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The probability of a behavioral response does not depend appreciably on the duration or intensity of a constant stimulation, whereas the response probability increases significantly upon injection of an irregular current. Biological mechanisms that can potentially suppress a constant input signal are present in the dynamics of both neurons and synapses and seem ideal candidates to explain these experimental findings. Here, we study a large network of integrate-and-fire neurons with several salient features of neuronal populations in the rat barrel cortex. The model includes cellular spike-frequency adaptation, experimentally constrained numbers and types of chemical synapses endowed with short-term plasticity, and gap junctions. Numerical simulations of this model indicate that cellular and synaptic adaptation mechanisms alone may not suffice to account for the experimental results if the local network activity is read out by an integrator. However, a circuit that approximates a differentiator can detect the single-cell stimulation with a reliability that barely depends on the length or intensity of the stimulus, but that increases when an irregular signal is used. This finding is in accordance with the experimental results obtained for the stimulation of a regularly-spiking excitatory cell.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1007831","type":"journal-article","created":{"date-parts":[[2021,2,8]],"date-time":"2021-02-08T21:22:37Z","timestamp":1612819357000},"page":"e1007831","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":18,"title":["A network model of the barrel cortex combined with a differentiator detector reproduces features of the behavioral response to single-neuron 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cortex function","volume":"103","author":"D Feldmeyer","year":"2013","journal-title":"Prog Neurobiol"},{"key":"pcbi.1007831.ref002","doi-asserted-by":"crossref","first-page":"20140204","DOI":"10.1098\/rstb.2014.0204","article-title":"What single-cell stimulation has told us about neural coding","volume":"370","author":"G Doron","year":"2015","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"issue":"6976","key":"pcbi.1007831.ref003","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1038\/nature02266","article-title":"Whisker movements evoked by stimulation of single pyramidal cells in rat motor cortex","volume":"427","author":"M Brecht","year":"2004","journal-title":"Nature"},{"issue":"7174","key":"pcbi.1007831.ref004","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1038\/nature06447","article-title":"Behavioural report of single neuron stimulation in somatosensory cortex","volume":"451","author":"AR 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Renart","year":"2010","journal-title":"Science"},{"issue":"9","key":"pcbi.1007831.ref017","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1038\/nrn3084","article-title":"Cortical state and attention","volume":"12","author":"KD Harris","year":"2011","journal-title":"Nat Rev Neurosci"},{"key":"pcbi.1007831.ref018","doi-asserted-by":"crossref","first-page":"258302","DOI":"10.1103\/PhysRevLett.121.258302","article-title":"Self-Consistent Correlations of Randomly Coupled Rotators in the Asynchronous State","volume":"121","author":"A van Meegen","year":"2018","journal-title":"Phys Rev Lett"},{"issue":"2","key":"pcbi.1007831.ref019","doi-asserted-by":"crossref","first-page":"023024","DOI":"10.1103\/PhysRevResearch.1.023024","article-title":"Theory of spike-train power spectra for multidimensional integrate-and-fire neurons","volume":"1","author":"S Vellmer","year":"2019","journal-title":"Phys Rev Research"},{"key":"pcbi.1007831.ref020","article-title":"Response nonlinearities in 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Ledoux","year":"2011","journal-title":"Front Comput Neurosci"},{"key":"pcbi.1007831.ref024","doi-asserted-by":"crossref","first-page":"011919","DOI":"10.1103\/PhysRevE.84.011919","article-title":"Rate response of neurons subject to fast or frozen noise: from stochastic and homogeneous to