{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,8]],"date-time":"2026-04-08T08:56:03Z","timestamp":1775638563781,"version":"3.50.1"},"reference-count":53,"publisher":"Public Library of Science (PLoS)","issue":"6","license":[{"start":{"date-parts":[[2020,6,9]],"date-time":"2020-06-09T00:00:00Z","timestamp":1591660800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"DOI":"10.1371\/journal.pcbi.1007936","type":"journal-article","created":{"date-parts":[[2020,6,9]],"date-time":"2020-06-09T13:30:18Z","timestamp":1591709418000},"page":"e1007936","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":28,"title":["A functional spiking-neuron model of activity-silent working memory in humans based on calcium-mediated short-term synaptic plasticity"],"prefix":"10.1371","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0640-2145","authenticated-orcid":true,"given":"Matthijs","family":"Pals","sequence":"first","affiliation":[]},{"given":"Terrence C.","family":"Stewart","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9111-5847","authenticated-orcid":true,"given":"Elkan G.","family":"Aky\u00fcrek","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4493-8223","authenticated-orcid":true,"given":"Jelmer P.","family":"Borst","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2020,6,9]]},"reference":[{"key":"pcbi.1007936.ref001","first-page":"47","volume-title":"Psychol. Learn. Motiv","author":"AD Baddeley","year":"1974"},{"key":"pcbi.1007936.ref002","doi-asserted-by":"crossref","first-page":"20","DOI":"10.1016\/j.conb.2013.10.008","article-title":"Working models of working memory","volume":"25","author":"O Barak","year":"2014","journal-title":"Curr. Opin. Neurobiol"},{"issue":"6","key":"pcbi.1007936.ref003","doi-asserted-by":"crossref","first-page":"864","DOI":"10.1038\/nn.4546","article-title":"Dynamic hidden states underlying working-memory-guided behavior","volume":"20","author":"MJ Wolff","year":"2017","journal-title":"Nat. Neurosci"},{"issue":"3","key":"pcbi.1007936.ref004","doi-asserted-by":"crossref","first-page":"e1004060","DOI":"10.1371\/journal.pcbi.1004060","article-title":"How attention can create synaptic tags for the learning of working memories in sequential tasks","volume":"11","author":"JO Rombouts","year":"2015","journal-title":"PLoS Comput. Biol"},{"issue":"3997","key":"pcbi.1007936.ref005","doi-asserted-by":"crossref","first-page":"652","DOI":"10.1126\/science.173.3997.652","article-title":"Neuron Activity Related to Short-Term Memory","volume":"173","author":"JMJ Fuster","year":"1971","journal-title":"Science"},{"issue":"3","key":"pcbi.1007936.ref006","doi-asserted-by":"crossref","first-page":"477","DOI":"10.1016\/0896-6273(95)90304-6","article-title":"Cellular basis of working memory","volume":"14","author":"PS Goldman-Rakic","year":"1995","journal-title":"Neuron"},{"issue":"32","key":"pcbi.1007936.ref007","doi-asserted-by":"crossref","first-page":"7013","DOI":"10.1523\/JNEUROSCI.2485-17.2018","article-title":"Working Memory: Delay Activity, Yes! Persistent Activity","volume":"38","author":"M Lundqvist","year":"2018","journal-title":"Maybe Not. J. Neurosci"},{"key":"pcbi.1007936.ref008","doi-asserted-by":"crossref","first-page":"1","DOI":"10.7554\/eLife.09000","article-title":"Testing sensory evidence against mnemonic templates","volume":"4","author":"NE Myers","year":"2015","journal-title":"Elife"},{"issue":"2","key":"pcbi.1007936.ref009","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1016\/j.tics.2013.12.001","article-title":"Revisiting the role of persistent neural activity during working memory","volume":"18","author":"KK Sreenivasan","year":"2014","journal-title":"Trends Cogn. Sci."