{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,3,20]],"date-time":"2026-03-20T05:00:08Z","timestamp":1773982808419,"version":"3.50.1"},"reference-count":62,"publisher":"Public Library of Science (PLoS)","issue":"7","license":[{"start":{"date-parts":[[2014,7,31]],"date-time":"2014-07-31T00:00:00Z","timestamp":1406764800000},"content-version":"unspecified","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.1003764","type":"journal-article","created":{"date-parts":[[2014,7,31]],"date-time":"2014-07-31T19:38:59Z","timestamp":1406835539000},"page":"e1003764","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":19,"title":["Dendritic Nonlinearities Reduce Network Size Requirements and Mediate ON and OFF States of Persistent Activity in a PFC Microcircuit Model"],"prefix":"10.1371","volume":"10","author":[{"given":"Athanasia","family":"Papoutsi","sequence":"first","affiliation":[]},{"given":"Kyriaki","family":"Sidiropoulou","sequence":"additional","affiliation":[]},{"given":"Panayiota","family":"Poirazi","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2014,7,31]]},"reference":[{"key":"ref1","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1016\/j.neuron.2009.03.020","article-title":"Reading the book of memory: sparse sampling versus dense mapping of connectomes","volume":"62","author":"HS Seung","year":"2009","journal-title":"Neuron"},{"key":"ref2","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.tins.2011.02.007","article-title":"Dissecting functional connectivity of neuronal microcircuits: experimental and theoretical insights","volume":"34","author":"S Feldt","year":"2011","journal-title":"Trends Neurosci"},{"key":"ref3","doi-asserted-by":"crossref","first-page":"87","DOI":"10.1038\/nature09880","article-title":"Functional specificity of local synaptic connections in neocortical networks","volume":"473","author":"H Ko","year":"2011","journal-title":"Nature"},{"key":"ref4","doi-asserted-by":"crossref","first-page":"1043","DOI":"10.1016\/j.neuron.2010.11.029","article-title":"An embedded subnetwork of highly active neurons in the neocortex","volume":"68","author":"L Yassin","year":"2010","journal-title":"Neuron"},{"key":"ref5","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1038\/nn.2337","article-title":"Replay of rule-learning related neural patterns in the prefrontal cortex during sleep","volume":"12","author":"A Peyrache","year":"2009","journal-title":"Nat Neurosci"},{"key":"ref6","doi-asserted-by":"crossref","first-page":"1","DOI":"10.3389\/fnana.2013.00001","article-title":"Computing the size and number of neuronal clusters in local circuits","volume":"7","author":"R Perin","year":"2013","journal-title":"Front Neuroanat"},{"key":"ref7","doi-asserted-by":"crossref","first-page":"11186","DOI":"10.1523\/JNEUROSCI.1921-08.2008","article-title":"Firing-pattern-dependent specificity of cortical excitatory feed-forward subnetworks","volume":"28","author":"T Otsuka","year":"2008","journal-title":"J Neurosci"},{"key":"ref8","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"},{"key":"ref9","doi-asserted-by":"crossref","first-page":"161","DOI":"10.3389\/fncir.2013.00161","article-title":"Induction and modulation of persistent activity in a layer V PFC microcircuit model","volume":"7","author":"A Papoutsi","year":"2013","journal-title":"Front Neural Circuits"},{"key":"ref10","doi-asserted-by":"crossref","first-page":"736","DOI":"10.1016\/j.neuron.2012.12.032","article-title":"NMDA Receptors Subserve Persistent Neuronal Firing during Working Memory in Dorsolateral Prefrontal Cortex","volume":"77","author":"M Wang","year":"2013","journal-title":"Neuron"},{"key":"ref11","doi-asserted-by":"crossref","first-page":"9587","DOI":"10.1523\/JNEUROSCI.19-21-09587.1999","article-title":"Synaptic basis of cortical persistent activity: the importance of NMDA receptors to working memory","volume":"19","author":"X-J Wang","year":"1999","journal-title":"J Neurosci"},{"key":"ref12","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1038\/nn.2245","article-title":"Dopamine modulates an mGluR5-mediated depolarization underlying prefrontal persistent activity","volume":"12","author":"K Sidiropoulou","year":"2009","journal-title":"Nat Neurosci"},{"key":"ref13","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1023\/A:1012837415096","article-title":"Turning on and off with excitation: the role of spike-timing asynchrony and synchrony in sustained neural activity","volume":"11","author":"BS Gutkin","year":"2001","journal-title":"J Comput Neurosci"},{"key":"ref14","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1016\/j.neuroscience.2005.06.011","article-title":"Computational