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Expectation propagation for approximate bayesian inference. arXiv. https:\/\/arxiv.org\/abs\/1301.2294."},{"key":"bib45","doi-asserted-by":"publisher","first-page":"926","DOI":"10.1126\/science.adq5874","type":"journal-article","article-title":"Hidden state inference requires abstract contextual representations in the ventral hippocampus","volume":"386","author":"Mishchanchuk","year":"2024","journal-title":"Science"},{"key":"bib46","doi-asserted-by":"publisher","first-page":"394","DOI":"10.1016\/j.neuron.2021.12.012","type":"journal-article","article-title":"Spatial goal coding in the hippocampal formation","volume":"110","author":"Nyberg","year":"2022","journal-title":"Neuron"},{"key":"bib47","doi-asserted-by":"publisher","DOI":"10.1101\/2025.01.08.632067","type":"preprint","article-title":"Replay builds an efficient cognitive map offline to avoid computation 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Brain-like slot representation for sequence working memory in recurrent neural networks. Second Workshop on Representational Alignment at ICLR."},{"key":"bib71","doi-asserted-by":"publisher","first-page":"1249","DOI":"10.1016\/j.cell.2020.10.024","type":"journal-article","article-title":"The tolman-eichenbaum machine: unifying space and relational memory through generalization in the hippocampal formation","volume":"183","author":"Whittington","year":"2020","journal-title":"Cell"},{"key":"bib72","doi-asserted-by":"publisher","DOI":"10.1101\/2023.11.05.565662","type":"preprint","article-title":"On prefrontal working memory and hippocampal episodic memory: unifying memories stored in weights and activity slots","volume-title":"bioRxiv","author":"Whittington","year":"2023"},{"key":"bib73","doi-asserted-by":"publisher","first-page":"1547","DOI":"10.1016\/j.neuron.2022.02.002","type":"journal-article","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":"bib74","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2025.115475","type":"journal-article","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 Reports"},{"key":"bib75","doi-asserted-by":"publisher","first-page":"632","DOI":"10.1126\/science.abm0204","type":"journal-article","article-title":"Geometry of sequence working memory in macaque prefrontal cortex","volume":"375","author":"Xie","year":"2022","journal-title":"Science"},{"key":"bib76","doi-asserted-by":"publisher","first-page":"2112","DOI":"10.1523\/JNEUROSCI.16-06-02112.1996","type":"journal-article","article-title":"Representation of spatial orientation by the intrinsic dynamics of the head-direction cell ensemble: a theory","volume":"16","author":"Zhang","year":"1996","journal-title":"The Journal of Neuroscience"},{"key":"bib77","type":"conference-paper","article-title":"Mice in the manhattan maze: Rapid learning, flexible routing and generalization, with and without cortex","author":"Zheng","year":"2024","unstructured":"Zheng J, Guimaraes R, Hu JY, Perona P, Meister M. 2024. Mice in the manhattan maze: Rapid learning, flexible routing and generalization, with and without cortex. Cognitive Computational Neuroscience."},{"key":"bib78","type":"preprint","article-title":"VariBAD: a very good method for bayes-adaptive deep RL via meta-learning","volume-title":"arXiv","author":"Zintgraf","year":"2019","unstructured":"Zintgraf L, Shiarlis K, Igl M, Schulze S, Gal Y, Hofmann K, Whiteson S. 2019. 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