{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T03:45:44Z","timestamp":1793504744911,"version":"4.5.0"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1012723","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2025,1,16]],"date-time":"2025-01-16T00:00:00Z","timestamp":1736985600000}}],"reference-count":96,"publisher":"Public Library of Science (PLoS)","issue":"1","license":[{"start":{"date-parts":[[2025,1,6]],"date-time":"2025-01-06T00:00:00Z","timestamp":1736121600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100010661","name":"Horizon 2020 Framework Programme","doi-asserted-by":"publisher","award":["860563"],"award-info":[{"award-number":["860563"]}],"id":[{"id":"10.13039\/100010661","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100018693","name":"HORIZON EUROPE Framework Programme","doi-asserted-by":"publisher","award":["101057655"],"award-info":[{"award-number":["101057655"]}],"id":[{"id":"10.13039\/100018693","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100018693","name":"HORIZON EUROPE Framework Programme","doi-asserted-by":"publisher","award":["101058516"],"award-info":[{"award-number":["101058516"]}],"id":[{"id":"10.13039\/100018693","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100018703","name":"HORIZON EUROPE European Innovation Council","doi-asserted-by":"publisher","award":["101058240"],"award-info":[{"award-number":["101058240"]}],"id":[{"id":"10.13039\/100018703","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100010661","name":"Horizon 2020 Framework Programme","doi-asserted-by":"publisher","award":["871767"],"award-info":[{"award-number":["871767"]}],"id":[{"id":"10.13039\/100010661","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100010661","name":"Horizon 2020 Framework Programme","doi-asserted-by":"publisher","award":["860563"],"award-info":[{"award-number":["860563"]}],"id":[{"id":"10.13039\/100010661","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>Although the primary function of excitatory-inhibitory (E-I) homeostasis is the maintenance of mean firing rates, the conjugation of multiple homeostatic mechanisms is thought to be pivotal to ensuring edge-of-bifurcation dynamics in cortical circuits. However, computational studies on E-I homeostasis have focused solely on the plasticity of inhibition, neglecting the impact of different modes of E-I homeostasis on cortical dynamics. Therefore, we investigate how the diverse mechanisms of E-I homeostasis employed by cortical networks shape oscillations and edge-of-bifurcation dynamics. Using the Wilson-Cowan model, we explore how distinct modes of E-I homeostasis maintain stable firing rates in models with varying levels of input and how it affects circuit dynamics. Our results confirm that E-I homeostasis can be leveraged to control edge-of-bifurcation dynamics and that some modes of homeostasis maintain mean firing rates under higher levels of input by modulating the distance to the bifurcation. Additionally, relying on multiple modes of homeostasis ensures stable activity while keeping oscillation frequencies within a physiological range. Our findings tie relevant features of cortical networks, such as E-I balance, the generation of gamma oscillations, and edge-of-bifurcation dynamics, under the framework of firing-rate homeostasis, providing a mechanistic explanation for the heterogeneity in the distance to the bifurcation found across cortical areas. In addition, we reveal the functional benefits of relying upon different homeostatic mechanisms, providing a robust method to regulate network dynamics with minimal perturbation to the generation of gamma rhythms and explaining the correlation between inhibition and gamma frequencies found in cortical networks.