and in vitro studies of persistent activity: edging towards cellular and synaptic mechanisms of working memory","volume":"139","author":"A Compte","year":"2006","journal-title":"Neuroscience"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"910","DOI":"10.1093\/cercor\/10.9.910","article-title":"Synaptic mechanisms and network dynamics underlying spatial working memory in a cortical network model","volume":"10","author":"A Compte","year":"2000","journal-title":"Cereb Cortex"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"173","DOI":"10.1038\/nature01171","article-title":"Graded persistent activity in entorhinal cortex neurons","volume":"420","author":"AV Egorov","year":"2002","journal-title":"Nature"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"10333","DOI":"10.1073\/pnas.0500717102","article-title":"Synaptic mechanisms of persistent reverberatory activity in neuronal networks","volume":"102","author":"P-M Lau","year":"2005","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref18","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1113\/jphysiol.2007.142315","article-title":"Voltage and calcium transients in basal dendrites of the rat prefrontal cortex","volume":"585","author":"BA Milojkovic","year":"2007","journal-title":"J Physiol"},{"key":"ref19","doi-asserted-by":"crossref","first-page":"16435","DOI":"10.1523\/JNEUROSCI.2777-11.2011","article-title":"Glutamate spillover promotes the generation of NMDA spikes","volume":"31","author":"JR Chalifoux","year":"2011","journal-title":"J Neurosci"},{"key":"ref20","doi-asserted-by":"crossref","first-page":"334","DOI":"10.3389\/fphys.2012.00334","article-title":"Extrasynaptic glutamate receptor activation as cellular bases for dynamic range compression in pyramidal neurons","volume":"3","author":"KD Oikonomou","year":"2012","journal-title":"Front Physiol"},{"key":"ref21","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1038\/35036228","article-title":"The prefrontal cortex and cognitive control","volume":"1","author":"EK Miller","year":"2000","journal-title":"Nat Rev Neurosci"},{"key":"ref22","doi-asserted-by":"crossref","first-page":"7513","DOI":"10.1523\/JNEUROSCI.6162-08.2009","article-title":"Distinct roles of GABA(A) and GABA(B) receptors in balancing and terminating persistent cortical activity","volume":"29","author":"EO Mann","year":"2009","journal-title":"J Neurosci"},{"key":"ref23","doi-asserted-by":"crossref","first-page":"e1002489","DOI":"10.1371\/journal.pcbi.1002489","article-title":"Predictive Features of Persistent Activity Emergence in Regular Spiking and Intrinsic Bursting Model Neurons","volume":"8","author":"K Sidiropoulou","year":"2012","journal-title":"PLoS Comput Biol"},{"key":"ref24","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1073\/pnas.98.1.301","article-title":"Dopamine D1\/D5 receptor modulation of excitatory synaptic inputs to layer V prefrontal cortex neurons","volume":"98","author":"JK Seamans","year":"2001","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref25","doi-asserted-by":"crossref","first-page":"452","DOI":"10.1093\/cercor\/11.5.452","article-title":"D(1) dopamine receptors potentiate nmda-mediated excitability increase in layer V prefrontal cortical pyramidal neurons","volume":"11","author":"J Wang","year":"2001","journal-title":"Cereb Cortex"},{"key":"ref26","doi-asserted-by":"crossref","first-page":"422","DOI":"10.1016\/j.neuron.2010.01.006","article-title":"Membrane potential dynamics of GABAergic neurons in the barrel cortex of behaving mice","volume":"65","author":"LJ Gentet","year":"2010","journal-title":"Neuron"},{"key":"ref27","doi-asserted-by":"crossref","first-page":"1160","DOI":"10.1016\/j.neuron.2011.02.022","article-title":"Synaptic mechanisms underlying sparse coding of active touch","volume":"69","author":"S Crochet","year":"2011","journal-title":"Neuron"},{"key":"ref28","doi-asserted-by":"crossref","first-page":"16791","DOI":"10.1073\/pnas.0804318105","article-title":"A specialized NMDA receptor function in layer 5 recurrent microcircuitry of the adult rat prefrontal cortex","volume":"105","author":"H Wang","year":"2008","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref29","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1093\/cercor\/bhh107","article-title":"Post-pubertal emergence of prefrontal cortical up states induced by D1-NMDA co-activation","volume":"15","author":"KY Tseng","year":"2005","journal-title":"Cereb Cortex"},{"key":"ref30","doi-asserted-by":"crossref","first-page":"288","DOI":"10.1038\/nature01616","article-title":"Turning on and off recurrent balanced cortical activity","volume":"423","author":"Y