CA3 - with a symmetric learning rule and recurrent weights - and CA1 - with an asymmetric learning rule and no recurrent weights. Here, when exposed repeatedly to a linear track, neurons in our model in CA3 show less shift of the centre of mass than those in CA1, in line with existing empirical findings. Investigating the functional benefits of such symmetry, we employ a simple reinforcement learning agent which may learn symmetric or classical successor representations. Here, we find that using a symmetric learning rule yields representations which afford better generalization, when the agent is probed to navigate to a new target without relearning the SR. This effect is reversed when the state space is not symmetric anymore. Thus, our results hint at a potential benefit of the inductive bias afforded by symmetric learning rules in areas employed in spatial navigation, where there naturally is a symmetry in the state space.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1013056","type":"journal-article","created":{"date-parts":[[2025,6,23]],"date-time":"2025-06-23T13:32:04Z","timestamp":1750685524000},"page":"e1013056","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":2,"title":["Impact of symmetry in local learning rules on predictive neural representations and generalization in spatial navigation"],"prefix":"10.1371","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0009-0009-8814-4933","authenticated-orcid":true,"given":"Janis","family":"Keck","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Caswell","family":"Barry","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christian F.","family":"Doeller","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5258-6590","authenticated-orcid":true,"given":"J\u00fcrgen","family":"Jost","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2025,6,23]]},"reference":[{"issue":"1","key":"pcbi.1013056.ref001","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1136\/jnnp.20.1.11","article-title":"Loss of recent memory after bilateral hippocampal lesions","volume":"20","author":"WB Scoville","year":"1957","journal-title":"J Neurol Neurosurg Psychiatry"},{"key":"pcbi.1013056.ref002","first-page":"570","article-title":"The hippocampus as a cognitive map","volume":"3","author":"J O\u2019Keefe","year":"1978","journal-title":"Hippocampus"},{"issue":"2","key":"pcbi.1013056.ref003","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1016\/S0896-6273(00)80773-4","article-title":"The hippocampus, memory, and place cells: is it spatial memory or a memory space?","volume":"23","author":"H Eichenbaum","year":"1999","journal-title":"Neuron"},{"key":"pcbi.1013056.ref004","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1146\/annurev.neuro.27.070203.144130","article-title":"The medial temporal lobe","volume":"27","author":"LR Squire","year":"2004","journal-title":"Annu Rev Neurosci"},{"issue":"7052","key":"pcbi.1013056.ref005","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":"1","key":"pcbi.1013056.ref006","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1038\/4580","article-title":"Predictive coding in the visual cortex: a functional interpretation of some extra-classical receptive-field effects","volume":"2","author":"RP Rao","year":"1999","journal-title":"Nat Neurosci"},{"issue":"2","key":"pcbi.1013056.ref007","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/S0301-0082(02)00076-X","article-title":"Functional integration and inference in the brain","volume":"68","author":"K Friston","year":"2002","journal-title":"Prog Neurobiol"},{"issue":"11","key":"pcbi.1013056.ref008","doi-asserted-by":"crossref","first-page":"1643","DOI":"10.1038\/nn.4650","article-title":"The hippocampus as a predictive map","volume":"20","author":"KL Stachenfeld","year":"2017","journal-title":"Nat Neurosci"},{"issue":"9","key":"pcbi.1013056.ref009","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1005768","article-title":"Predictive representations can link model-based reinforcement learning to model-free mechanisms","volume":"13","author":"EM