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1012723","type":"journal-article","created":{"date-parts":[[2025,1,6]],"date-time":"2025-01-06T13:40:10Z","timestamp":1736170810000},"page":"e1012723","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":10,"title":["Excitatory-inhibitory homeostasis and bifurcation control in the Wilson-Cowan model of cortical dynamics"],"prefix":"10.1371","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3199-4365","authenticated-orcid":true,"given":"Francisco","family":"P\u00e1scoa dos Santos","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul F. M. J.","family":"Verschure","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2025,1,6]]},"reference":[{"issue":"4","key":"pcbi.1012723.ref001","doi-asserted-by":"crossref","first-page":"480","DOI":"10.1162\/neco.1989.1.4.480","article-title":"A Canonical Microcircuit for Neocortex","volume":"1","author":"RJ Douglas","year":"1989","journal-title":"Neural Computation"},{"issue":"5","key":"pcbi.1012723.ref002","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1113\/expphysiol.1988.sp003190","article-title":"The Wellcome Prize lecture. From single cells to simple circuits in the cerebral cortex","volume":"73","author":"KA Martin","year":"1988","journal-title":"Quarterly Journal of Experimental Physiology (Cambridge, England)"},{"key":"pcbi.1012723.ref003","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1016\/j.neuroscience.2017.05.027","article-title":"Inhibitory interneurons and their circuit motifs in the many layers of the barrel cortex","volume":"368","author":"D Feldmeyer","year":"2018","journal-title":"Neuroscience"},{"key":"pcbi.1012723.ref004","doi-asserted-by":"crossref","first-page":"419","DOI":"10.1146\/annurev.neuro.27.070203.144152","article-title":"Neuronal circuits of the neocortex","volume":"27","author":"RJ Douglas","year":"2004","journal-title":"Annual Review of Neuroscience"},{"issue":"5","key":"pcbi.1012723.ref005","doi-asserted-by":"crossref","first-page":"051909","DOI":"10.1103\/PhysRevE.70.051909","article-title":"High-order events in cortical networks: A lower bound","volume":"70","author":"A Benucci","year":"2004","journal-title":"Physical Review E"},{"issue":"2","key":"pcbi.1012723.ref006","first-page":"187","article-title":"Relationships of local inhibitory and excitatory circuits to orientation preference maps in ferret visual cortex","volume":"12","author":"B Roerig","year":"2002","journal-title":"Cerebral Cortex (New York, NY: 1991)"},{"issue":"3","key":"pcbi.1012723.ref007","doi-asserted-by":"crossref","first-page":"169","DOI":"10.1038\/s41583-020-0262-x","article-title":"Macroscopic gradients of synaptic excitation and inhibition in the neocortex","volume":"21","author":"XJ Wang","year":"2020","journal-title":"Nature Reviews Neuroscience"},{"issue":"12","key":"pcbi.1012723.ref008","doi-asserted-by":"crossref","first-page":"1690","DOI":"10.1038\/nn.4415","article-title":"Synaptic scaling rule preserves excitatory-inhibitory balance and salient neuronal network dynamics","volume":"19","author":"J Barral","year":"2016","journal-title":"Nature Neuroscience"},{"issue":"1","key":"pcbi.1012723.ref009","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1113\/jphysiol.1991.sp018733","article-title":"A functional microcircuit for cat visual cortex","volume":"440","author":"RJ Douglas","year":"1991","journal-title":"The Journal of Physiology"},{"issue":"6965","key":"pcbi.1012723.ref010","doi-asserted-by":"crossref","first-page":"442","DOI":"10.1038\/nature02116","article-title":"Balanced inhibition underlies tuning and sharpens spike timing in auditory cortex","volume":"426","author":"M Wehr","year":"2003","journal-title":"Nature"},{"issue":"5","key":"pcbi.1012723.ref011","doi-asserted-by":"crossref","first-page":"535","DOI":"10.1038\/nn.2105","article-title":"Instantaneous correlation of excitation and inhibition during ongoing and sensory-evoked activities","volume":"11","author":"M Okun","year":"2008","journal-title":"Nature Neuroscience"},{"issue":"7511","key":"pcbi.1012723.ref012","doi-asserted-by":"crossref","first-page":"596","DOI":"10.1038\/nature13321","article-title":"Equalizing excitation-inhibition ratios across visual cortical neurons","volume":"511","author":"M Xue","year":"2014","journal-title":"Nature"},{"issue":"5293","key":"pcbi.1012723.ref013","doi-asserted-by":"crossref","first-page":"1724","DOI":"10.1126\/science.274.5293.1724","article-title":"Chaos in neuronal networks with balanced excitatory and inhibitory activity","volume":"274","author":"C van Vreeswijk","year":"1996","journal-title":"Science (New York, NY)"},{"issue":"6062","key":"pcbi.1012723.ref014","doi-asserted-by":"crossref","first-page":"1569","DOI":"10.1126\/science.1211095","article-title":"Inhibitory Plasticity Balances Excitation and Inhibition in Sensory Pathways and Memory Networks","volume":"334","author":"TP