Shu","year":"2003","journal-title":"Nature"},{"key":"ref31","doi-asserted-by":"crossref","first-page":"7298","DOI":"10.1073\/pnas.0800360105","article-title":"Hyperpolarization-activated graded persistent activity in the prefrontal cortex","volume":"105","author":"M Winograd","year":"2008","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1146\/annurev-neuro-062111-150343","article-title":"Active properties of neocortical pyramidal neuron dendrites","volume":"36","author":"G Major","year":"2013","journal-title":"Annu Rev Neurosci"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"1219","DOI":"10.1093\/cercor\/bhg104","article-title":"Persistent cortical activity: mechanisms of generation and effects on neuronal excitability","volume":"13","author":"DA McCormick","year":"2003","journal-title":"Cereb Cortex"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"331","DOI":"10.1152\/jn.1989.61.2.331","article-title":"Mnemonic coding of visual space in the monkey&apos;s dorsolateral prefrontal cortex","volume":"61","author":"S Funahashi","year":"1989","journal-title":"J Neurophysiol"},{"key":"ref35","doi-asserted-by":"crossref","first-page":"5154","DOI":"10.1523\/JNEUROSCI.16-16-05154.1996","article-title":"Neural mechanisms of visual working memory in prefrontal cortex of the macaque","volume":"16","author":"EK Miller","year":"1996","journal-title":"J Neurosci"},{"key":"ref36","doi-asserted-by":"crossref","first-page":"7737","DOI":"10.1523\/JNEUROSCI.1347-08.2008","article-title":"Complementary contributions of prefrontal neuron classes in abstract numerical categorization","volume":"28","author":"I Diester","year":"2008","journal-title":"J Neurosci"},{"key":"ref37","doi-asserted-by":"crossref","first-page":"771","DOI":"10.1152\/jn.00070.2003","article-title":"The NMDA-to-AMPA ratio at synapses onto layer 2\/3 pyramidal neurons is conserved across prefrontal and visual cortices","volume":"90","author":"CIO Myme","year":"2003","journal-title":"J Neurophysiol"},{"key":"ref38","doi-asserted-by":"crossref","first-page":"1016","DOI":"10.1038\/nm1478","article-title":"Pathophysiologically based treatment interventions in schizophrenia","volume":"12","author":"DA Lewis","year":"2006","journal-title":"Nat Med"},{"key":"ref39","doi-asserted-by":"crossref","first-page":"1306","DOI":"10.1038\/nn.3492","article-title":"Balanced cortical microcircuitry for maintaining information in working memory","volume":"16","author":"S Lim","year":"2013","journal-title":"Nat Neurosci"},{"key":"ref40","doi-asserted-by":"crossref","first-page":"285","DOI":"10.1038\/35005094","article-title":"NMDA spikes in basal dendrites of cortical pyramidal neurons","volume":"404","author":"J Schiller","year":"2000","journal-title":"Nature"},{"key":"ref41","doi-asserted-by":"crossref","first-page":"2584","DOI":"10.1152\/jn.00011.2008","article-title":"Spatiotemporally graded NMDA spike\/plateau potentials in basal dendrites of neocortical pyramidal neurons","volume":"99","author":"G Major","year":"2008","journal-title":"J Neurophysiol"},{"key":"ref42","doi-asserted-by":"crossref","first-page":"756","DOI":"10.1126\/science.1171958","article-title":"Synaptic integration in tuft dendrites of layer 5 pyramidal neurons: a new unifying principle","volume":"325","author":"ME Larkum","year":"2009","journal-title":"Science"},{"key":"ref43","first-page":"1","article-title":"NMDA spikes enhance action potential generation during sensory input","author":"LM Palmer","year":"2014","journal-title":"Nat Neurosci"},{"key":"ref44","doi-asserted-by":"crossref","first-page":"1137","DOI":"10.1038\/nature07663","article-title":"Dendritic encoding of sensory stimuli controlled by deep cortical interneurons","volume":"457","author":"M Murayama","year":"2009","journal-title":"Nature"},{"key":"ref45","doi-asserted-by":"crossref","first-page":"115","DOI":"10.1038\/nature12600","article-title":"Dendritic spikes enhance stimulus selectivity in cortical neurons in vivo","volume":"503","author":"SL Smith","year":"2013","journal-title":"Nature"},{"key":"ref46","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1038\/nature11601","article-title":"Nonlinear dendritic integration of sensory and motor input during an active sensing task","volume":"492","author":"N Xu","year":"2012","journal-title":"Nature"},{"key":"ref47","doi-asserted-by":"crossref","first-page":"e1002362","DOI":"10.1371\/journal.pcbi.1002362","article-title":"Minimal size of cell assemblies coordinated by gamma oscillations","volume":"8","author":"C B\u00f6rgers","year":"2012","journal-title":"PLoS Comput Biol"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"10321","DOI":"10.1523\/JNEUROSCI.1703-09.2009","article-title":"Spatial