deterministic and heterogeneous populations","volume":"84","author":"AK Alijani","year":"2011","journal-title":"Phys Rev E"},{"key":"pcbi.1007831.ref025","doi-asserted-by":"crossref","first-page":"268301","DOI":"10.1103\/PhysRevLett.118.268301","article-title":"Optimal detection of a localized perturbation in random networks of integrate-and-fire neurons","volume":"118","author":"D Bernardi","year":"2017","journal-title":"Phys Rev Lett"},{"issue":"3","key":"pcbi.1007831.ref026","doi-asserted-by":"crossref","first-page":"032304","DOI":"10.1103\/PhysRevE.99.032304","article-title":"Detecting single-cell stimulation in a large network of integrate-and-fire neurons","volume":"99","author":"D Bernardi","year":"2019","journal-title":"Phys Rev E"},{"issue":"11","key":"pcbi.1007831.ref027","doi-asserted-by":"crossref","first-page":"3116","DOI":"10.1152\/jn.00917.2011","article-title":"Microcircuits of excitatory and inhibitory neurons in layer 2\/3 of mouse barrel cortex","volume":"107","author":"M Avermann","year":"2012","journal-title":"J Neurophysiol"},{"issue":"20","key":"pcbi.1007831.ref028","doi-asserted-by":"crossref","first-page":"6746","DOI":"10.1523\/JNEUROSCI.0305-14.2014","article-title":"Comparative strength and dendritic organization of thalamocortical and corticocortical synapses onto excitatory layer 4 neurons","volume":"34","author":"CE Schoonover","year":"2014","journal-title":"J Neurosci"},{"issue":"5","key":"pcbi.1007831.ref029","doi-asserted-by":"crossref","first-page":"2987","DOI":"10.1152\/jn.00283.2003","article-title":"Two dynamically distinct inhibitory networks in layer 4 of the neocortex","volume":"90","author":"M Beierlein","year":"2003","journal-title":"J Neurophysiol"},{"issue":"40","key":"pcbi.1007831.ref030","doi-asserted-by":"crossref","first-page":"16807","DOI":"10.1073\/pnas.1113648108","article-title":"Inhibitory interneurons in a cortical column form hot zones of inhibition in layers 2 and 5A","volume":"108","author":"HS Meyer","year":"2011","journal-title":"Proc Natl Acad Sci USA"},{"issue":"8","key":"pcbi.1007831.ref031","doi-asserted-by":"crossref","first-page":"e1004165","DOI":"10.1371\/journal.pcbi.1004165","article-title":"Experimentally verified parameter sets for modelling heterogeneous neocortical pyramidal-cell populations","volume":"11","author":"PM Harrison","year":"2015","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1007831.ref032","doi-asserted-by":"crossref","first-page":"904","DOI":"10.1038\/78809","article-title":"A network of electrically coupled interneurons drives synchronized inhibition in neocortex","volume":"3","author":"M Beierlein","year":"2000","journal-title":"Nat Neurosci"},{"issue":"8","key":"pcbi.1007831.ref033","doi-asserted-by":"crossref","first-page":"1068","DOI":"10.1038\/nn.3446","article-title":"Inhibition of inhibition in visual cortex: the logic of connections between molecularly distinct interneurons","volume":"16","author":"CK Pfeffer","year":"2013","journal-title":"Nat Neurosci"},{"issue":"6757","key":"pcbi.1007831.ref034","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1038\/47029","article-title":"A network of fast-spiking cells in the neocortex connected by electrical synapses","volume":"402","author":"M Galarreta","year":"1999","journal-title":"Nature"},{"issue":"12","key":"pcbi.1007831.ref035","doi-asserted-by":"crossref","first-page":"3446","DOI":"10.1111\/ejn.14267","article-title":"Interneuronal gap junctions increase synchrony and robustness of hippocampal ripple oscillations","volume":"48","author":"A Holzbecher","year":"2018","journal-title":"Eur J 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inhibition during sparse activity in the somatosensory cortex","volume":"10","author":"C Kapfer","year":"2007","journal-title":"Nat Neurosci"},{"key":"pcbi.1007831.ref043","doi-asserted-by":"crossref","first-page":"3869","DOI":"10.1093\/cercor\/bhx094","article-title":"Layer-Dependent Short-Term Synaptic Plasticity Between Excitatory Neurons in the C2 Barrel Column of Mouse Primary Somatosensory Cortex","volume":"27","author":"S Lefort","year":"2017","journal-title":"Cereb