},{"issue":"4","key":"pcbi.1007936.ref010","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1038\/nn.3667","article-title":"Neural mechanisms of dual-task interference and cognitive capacity limitation in the prefrontal cortex","volume":"17","author":"K Watanabe","year":"2014","journal-title":"Nat. Neurosci"},{"key":"pcbi.1007936.ref011","doi-asserted-by":"crossref","first-page":"152","DOI":"10.1016\/j.neuron.2016.02.028","article-title":"Gamma and Beta Bursts Underlie Working Memory","volume":"90","author":"M Lundqvist","year":"2016","journal-title":"Neuron"},{"issue":"5869","key":"pcbi.1007936.ref012","doi-asserted-by":"crossref","first-page":"1543","DOI":"10.1126\/science.1150769","article-title":"Synaptic Theory of Working Memory","volume":"319","author":"G Mongillo","year":"2008","journal-title":"Science"},{"issue":"7","key":"pcbi.1007936.ref013","doi-asserted-by":"crossref","first-page":"394","DOI":"10.1016\/j.tics.2015.05.004","article-title":"\u2018Activity-silent\u2019 working memory in prefrontal cortex: a dynamic coding framework","volume":"19","author":"MG Stokes","year":"2015","journal-title":"Trends Cogn. Sci"},{"issue":"5","key":"pcbi.1007936.ref014","doi-asserted-by":"crossref","first-page":"1628","DOI":"10.1073\/pnas.1221572110","article-title":"Using Model-Based functional MRI to locate Working Memory Updates and Declarative Memory Retrievals in the Fronto-Parietal Network","volume":"110","author":"JP Borst","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"issue":"2","key":"pcbi.1007936.ref015","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1037\/a0018106","article-title":"The Problem State: A Cognitive Bottleneck in Multitasking","volume":"36","author":"JP Borst","year":"2010","journal-title":"J. Exp. Psychol. Learn. Mem. Cogn"},{"issue":"3","key":"pcbi.1007936.ref016","doi-asserted-by":"crossref","first-page":"411","DOI":"10.1037\/0278-7393.28.3.411","article-title":"Access to information in working memory: Exploring the focus of attention","volume":"28","author":"K Oberauer","year":"2002","journal-title":"J. Exp. Psychol. Learn. Mem. Cogn"},{"issue":"7","key":"pcbi.1007936.ref017","first-page":"327","article-title":"Different states in visual working memory: when it guides attention and when it does not","volume":"15","author":"CNL Olivers","year":"2011","journal-title":"Trends Cogn. Sci"},{"issue":"28","key":"pcbi.1007936.ref018","doi-asserted-by":"crossref","first-page":"9424","DOI":"10.1523\/JNEUROSCI.1875-10.2010","article-title":"Neuronal population coding of parametric working memory","volume":"30","author":"O Barak","year":"2010","journal-title":"J. Neurosci"},{"issue":"1","key":"pcbi.1007936.ref019","doi-asserted-by":"crossref","first-page":"355","DOI":"10.1146\/annurev.physiol.64.092501.114547","article-title":"Short-Term Synaptic Plasticity","volume":"64","author":"RS Zucker","year":"2002","journal-title":"Annu. Rev. Physiol"},{"issue":"10","key":"pcbi.1007936.ref020","doi-asserted-by":"crossref","first-page":"821","DOI":"10.1162\/089976698300017502","article-title":"Neural Networks with Dynamic Synapses","author":"M Tsodyks","year":"1998","journal-title":"Neural Comput"},{"issue":"4","key":"pcbi.1007936.ref021","doi-asserted-by":"crossref","first-page":"534","DOI":"10.1038\/nn1670","article-title":"Heterogeneity in the pyramidal network of the medial prefrontal cortex","volume":"9","author":"Y Wang","year":"2006","journal-title":"Nat. Neurosci"},{"issue":"3","key":"pcbi.1007936.ref022","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1016\/j.neuron.2017.02.047","article-title":"The Mechanisms and Functions of Synaptic Facilitation","volume":"94","author":"SL Jackman","year":"2017","journal-title":"Neuron"},{"key":"pcbi.1007936.ref023","doi-asserted-by":"crossref","first-page":"1","DOI":"10.7554\/eLife.09000","article-title":"Testing sensory evidence against mnemonic templates","volume":"4","author":"NE Myers","year":"2015","journal-title":"eLife"},{"issue":"123","key":"pcbi.1007936.ref024","first-page":"1","article-title":"Revealing hidden states in visual working memory using electroencephalography","volume":"9","author":"MJ Wolff","year":"2015","journal-title":"Front. Syst. Neurosci"},{"key":"pcbi.1007936.ref025","doi-asserted-by":"crossref","first-page":"48","DOI":"10.3389\/fninf.2013.00048","article-title":"Nengo: a Python tool for building large-scale functional brain models","volume":"7","author":"T Bekolay","year":"2014","journal-title":"Front. Neuroinformatics"},{"key":"pcbi.1007936.ref026","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780199794546.001.0001","volume-title":"How to Build a Brain: A Neural Architecture for Biological Cognition","author":"C Eliasmith","year":"2013"},{"issue":"6111","key":"pcbi.1007936.ref027","doi-asserted-by":"crossref","first-page":"1202","DOI":"10.1126\/science.1225266","article-title":"A large-scale model of the functioning brain","volume":"338","author":"C Eliasmith","year":"2012","journal-title":"Science"},{"key":"pcbi.1007936.ref028","first-page":"169","author":"T Bekolay","year":"2013"},{"issue":"1","key":"pcbi.1007936.ref029","doi-asserted-by":"crossref","first-page":"455","DOI":"10.1152\/jn.2002.88.1.455","article-title":"Spatial Structure and Symmetry of Simple-Cell Receptive Fields in Macaque Primary Visual Cortex","volume":"88","author":"DL Ringach","year":"2002","journal-title":"J. Neurophysiol"},{"issue":"6","key":"pcbi.1007936.ref030","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1152\/jn.1987.58.6.1233","article-title":"An Evaluation of the Two-Dimensional Gabor Filter Model of Simple Receptive Fields in Cat Striate Cortex","volume":"58","author":"JP Jones","year":"1987","journal-title":"J. Neurophysiol"},{"key":"pcbi.1007936.ref031","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1023\/A:1008837311948","article-title":"A model of visuospatial short-term memory in prefrontal cortex: recurrent network and cellular bistability","volume":"5","author":"M Camperi","year":"1998","journal-title":"J. Comput. Neurosci"},{"issue":"11","key":"pcbi.1007936.ref032","doi-asserted-by":"crossref","first-page":"1208","DOI":"10.1093\/cercor\/bhg101","article-title":"A Recurrent Network Model of Somatosensory Parametric Working Memory in the Prefrontal Cortex","volume":"13","author":"P Miller","year":"2003","journal-title":"Cereb. Cortex"},{"key":"pcbi.1007936.ref033","first-page":"320","volume-title":"Oxf. Handb. Comput. Math. Psychol","author":"TJ Palmeri","year":"2015"},{"key":"pcbi.1007936.ref034","doi-asserted-by":"crossref","DOI":"10.3389\/fnins.2012.00002","article-title":"Learning to Select Actions with Spiking Neurons in the Basal Ganglia","volume":"6","author":"TC Stewart","year":"2012","journal-title":"Front. Neurosci"},{"key":"pcbi.1007936.ref035","author":"J Fuster","year":"2015","edition":"5"},{"issue":"5","key":"pcbi.1007936.ref036","doi-asserted-by":"crossref","first-page":"1141","DOI":"10.1162\/jocn_a_00556","article-title":"Distributed and dynamic storage of working memory stimulus information in extrastriate cortex","volume":"26","author":"KK Sreenivasan","year":"2014","journal-title":"J. Cogn. Neurosci"},{"issue":"2","key":"pcbi.1007936.ref037","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1016\/j.neuron.2013.01.039","article-title":"Dynamic Coding for Cognitive Control in Prefrontal Cortex","volume":"78","author":"MG Stokes","year":"2013","journal-title":"Neuron"},{"issue":"2","key":"pcbi.1007936.ref038","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1038\/nrn1603","article-title":"Working memory in primate sensory systems","volume":"6","author":"T Pasternak","year":"2005","journal-title":"Nat. Rev. Neurosci"},{"key":"pcbi.1007936.ref039","doi-asserted-by":"crossref","first-page":"53","DOI":"10.1016\/j.intell.2013.10.003","article-title":"A spiking neural model applied to the study of human performance and cognitive decline on Raven\u2019s Advanced Progressive Matrices","volume":"42","author":"D Rasmussen","year":"2014","journal-title":"Intelligence"},{"key":"pcbi.1007936.ref040","doi-asserted-by":"crossref","first-page":"48","DOI":"10.3389\/fpsyg.2017.00099","article-title":"A Spiking Neuron Model of Word Associations for the Remote Associates