Russek","year":"2017","journal-title":"PLoS Comput Biol"},{"issue":"2","key":"pcbi.1013056.ref010","doi-asserted-by":"crossref","first-page":"490","DOI":"10.1016\/j.neuron.2018.10.002","article-title":"What is a cognitive map? Organizing knowledge for flexible behavior","volume":"100","author":"TEJ Behrens","year":"2018","journal-title":"Neuron"},{"issue":"4","key":"pcbi.1013056.ref011","doi-asserted-by":"crossref","first-page":"613","DOI":"10.1162\/neco.1993.5.4.613","article-title":"Improving generalization for temporal difference learning: the successor representation","volume":"5","author":"P Dayan","year":"1993","journal-title":"Neural Computation"},{"key":"pcbi.1013056.ref012","article-title":"Successor features for transfer in reinforcement learning","volume":"30","author":"A Barreto","year":"2017","journal-title":"Adv Neural Inf Process Syst"},{"issue":"9","key":"pcbi.1013056.ref013","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1038\/s41562-017-0180-8","article-title":"The successor representation in human reinforcement learning","volume":"1","author":"I Momennejad","year":"2017","journal-title":"Nat Hum Behav"},{"issue":"33","key":"pcbi.1013056.ref014","doi-asserted-by":"crossref","first-page":"7193","DOI":"10.1523\/JNEUROSCI.0151-18.2018","article-title":"The successor representation: its computational logic and neural substrates","volume":"38","author":"SJ Gershman","year":"2018","journal-title":"J Neurosci"},{"key":"pcbi.1013056.ref015","article-title":"Eigenoption discovery through the deep successor representation","author":"MC Machado","year":"2017","journal-title":"arXiv preprint"},{"key":"pcbi.1013056.ref016","article-title":"Successor-predecessor intrinsic exploration","author":"C Yu","year":"2023","journal-title":"arXiv preprint"},{"key":"pcbi.1013056.ref017","article-title":"Predictive representations: building blocks of intelligence","author":"W Carvalho","year":"2024","journal-title":"arXiv preprint"},{"key":"pcbi.1013056.ref018","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.10094","article-title":"Extracting grid cell characteristics from place cell inputs using non-negative principal component analysis","volume":"5","author":"Y Dordek","year":"2016","journal-title":"Elife"},{"key":"pcbi.1013056.ref019","article-title":"Attractor network dynamics enable preplay and rapid path planning in maze\u2013like environments","volume":"28","author":"DS Corneil","year":"2015","journal-title":"Adv Neural Inf Process Syst"},{"issue":"12","key":"pcbi.1013056.ref020","doi-asserted-by":"crossref","first-page":"1347","DOI":"10.1002\/hipo.23246","article-title":"Neurobiological successor features for spatial navigation","volume":"30","author":"W de Cothi","year":"2020","journal-title":"Hippocampus"},{"issue":"17","key":"pcbi.1013056.ref021","doi-asserted-by":"crossref","DOI":"10.1016\/j.cub.2022.06.090","article-title":"Predictive maps in rats and humans for spatial navigation","volume":"32","author":"W de Cothi","year":"2022","journal-title":"Curr Biol"},{"key":"pcbi.1013056.ref022","article-title":"A neurally plausible model learns successor representations in partially observable environments","volume":"32","author":"E V\u00e9rtes","year":"2019","journal-title":"Adv Neural Inf Process Syst"},{"key":"pcbi.1013056.ref023","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.80663","article-title":"Rapid learning of predictive maps with STDP and theta phase precession","volume":"12","author":"TM George","year":"2023","journal-title":"Elife"},{"key":"pcbi.1013056.ref024","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.80671","article-title":"Learning predictive cognitive maps with spiking neurons during behavior and replays","volume":"12","author":"J Bono","year":"2023","journal-title":"Elife"},{"key":"pcbi.1013056.ref025","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.80680","article-title":"Neural learning rules for generating flexible predictions and computing the successor representation","volume":"12","author":"C