Cortex"},{"key":"pcbi.1007831.ref044","doi-asserted-by":"crossref","first-page":"719","DOI":"10.1073\/pnas.94.2.719","article-title":"The neural code between neocortical pyramidal neurons depends on neurotransmitter release probability","volume":"94","author":"MV Tsodyks","year":"1997","journal-title":"Proc Natl Acad Sci USA"},{"key":"pcbi.1007831.ref045","doi-asserted-by":"crossref","first-page":"821","DOI":"10.1162\/089976698300017502","article-title":"Neural networks with dynamic synapses","volume":"10","author":"M Tsodyks","year":"1998","journal-title":"Neural Comput"},{"key":"pcbi.1007831.ref046","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1523\/JNEUROSCI.3474-11.2012","article-title":"Neural correlation is stimulus modulated by feedforward inhibitory circuitry","volume":"32","author":"JW Middleton","year":"2012","journal-title":"J Neurosci"},{"key":"pcbi.1007831.ref047","doi-asserted-by":"crossref","first-page":"607","DOI":"10.1038\/nn.3051","article-title":"Unique functional properties of somatostatin-expressing GABAergic neurons in mouse barrel cortex","volume":"15","author":"LJ Gentet","year":"2012","journal-title":"Nat Neurosci"},{"key":"pcbi.1007831.ref048","article-title":"Psychophysical characterization of single neuron stimulation effects in rat barrel cortex","author":"G Doron","year":"2012","journal-title":"Humboldt-Universit\u00e4t zu Berlin, Mathematisch-Naturwissenschaftliche 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labeling of a single neuron recorded extracellularly","volume":"170","author":"D Pinault","year":"1994","journal-title":"Neurosci Lett"},{"key":"pcbi.1007831.ref059","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1152\/jn.1994.72.1.392","article-title":"OFF response transformations in the whisker\/barrel system","volume":"72","author":"HT Kyriazi","year":"1994","journal-title":"J Neurophysiol"},{"key":"pcbi.1007831.ref060","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1152\/jn.1996.76.1.130","article-title":"Spatial gradients and inhibitory summation in the rat whisker barrel system","volume":"76","author":"JC Brumberg","year":"1996","journal-title":"J Neurophysiol"},{"key":"pcbi.1007831.ref061","doi-asserted-by":"crossref","first-page":"1808","DOI":"10.1152\/jn.1999.82.4.1808","article-title":"Cortical columnar processing in the rat whisker-to-barrel system","volume":"82","author":"JC Brumberg","year":"1999","journal-title":"J 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Cortex"},{"key":"pcbi.1007831.ref071","doi-asserted-by":"crossref","first-page":"420","DOI":"10.1038\/nn1643","article-title":"The tempotron: a neuron that learns spike timing-based decisions","volume":"9","author":"R G\u00fctig","year":"2006","journal-title":"Nat Neurosci"},{"issue":"26","key":"pcbi.1007831.ref072","doi-asserted-by":"crossref","first-page":"268104","DOI":"10.1103\/PhysRevLett.105.268104","article-title":"Dynamical entropy production in spiking neuron networks in the balanced state","volume":"105","author":"M Monteforte","year":"2010","journal-title":"Phys Rev Lett"},{"issue":"4","key":"pcbi.1007831.ref073","first-page":"041007","article-title":"Dynamic flux tubes form reservoirs of stability in neuronal circuits","volume":"2","author":"M Monteforte","year":"2012","journal-title":"Phys Rev X"},{"key":"pcbi.1007831.ref074","doi-asserted-by":"crossref","first-page":"e1005258","DOI":"10.1371\/journal.pcbi.1005258","article-title":"Encoding in Balanced Networks: Revisiting Spike Patterns and Chaos in Stimulus-Driven Systems","volume":"12","author":"G Lajoie","year":"2016","journal-title":"PLoS Comput Biol"},{"issue":"5","key":"pcbi.1007831.ref075","doi-asserted-by":"crossref","first-page":"3024","DOI":"10.1152\/jn.00245.2003","article-title":"Spatiotemporal effects of microstimulation in rat neocortex: a parametric study using multielectrode recordings","volume":"90","author":"S Butovas","year":"2003","journal-title":"J Neurophysiol"},{"issue":"3","key":"pcbi.1007831.ref076","doi-asserted-by":"crossref","first-page":"1227","DOI":"10.1152\/jn.01170.2005","article-title":"Effects of electrically coupled inhibitory networks on local