profile and differential recruitment of GABAB modulate oscillatory activity in auditory cortex","volume":"29","author":"AM Oswald","year":"2009","journal-title":"J Neurosci"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"1498","DOI":"10.1038\/nn.3220","article-title":"Slow dynamics and high variability in balanced cortical networks with clustered connections","volume":"15","author":"A Litwin-Kumar","year":"2012","journal-title":"Nat Neurosci"},{"key":"ref50","doi-asserted-by":"crossref","first-page":"1027","DOI":"10.1038\/79848","article-title":"Cellular and network mechanisms of rhythmic recurrent activity in neocortex","volume":"3","author":"MV Sanchez-Vives","year":"2000","journal-title":"Nat Neurosci"},{"key":"ref51","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1126\/science.1169957","article-title":"Burst spiking of a single cortical neuron modifies global brain state","volume":"324","author":"CY Li","year":"2009","journal-title":"Science"},{"key":"ref52","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1016\/j.neures.2010.02.011","article-title":"Rule-dependent anticipatory activity in prefrontal neurons","volume":"67","author":"M Yamada","year":"2010","journal-title":"Neurosci Res"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1111\/j.1749-6632.2011.06270.x","article-title":"The orbitofrontal cortex, predicted value, and choice","volume":"1239","author":"BW Balleine","year":"2011","journal-title":"Ann N Y Acad Sci"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"6154","DOI":"10.1523\/JNEUROSCI.0581-09.2009","article-title":"The role of the monkey dorsal pontine nuclei in goal-directed eye and hand movements","volume":"29","author":"K Tziridis","year":"2009","journal-title":"J Neurosci"},{"key":"ref55","doi-asserted-by":"crossref","first-page":"424","DOI":"10.1523\/JNEUROSCI.1854-12.2013","article-title":"NMDA and GABAB (KIR) conductances: the \u201cperfect couple\u201d for bistability","volume":"33","author":"H Sanders","year":"2013","journal-title":"J Neurosci"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1016\/j.neuron.2012.09.039","article-title":"NMDA receptors control cue-outcome selectivity and plasticity of orbitofrontal firing patterns during associative stimulus-reward learning","volume":"76","author":"M Van Wingerden","year":"2012","journal-title":"Neuron"},{"key":"ref57","doi-asserted-by":"crossref","first-page":"4535","DOI":"10.1523\/JNEUROSCI.5297-05.2006","article-title":"Neocortical network activity in vivo is generated through a dynamic balance of excitation and inhibition","volume":"26","author":"B Haider","year":"2006","journal-title":"J Neurosci"},{"key":"ref58","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1177\/107385840100700207","article-title":"NEURON: a tool for neuroscientists","volume":"7","author":"ML Hines","year":"2001","journal-title":"Neuroscientist"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/S0301-0082(97)00070-1","article-title":"An ultrastructural study of the neural circuit between the prefrontal cortex and the mediodorsal nucleus of the thalamus","volume":"54","author":"M Kuroda","year":"1998","journal-title":"Prog Neurobiol"},{"key":"ref60","doi-asserted-by":"crossref","first-page":"868","DOI":"10.1038\/nature03252","article-title":"Excitatory cortical neurons form fine-scale functional networks","volume":"433","author":"Y Yoshimura","year":"2005","journal-title":"Nature"},{"key":"ref61","doi-asserted-by":"crossref","first-page":"1142","DOI":"10.1038\/nature07709","article-title":"The subcellular organization of neocortical excitatory connections","volume":"457","author":"L Petreanu","year":"2009","journal-title":"Nature"},{"key":"ref62","doi-asserted-by":"crossref","first-page":"1477","DOI":"10.1016\/j.neuron.2013.10.059","article-title":"Membrane potential dynamics of neocortical projection neurons driving target-specific signals","volume":"80","author":"T Yamashita","year":"2013","journal-title":"Neuron"}],"container-title":["PLoS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pcbi.1003764","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,5,9]],"date-time":"2020-05-09T01:53:52Z","timestamp":1588989232000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1003764"}},"subtitle":[],"editor":[{"given":"Aldo A.","family":"Faisal","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2014,7,31]]},"references-count":62,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2014,7,31]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1003764","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2014,7,31]]}}}