Fang","year":"2023","journal-title":"Elife"},{"issue":"23","key":"pcbi.1013056.ref026","doi-asserted-by":"crossref","DOI":"10.1016\/j.cub.2015.10.049","article-title":"The hippocampus","volume":"25","author":"JJ Knierim","year":"2015","journal-title":"Curr Biol"},{"issue":"5688","key":"pcbi.1013056.ref027","doi-asserted-by":"crossref","first-page":"1295","DOI":"10.1126\/science.1100265","article-title":"Distinct ensemble codes in hippocampal areas CA3 and CA1","volume":"305","author":"S Leutgeb","year":"2004","journal-title":"Science"},{"issue":"24","key":"pcbi.1013056.ref028","doi-asserted-by":"crossref","first-page":"6610","DOI":"10.1523\/JNEUROSCI.5388-05.2006","article-title":"Malleability of spike-timing-dependent plasticity at the CA3-CA1 synapse","volume":"26","author":"GM Wittenberg","year":"2006","journal-title":"J Neurosci"},{"key":"pcbi.1013056.ref029","doi-asserted-by":"crossref","first-page":"4","DOI":"10.3389\/fnsyn.2011.00004","article-title":"A history of spike-timing-dependent plasticity","volume":"3","author":"H Markram","year":"2011","journal-title":"Front Synaptic Neurosci"},{"issue":"5297","key":"pcbi.1013056.ref030","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1126\/science.275.5297.213","article-title":"Regulation of synaptic efficacy by coincidence of postsynaptic APs and EPSPs","volume":"275","author":"H Markram","year":"1997","journal-title":"Science"},{"issue":"24","key":"pcbi.1013056.ref031","doi-asserted-by":"crossref","first-page":"10464","DOI":"10.1523\/JNEUROSCI.18-24-10464.1998","article-title":"Synaptic modifications in cultured hippocampal neurons: dependence on spike timing, synaptic strength, and postsynaptic cell type","volume":"18","author":"GQ Bi","year":"1998","journal-title":"J Neurosci"},{"key":"pcbi.1013056.ref032","doi-asserted-by":"crossref","first-page":"11552","DOI":"10.1038\/ncomms11552","article-title":"Symmetric spike timing-dependent plasticity at CA3-CA3 synapses optimizes storage and recall in autoassociative networks","volume":"7","author":"RK Mishra","year":"2016","journal-title":"Nat Commun"},{"issue":"5","key":"pcbi.1013056.ref033","doi-asserted-by":"crossref","first-page":"803","DOI":"10.1016\/j.neuron.2004.05.010","article-title":"A double dissociation between hippocampal subfields: differential time course of CA3 and CA1 place cells for processing changed environments","volume":"42","author":"I Lee","year":"2004","journal-title":"Neuron"},{"issue":"6998","key":"pcbi.1013056.ref034","doi-asserted-by":"crossref","first-page":"456","DOI":"10.1038\/nature02739","article-title":"Comparison of population coherence of place cells in hippocampal subfields CA1 and CA3","volume":"430","author":"I Lee","year":"2004","journal-title":"Nature"},{"issue":"16","key":"pcbi.1013056.ref035","doi-asserted-by":"crossref","first-page":"8918","DOI":"10.1073\/pnas.94.16.8918","article-title":"Experience-dependent, asymmetric expansion of hippocampal place fields","volume":"94","author":"MR Mehta","year":"1997","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"4","key":"pcbi.1013056.ref036","article-title":"Functional differences in the backward shifts of CA1 and CA3 place fields in novel and familiar environments","volume":"7","author":"ED Roth","year":"2012","journal-title":"PLoS One"},{"issue":"1","key":"pcbi.1013056.ref037","doi-asserted-by":"crossref","first-page":"2977","DOI":"10.1038\/s41467-021-23260-3","article-title":"Distinct place cell dynamics in CA1 and CA3 encode experience in new environments","volume":"12","author":"C Dong","year":"2021","journal-title":"Nat Commun"},{"key":"pcbi.1013056.ref038","volume-title":"Reinforcement learning: an introduction","author":"RS Sutton","year":"2018"},{"key":"pcbi.1013056.ref039","article-title":"Endotaxis: a neuromorphic algorithm for mapping, goal-learning, navigation, and patrolling","author":"T Zhang","year":"2021","journal-title":"bioRxiv"},{"issue":"8","key":"pcbi.1013056.ref040","doi-asserted-by":"crossref","first-page":"1659","DOI":"10.1002\/hipo.22002","article-title":"Activity dynamics and behavioral correlates