neuronal responses to intracortical microstimulation","volume":"96","author":"S Butovas","year":"2006","journal-title":"J Neurophysiol"},{"issue":"4","key":"pcbi.1007831.ref077","doi-asserted-by":"crossref","first-page":"508","DOI":"10.1016\/j.neuron.2009.07.016","article-title":"Direct activation of sparse, distributed populations of cortical neurons by electrical microstimulation","volume":"63","author":"MH Histed","year":"2009","journal-title":"Neuron"},{"issue":"6","key":"pcbi.1007831.ref078","doi-asserted-by":"crossref","first-page":"066016","DOI":"10.1088\/1741-2560\/10\/6\/066016","article-title":"Computational modeling of direct neuronal recruitment during intracortical microstimulation in somatosensory cortex","volume":"10","author":"CK Overstreet","year":"2013","journal-title":"J Neural Eng"},{"key":"pcbi.1007831.ref079","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1016\/j.neuron.2016.06.033","article-title":"GABAergic Interneurons in the Neocortex: From Cellular Properties to Circuits","volume":"91","author":"R Tremblay","year":"2016","journal-title":"Neuron"},{"issue":"5","key":"pcbi.1007831.ref080","doi-asserted-by":"crossref","first-page":"e1006999","DOI":"10.1371\/journal.pcbi.1006999","article-title":"Amplifying the redistribution of somato-dendritic inhibition by the interplay of three interneuron types","volume":"15","author":"L Hert\u00e4g","year":"2019","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1007831.ref081","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1007\/s10827-008-0135-1","article-title":"Correlations in spiking neuronal networks with distance dependent connections","volume":"27","author":"B Kriener","year":"2009","journal-title":"J Comput Neurosci"},{"issue":"3","key":"pcbi.1007831.ref082","doi-asserted-by":"crossref","first-page":"e1002408","DOI":"10.1371\/journal.pcbi.1002408","article-title":"Impact of network structure and cellular response on spike time correlations","volume":"8","author":"J Trousdale","year":"2012","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1007831.ref083","doi-asserted-by":"crossref","first-page":"031916","DOI":"10.1103\/PhysRevE.85.031916","article-title":"Recurrent interactions in spiking networks with arbitrary topology","volume":"85","author":"V Pernice","year":"2012","journal-title":"Phys Rev E"},{"issue":"1","key":"pcbi.1007831.ref084","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1038\/nn.4433","article-title":"The spatial structure of correlated neuronal variability","volume":"20","author":"R Rosenbaum","year":"2017","journal-title":"Nat Neurosci"},{"key":"pcbi.1007831.ref085","doi-asserted-by":"crossref","first-page":"023002","DOI":"10.1088\/1367-2630\/15\/2\/023002","article-title":"Echoes in correlated neural systems","volume":"15","author":"M Helias","year":"2013","journal-title":"New J Phys"},{"key":"pcbi.1007831.ref086","doi-asserted-by":"crossref","first-page":"139","DOI":"10.3389\/fncom.2013.00139","article-title":"Correlations in background activity control persistent state stability and allow execution of working memory tasks","volume":"7","author":"M Dipoppa","year":"2013","journal-title":"Front Comput Neurosci"},{"issue":"1","key":"pcbi.1007831.ref087","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1371\/journal.pcbi.1003428","article-title":"The Correlation Structure of Local Neuronal Networks Intrinsically Results from Recurrent Dynamics","volume":"10","author":"M Helias","year":"2014","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1007831.ref088","doi-asserted-by":"crossref","first-page":"e1005534","DOI":"10.1371\/journal.pcbi.1005534","article-title":"Locking of correlated neural activity to ongoing oscillations","volume":"13","author":"T K\u00fchn","year":"2017","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1007831.ref089","doi-asserted-by":"crossref","first-page":"e1005497","DOI":"10.1371\/journal.pcbi.1005497","article-title":"Robust information propagation through noisy neural circuits","volume":"13","author":"J Zylberberg","year":"2017","journal-title":"PLoS Comput Biol"},{"key":"pcbi.1007831.ref090","doi-asserted-by":"crossref","DOI":"10.1017\/CBO9781107447615","volume-title":"Neuronal