Test","volume":"8","author":"I Kaji\u0107","year":"2017","journal-title":"Front. Psychol"},{"issue":"1","key":"pcbi.1007936.ref041","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1162\/jocn_a_00140","article-title":"Neural evidence for a distinction between short-term memory and the focus of attention","volume":"24","author":"JA Lewis-Peacock","year":"2012","journal-title":"J. Cogn. Neurosci"},{"key":"pcbi.1007936.ref042","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/S0079-7421(09)51002-X","volume-title":"Psychol. Learn. Motiv.","author":"K Oberauer","year":"2009"},{"key":"pcbi.1007936.ref043","article-title":"Attention and memory: An integrated framework","author":"N Cowan","year":"1995","journal-title":"New York: Oxford University Press"},{"issue":"3","key":"pcbi.1007936.ref044","doi-asserted-by":"crossref","first-page":"817","DOI":"10.1037\/0278-7393.27.3.817","article-title":"Working memory and focal attention","volume":"27","author":"B McElree","year":"2001","journal-title":"J. Exp. Psychol. Learn. Mem. Cogn"},{"key":"pcbi.1007936.ref045","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780195324259.001.0001","volume-title":"How Can the Human Mind Occur in the Physical Universe?","author":"JR Anderson","year":"2007"},{"key":"pcbi.1007936.ref046","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.cogpsych.2016.01.003","article-title":"Contrasting Single and Multi-Component Working-Memory Systems in Dual Tasking","volume":"86","author":"M Nijboer","year":"2016","journal-title":"Cognit. Psychol"},{"issue":"9","key":"pcbi.1007936.ref047","doi-asserted-by":"crossref","first-page":"e12966","DOI":"10.1371\/journal.pone.0012966","article-title":"The Neural Correlates of Problem States: Testing fMRI Predictions of a Computational Model of Multitasking","volume":"5","author":"JP Borst","year":"2010","journal-title":"PLoS ONE"},{"key":"pcbi.1007936.ref048","volume-title":"Proc CHI. Seoul","author":"JP Borst","year":"2015"},{"key":"pcbi.1007936.ref049","doi-asserted-by":"crossref","first-page":"5","DOI":"10.3389\/fnhum.2014.00005","article-title":"Multiple neural states of representation in short-term memory? It\u2019s a matter of attention","volume":"8","author":"JJ Larocque","year":"2014","journal-title":"Front. Hum. Neurosci"},{"issue":"14","key":"pcbi.1007936.ref050","doi-asserted-by":"crossref","first-page":"3667","DOI":"10.1523\/JNEUROSCI.4864-05.2006","article-title":"Higher-dimensional neurons explain the tuning and dynamics of working memory cells","volume":"26","author":"R Singh","year":"2006","journal-title":"J. Neurosci"},{"issue":"3","key":"pcbi.1007936.ref051","doi-asserted-by":"crossref","first-page":"621","DOI":"10.1162\/neco.2009.02-09-970","article-title":"Population models of temporal differentiation","volume":"22","author":"BP Tripp","year":"2010","journal-title":"Neural Comput"},{"key":"pcbi.1007936.ref052","volume-title":"Neural Engineering: Computation, Representation, and Dynamics in Neurobiological Systems","author":"C Eliasmith","year":"2002"},{"issue":"1","key":"pcbi.1007936.ref053","doi-asserted-by":"crossref","first-page":"195","DOI":"10.3758\/s13428-018-01193-y","article-title":"PsychoPy2: Experiments in behavior made easy","volume":"51","author":"J Peirce","year":"2019","journal-title":"Behav. Res. Methods"}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1007936","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,6,9]],"date-time":"2020-06-09T13:31:16Z","timestamp":1591709476000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1007936"}},"subtitle":[],"editor":[{"given":"Abigail","family":"Morrison","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2020,6,9]]},"references-count":53,"journal-issue":{"issue":"6","published-online":{"date-parts":[[2020,6,9]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1007936","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/823559","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2020,6,9]]}}}