of CA3 and CA1 hippocampal pyramidal neurons","volume":"22","author":"K Mizuseki","year":"2012","journal-title":"Hippocampus"},{"key":"pcbi.1013056.ref041","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.85274","article-title":"RatInABox, a toolkit for modelling locomotion and neuronal activity in continuous environments","volume":"13","author":"TM George","year":"2024","journal-title":"Elife"},{"key":"pcbi.1013056.ref042","first-page":"501","article-title":"Transfer in deep reinforcement learning using successor features and generalised policy improvement.","volume-title":"International Conference on Machine Learning","author":"A Barreto","year":"2018"},{"issue":"1","key":"pcbi.1013056.ref043","doi-asserted-by":"crossref","first-page":"4942","DOI":"10.1038\/s41467-021-25123-3","article-title":"Linear reinforcement learning in planning, grid fields, and cognitive control","volume":"12","author":"P Piray","year":"2021","journal-title":"Nat Commun"},{"key":"pcbi.1013056.ref044","doi-asserted-by":"crossref","first-page":"836498","DOI":"10.3389\/fncom.2022.836498","article-title":"Symmetry-based representations for artificial and biological general intelligence","volume":"16","author":"I Higgins","year":"2022","journal-title":"Front Comput Neurosci"},{"key":"pcbi.1013056.ref045","first-page":"31497","article-title":"Symmetry-induced disentanglement on graphs","volume":"35","author":"G Mercatali","year":"2022","journal-title":"Adv Neural Inf Process Syst"},{"key":"pcbi.1013056.ref046","article-title":"Plannable approximations to mdp homomorphisms: equivariance under actions","author":"E Van der Pol","year":"2020","journal-title":"arXiv preprint"},{"key":"pcbi.1013056.ref047","article-title":"A simple approach for state-action abstraction using a learned mdp homomorphism","author":"AN Mavor-Parker","year":"2022","journal-title":"arXiv preprint"},{"key":"pcbi.1013056.ref048","first-page":"4199","article-title":"Mdp homomorphic networks: group symmetries in reinforcement learning","volume":"33","author":"E Van der Pol","year":"2020","journal-title":"Adv Neural Inf Process Syst"},{"key":"pcbi.1013056.ref049","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1016\/S0004-3702(02)00376-4","article-title":"Equivalence notions and model minimization in Markov decision processes","volume":"147","author":"R Givan","year":"2003","journal-title":"Artif Intell"},{"key":"pcbi.1013056.ref050","article-title":"Approximate temporal difference learning is a gradient descent for reversible policies","author":"Y Ollivier","year":"2018","journal-title":"arXiv preprint"},{"issue":"24","key":"pcbi.1013056.ref051","doi-asserted-by":"crossref","first-page":"10664","DOI":"10.1523\/JNEUROSCI.19-24-10664.1999","article-title":"Heterogeneity of synaptic plasticity at unitary CA3-CA1 and CA3-CA3 connections in rat hippocampal slice cultures","volume":"19","author":"D Debanne","year":"1999","journal-title":"J Neurosci"},{"issue":"22","key":"pcbi.1013056.ref052","doi-asserted-by":"crossref","first-page":"3552","DOI":"10.1016\/j.neuron.2021.09.034","article-title":"The learning of prospective and retrospective cognitive maps within neural circuits","volume":"109","author":"VM K Namboodiri","year":"2021","journal-title":"Neuron"},{"issue":"4","key":"pcbi.1013056.ref053","doi-asserted-by":"crossref","first-page":"563","DOI":"10.1016\/j.neuron.2019.05.041","article-title":"Neuromodulation of spike-timing-dependent plasticity: past, present, and future","volume":"103","author":"Z Brzosko","year":"2019","journal-title":"Neuron"},{"issue":"6","key":"pcbi.1013056.ref054","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1016\/j.neuron.2007.08.013","article-title":"Neuromodulators control the polarity of spike-timing-dependent synaptic plasticity","volume":"55","author":"GH Seol","year":"2007","journal-title":"Neuron"},{"key":"pcbi.1013056.ref055","doi-asserted-by":"crossref","first-page":"146","DOI":"10.3389\/fnsyn.2010.00146","article-title":"Timing is not everything: neuromodulation opens the STDP gate","volume":"2","author":"V