dynamics: From single neurons to networks and models of cognition","author":"W Gerstner","year":"2014"},{"issue":"2","key":"pcbi.1007831.ref091","doi-asserted-by":"crossref","first-page":"301","DOI":"10.1016\/j.neuron.2008.12.020","article-title":"The excitatory neuronal network of the C2 barrel column in mouse primary somatosensory cortex","volume":"61","author":"S Lefort","year":"2009","journal-title":"Neuron"},{"key":"pcbi.1007831.ref092","first-page":"13260","volume":"31","author":"AM Packer","year":"2011","journal-title":"Dense, unspecific connectivity of neocortical parvalbumin-positive interneurons: a canonical microcircuit for inhibition? J Neurosci"},{"issue":"3","key":"pcbi.1007831.ref093","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1093\/cercor\/bht263","article-title":"A barrel-related interneuron in layer 4 of rat somatosensory cortex with a high intrabarrel connectivity","volume":"25","author":"C Koelbl","year":"2015","journal-title":"Cereb Cortex"},{"issue":"10","key":"pcbi.1007831.ref094","doi-asserted-by":"crossref","first-page":"4142","DOI":"10.1523\/JNEUROSCI.22-10-04142.2002","article-title":"The spatial dimensions of electrically coupled networks of interneurons in the neocortex","volume":"22","author":"Y Amitai","year":"2002","journal-title":"J Neurosci"},{"issue":"3","key":"pcbi.1007831.ref095","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1023\/A:1023265027714","article-title":"Dynamics of spiking neurons connected by both inhibitory and electrical coupling","volume":"14","author":"TJ Lewis","year":"2003","journal-title":"J Comput Neurosci"},{"issue":"3","key":"pcbi.1007831.ref096","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1007\/s10827-008-0117-3","article-title":"Synchronization properties of networks of electrically coupled neurons in the presence of noise and heterogeneities","volume":"26","author":"S Ostojic","year":"2009","journal-title":"J Comput Neurosci"},{"issue":"4","key":"pcbi.1007831.ref097","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1038\/73936","article-title":"Proximally targeted GABAergic synapses and gap junctions synchronize cortical interneurons","volume":"3","author":"G Tam\u00e1s","year":"2000","journal-title":"Nat Neurosci"},{"issue":"4","key":"pcbi.1007831.ref098","doi-asserted-by":"crossref","first-page":"1219","DOI":"10.1523\/JNEUROSCI.4727-04.2006","article-title":"Barrel cortex microcircuits: thalamocortical feedforward inhibition in spiny stellate cells is mediated by a small number of fast-spiking interneurons","volume":"26","author":"QQ Sun","year":"2006","journal-title":"J Neurosci"},{"key":"pcbi.1007831.ref099","doi-asserted-by":"crossref","first-page":"5323","DOI":"10.1073\/pnas.95.9.5323","article-title":"Differential signaling via the same axon of neocortical pyramidal neurons","volume":"95","author":"H Markram","year":"1998","journal-title":"Proc Natl Acad Sci USA"},{"issue":"17","key":"pcbi.1007831.ref100","doi-asserted-by":"crossref","first-page":"178102","DOI":"10.1103\/PhysRevLett.105.178102","article-title":"Firing-rate response of a neuron receiving excitatory and inhibitory synaptic shot noise","volume":"105","author":"MJE Richardson","year":"2010","journal-title":"Phys Rev Lett"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1007831","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2021,2,19]],"date-time":"2021-02-19T00:00:00Z","timestamp":1613692800000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1007831","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,2,20]],"date-time":"2021-02-20T00:33:50Z","timestamp":1613781230000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1007831"}},"subtitle":[],"editor":[{"given":"Jonathan","family":"Rubin","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2021,2,8]]},"references-count":100,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2021,2,8]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1007831","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2020.03.30.016261","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,2,8]]}}}