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In line with these observations, here we develop a modelling approach to provide a mechanistic understanding of how hippocampal neural assemblies evolve differently, depending on the frequency of presentation of the stimuli. For this, we added an online Hebbian learning rule, background firing activity, neural adaptation and heterosynaptic plasticity to a rate attractor network model, thus creating dynamic memory representations that can persist, increase or fade according to the frequency of presentation of the corresponding memory patterns. Specifically, we show that a dynamic interplay between Hebbian learning and background firing activity can explain the relationship between the memory assembly sizes and their frequency of stimulation. Frequently stimulated assemblies increase their size independently from each other (i.e. creating orthogonal representations that do not share neurons, thus avoiding interference). Importantly, connections between neurons of assemblies that are not further stimulated become labile so that these neurons can be recruited by other assemblies, providing a neuronal mechanism of forgetting.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1011727","type":"journal-article","created":{"date-parts":[[2023,12,20]],"date-time":"2023-12-20T13:44:01Z","timestamp":1703079841000},"page":"e1011727","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":7,"title":["A dynamic attractor network model of memory formation, reinforcement and forgetting"],"prefix":"10.1371","volume":"19","author":[{"ORCID":"https:\/\/orcid.org\/0009-0005-1298-4662","authenticated-orcid":true,"given":"Marta","family":"Boscaglia","sequence":"first","affiliation":[]},{"given":"Chiara","family":"Gastaldi","sequence":"additional","affiliation":[]},{"given":"Wulfram","family":"Gerstner","sequence":"additional","affiliation":[]},{"given":"Rodrigo","family":"Quian Quiroga","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2023,12,20]]},"reference":[{"key":"pcbi.1011727.ref001","doi-asserted-by":"crossref","DOI":"10.4324\/9780429449642","volume-title":"Memory","author":"A. 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Zenke","year":"2017","journal-title":"Biological Sciences"},{"issue":"12","key":"pcbi.1011727.ref038","doi-asserted-by":"crossref","first-page":"e1009691","DOI":"10.1371\/journal.pcbi.1009691","article-title":"When shared concept cells support associations: Theory of overlapping memory engrams","volume":"17","author":"C. Gastaldi","year":"2021","journal-title":"PLOS Computational Biology"},{"issue":"4","key":"pcbi.1011727.ref039","doi-asserted-by":"crossref","first-page":"439","DOI":"10.1016\/j.conb.2009.07.005","article-title":"Spontaneous and driven cortical activity: implications for computation","volume":"19","author":"D.L. Ringach","year":"2009","journal-title":"Current Opinion in Neurobiology"},{"key":"pcbi.1011727.ref040","doi-asserted-by":"crossref","first-page":"292","DOI":"10.1038\/nrn2258","article-title":"Noise in the nervous system","volume":"9","author":"A.A. Faisal","year":"2008","journal-title":"Nature Reviews Neuroscience"},{"issue":"1","key":"pcbi.1011727.ref041","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1007\/s10827-006-7074-5","article-title":"Predicting spike timing of neocortical pyramidal neurons by simple threshold models","volume":"21","author":"R. Jolivet","year":"2006","journal-title":"Journal of Computational Neuroscience"},{"issue":"015008","key":"pcbi.1011727.ref042","first-page":"1","article-title":"Free association transitions in models of cortical latching dynamics","volume":"10","author":"E. Russo","year":"2008","journal-title":"New Journal of Physics"},{"issue":"10","key":"pcbi.1011727.ref043","doi-asserted-by":"crossref","first-page":"2523","DOI":"10.1162\/NECO_a_00499","article-title":"Scaling laws of associative memory retrieval","volume":"25","author":"S. Romani","year":"2013","journal-title":"Neural computation"},{"key":"pcbi.1011727.ref044","doi-asserted-by":"crossref","first-page":"942","DOI":"10.1038\/nn.3431","article-title":"Temporal whitening by power-law adaptation in neocortical neurons","volume":"16","author":"C. Pozzorini","year":"2013","journal-title":"Nature Neuroscience"},{"key":"pcbi.1011727.ref045","volume-title":"Memory: A Contribution to Experimental Psychology","author":"H.E. Ebbinghaus"},{"key":"pcbi.1011727.ref046","doi-asserted-by":"crossref","first-page":"734","DOI":"10.1037\/0033-295X.103.4.734","article-title":"One hundred years of forgetting: a quantitative description of retention","volume":"103","author":"D.C. Rubin","year":"1996","journal-title":"Psychological Review"},{"issue":"1","key":"pcbi.1011727.ref047","doi-asserted-by":"crossref","first-page":"100","DOI":"10.1162\/neco.1994.6.1.100","article-title":"The role of constraints in Hebbian learning","volume":"6","author":"K.D. 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Quian Quiroga","year":"2019","journal-title":"Cell"},{"issue":"1","key":"pcbi.1011727.ref054","doi-asserted-by":"crossref","first-page":"13107","DOI":"10.1038\/s41598-022-16407-9","article-title":"Stochastic consolidation of lifelong memory","volume":"12","author":"N. Shaham","year":"2022","journal-title":"Scientific Reports"},{"key":"pcbi.1011727.ref055","doi-asserted-by":"crossref","first-page":"64","DOI":"10.3389\/fncom.2014.00064","article-title":"Memory consolidation from seconds to weeks: a three-stage neural network model with autonomous reinstatement dynamics","volume":"8","author":"F. Fiebig","year":"2014","journal-title":"Frontiers in Computational Neuroscience"},{"key":"pcbi.1011727.ref056","doi-asserted-by":"crossref","first-page":"e43717","DOI":"10.7554\/eLife.43717","article-title":"Self-organized reactivation maintains and reinforces memories despite synaptic turnover.","volume":"8","author":"M.J. 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