Pawlak","year":"2010","journal-title":"Front Synaptic Neurosci"},{"key":"pcbi.1013056.ref056","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.09685","article-title":"Retroactive modulation of spike timing-dependent plasticity by dopamine","volume":"4","author":"Z Brzosko","year":"2015","journal-title":"Elife"},{"key":"pcbi.1013056.ref057","doi-asserted-by":"crossref","first-page":"91","DOI":"10.1016\/j.neuroscience.2011.06.064","article-title":"Cholinergic modulation on spike timing-dependent plasticity in hippocampal CA1 network","volume":"192","author":"E Sugisaki","year":"2011","journal-title":"Neuroscience"},{"issue":"1","key":"pcbi.1013056.ref058","doi-asserted-by":"crossref","first-page":"9486","DOI":"10.1038\/s41598-018-27393-2","article-title":"Acetylcholine-modulated plasticity in reward-driven navigation: a computational study","volume":"8","author":"S Zannone","year":"2018","journal-title":"Sci Rep"},{"issue":"10","key":"pcbi.1013056.ref059","doi-asserted-by":"crossref","first-page":"1241","DOI":"10.1038\/nn1961","article-title":"Forward and reverse hippocampal place-cell sequences during ripples","volume":"10","author":"K Diba","year":"2007","journal-title":"Nat Neurosci"},{"issue":"7084","key":"pcbi.1013056.ref060","doi-asserted-by":"crossref","first-page":"680","DOI":"10.1038\/nature04587","article-title":"Reverse replay of behavioural sequences in hippocampal place cells during the awake state","volume":"440","author":"DJ Foster","year":"2006","journal-title":"Nature"},{"issue":"12","key":"pcbi.1013056.ref061","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1003383","article-title":"Forward and backward inference in spatial cognition","volume":"9","author":"WD Penny","year":"2013","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1013056.ref062","article-title":"Learning state representations via retracing in reinforcement learning","author":"C Yu","year":"2021","journal-title":"arXiv preprint"},{"key":"pcbi.1013056.ref063","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.34171","article-title":"Recurrent network model for learning goal-directed sequences through reverse replay","volume":"7","author":"T Haga","year":"2018","journal-title":"Elife"},{"issue":"3","key":"pcbi.1013056.ref064","doi-asserted-by":"crossref","first-page":"573","DOI":"10.1037\/a0028681","article-title":"Generalization through the recurrent interaction of episodic memories: a model of the hippocampal system","volume":"119","author":"D Kumaran","year":"2012","journal-title":"Psychol Rev"},{"key":"pcbi.1013056.ref065","doi-asserted-by":"crossref","DOI":"10.1155\/2008\/381243","article-title":"What does the anatomical organization of the entorhinal cortex tell us?","volume":"2008","author":"CB Canto","year":"2008","journal-title":"Neural plasticity"},{"key":"pcbi.1013056.ref066","first-page":"1644","article-title":"A generative model of the hippocampal formation trained with theta driven local learning rules","volume":"36","author":"TM George","year":"2023","journal-title":"Adv Neural Inf Process Syst"},{"issue":"5","key":"pcbi.1013056.ref067","article-title":"The Tolman-Eichenbaum machine: unifying space and relational memory through generalization in the hippocampal formation","volume":"183","author":"JCR Whittington","year":"2020","journal-title":"Cell"},{"issue":"27","key":"pcbi.1013056.ref068","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":"IR Fiete","year":"2008","journal-title":"J Neurosci"},{"issue":"6","key":"pcbi.1013056.ref069","doi-asserted-by":"crossref","first-page":"2112","DOI":"10.1523\/JNEUROSCI.16-06-02112.1996","article-title":"Representation of spatial orientation by the intrinsic dynamics of the head-direction cell ensemble: a theory","volume":"16","author":"K Zhang","year":"1996","journal-title":"J Neurosci"},{"issue":"15","key":"pcbi.1013056.ref070","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":"A Samsonovich","year":"1997","journal-title":"J Neurosci"},{"issue":"12","key":"pcbi.1013056.ref071","doi-asserted-by":"crossref","first-page":"744","DOI":"10.1038\/s41583-022-00642-0","article-title":"Attractor and integrator networks in the brain","volume":"23","author":"M Khona","year":"2022","journal-title":"Nat Rev Neurosci"},{"issue":"36","key":"pcbi.1013056.ref072","doi-asserted-by":"crossref","first-page":"7675","DOI":"10.1523\/JNEUROSCI.0657-21.2021","article-title":"Learning and representation of hierarchical concepts in hippocampus and prefrontal cortex","volume":"41","author":"S Theves","year":"2021","journal-title":"J Neurosci"},{"issue":"4","key":"pcbi.1013056.ref073","doi-asserted-by":"crossref","first-page":"615","DOI":"10.1038\/s41593-023-01283-x","article-title":"Hippocampal spatio-predictive cognitive maps adaptively guide reward generalization","volume":"26","author":"MM Garvert","year":"2023","journal-title":"Nat Neurosci"},{"key":"pcbi.1013056.ref074","article-title":"Grid-like entorhinal representation of an abstract value space during prospective decision making","author":"A Nitsch","year":"2023","journal-title":"bioRxiv"},{"issue":"8","key":"pcbi.1013056.ref075","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1016\/j.tics.2020.05.008","article-title":"Knowledge across reference frames: cognitive maps and image spaces","volume":"24","author":"R Bottini","year":"2020","journal-title":"Trends Cogn Sci"},{"issue":"6292","key":"pcbi.1013056.ref076","doi-asserted-by":"crossref","first-page":"1464","DOI":"10.1126\/science.aaf0941","article-title":"Organizing conceptual knowledge in humans with a gridlike code","volume":"352","author":"AO Constantinescu","year":"2016","journal-title":"Science"},{"issue":"6","key":"pcbi.1013056.ref077","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":"ME Hasselmo","year":"2006","journal-title":"Curr Opin Neurobiol"},{"issue":"4","key":"pcbi.1013056.ref078","doi-asserted-by":"crossref","first-page":"580","DOI":"10.1002\/cne.902950407","article-title":"Organization of intrahippocampal projections originating from CA3 pyramidal cells in the rat","volume":"295","author":"N Ishizuka","year":"1990","journal-title":"J Comp Neurol"},{"issue":"11","key":"pcbi.1013056.ref079","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1101\/lm.725207","article-title":"Intrinsic and extrinsic wiring of CA3: indications for connectional heterogeneity","volume":"14","author":"MP Witter","year":"2007","journal-title":"Learn Memory"},{"key":"pcbi.1013056.ref080","volume-title":"Markov chains","author":"JR Norris","year":"1998"},{"issue":"11","key":"pcbi.1013056.ref081","doi-asserted-by":"crossref","first-page":"1713","DOI":"10.1162\/neco_a_01611","article-title":"A tutorial on the spectral theory of Markov chains","volume":"35","author":"E Seabrook","year":"2023","journal-title":"Neural Comput"},{"key":"pcbi.1013056.ref082","doi-asserted-by":"crossref","DOI":"10.1090\/mbk\/107","volume-title":"Markov chains and mixing times","author":"DA Levin","year":"2017"},{"issue":"49","key":"pcbi.1013056.ref083","doi-asserted-by":"crossref","first-page":"31427","DOI":"10.1073\/pnas.2007981117","article-title":"A general model of hippocampal and dorsal striatal learning and decision making","volume":"117","author":"JP Geerts","year":"2020","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1013056.ref084","article-title":"Neuro-Nav: a library for neurally-plausible reinforcement learning","author":"A Juliani","year":"2022","journal-title":"arXiv preprint"}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1013056","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,6,23]],"date-time":"2025-06-23T13:32:23Z","timestamp":1750685543000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1013056"}},"subtitle":[],"editor":[{"given":"Suhita","family":"Nadkarni","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2025,6,23]]},"references-count":84,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2025,6,23]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1013056","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2024.05.27.595705","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,6,23]]}}}