Vogels","year":"2011","journal-title":"Science"},{"key":"pcbi.1012723.ref015","doi-asserted-by":"crossref","DOI":"10.3389\/fphys.2012.00163","article-title":"Being Critical of Criticality in the Brain","volume":"3","author":"J Beggs","year":"2012","journal-title":"Frontiers in Physiology"},{"issue":"45","key":"pcbi.1012723.ref016","doi-asserted-by":"crossref","first-page":"eabg8411","DOI":"10.1126\/sciadv.abg8411","article-title":"Excitatory-inhibitory balance modulates the formation and dynamics of neuronal assemblies in cortical networks","volume":"7","author":"S Sadeh","year":"2021","journal-title":"Science Advances"},{"key":"pcbi.1012723.ref017","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1146\/annurev-neuro-060909-153238","article-title":"Too many cooks? Intrinsic and synaptic homeostatic mechanisms in cortical circuit refinement","volume":"34","author":"G Turrigiano","year":"2011","journal-title":"Annual Review of Neuroscience"},{"key":"pcbi.1012723.ref018","doi-asserted-by":"crossref","DOI":"10.1146\/annurev-neuro-092523-110001","article-title":"Keeping Your Brain in Balance: Homeostatic Regulation of Network Function","author":"W Wen","year":"2024","journal-title":"Annual Review of Neuroscience"},{"issue":"6670","key":"pcbi.1012723.ref019","doi-asserted-by":"crossref","first-page":"892","DOI":"10.1038\/36103","article-title":"Activity-dependent scaling of quantal amplitude in neocortical neurons","volume":"391","author":"GG Turrigiano","year":"1998","journal-title":"Nature"},{"issue":"17","key":"pcbi.1012723.ref020","doi-asserted-by":"crossref","first-page":"4377","DOI":"10.1523\/JNEUROSCI.5298-07.2008","article-title":"Multiple Modes of Network Homeostasis in Visual Cortical Layer 2\/3","volume":"28","author":"A Maffei","year":"2008","journal-title":"Journal of Neuroscience"},{"issue":"12","key":"pcbi.1012723.ref021","doi-asserted-by":"crossref","first-page":"1353","DOI":"10.1038\/nn1351","article-title":"Selective reconfiguration of layer 4 visual cortical circuitry by visual deprivation","volume":"7","author":"A Maffei","year":"2004","journal-title":"Nature Neuroscience"},{"issue":"2","key":"pcbi.1012723.ref022","doi-asserted-by":"crossref","first-page":"327","DOI":"10.1016\/j.neuron.2013.08.018","article-title":"Synaptic scaling and homeostatic plasticity in the mouse visual cortex in vivo","volume":"80","author":"T Keck","year":"2013","journal-title":"Neuron"},{"issue":"6","key":"pcbi.1012723.ref023","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1038\/9165","article-title":"Plasticity in the intrinsic excitability of cortical pyramidal neurons","volume":"2","author":"NS Desai","year":"1999","journal-title":"Nature Neuroscience"},{"issue":"4","key":"pcbi.1012723.ref024","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1016\/j.neuron.2010.09.033","article-title":"Visual deprivation suppresses L5 pyramidal neuron excitability by preventing the induction of intrinsic plasticity","volume":"68","author":"K Nataraj","year":"2010","journal-title":"Neuron"},{"issue":"48","key":"pcbi.1012723.ref025","doi-asserted-by":"crossref","first-page":"9891","DOI":"10.1523\/JNEUROSCI.1200-21.2021","article-title":"Developmental Regulation of Homeostatic Plasticity in Mouse Primary Visual Cortex","volume":"41","author":"W Wen","year":"2021","journal-title":"The Journal of Neuroscience"},{"issue":"4","key":"pcbi.1012723.ref026","doi-asserted-by":"crossref","first-page":"655","DOI":"10.1016\/j.neuron.2019.08.031","article-title":"Cortical Circuit Dynamics Are Homeostatically Tuned to Criticality In Vivo","volume":"104","author":"Z Ma","year":"2019","journal-title":"Neuron"},{"issue":"6","key":"pcbi.1012723.ref027","doi-asserted-by":"crossref","first-page":"350","DOI":"10.1007\/s00422-005-0563-1","article-title":"A field-theoretic approach to understanding scale-free neocortical dynamics","volume":"92","author":"WJ Freeman","year":"2005","journal-title":"Biological Cybernetics"},{"key":"pcbi.1012723.ref028","doi-asserted-by":"crossref","first-page":"770","DOI":"10.1002\/hbm.20590","article-title":"Avalanche dynamics of human brain oscillations: Relation to critical branching processes and temporal correlations","volume":"29","author":"SS Poil","year":"2008","journal-title":"Human Brain Mapping"},{"issue":"4","key":"pcbi.1012723.ref029","doi-asserted-by":"crossref","first-page":"112254","DOI":"10.1016\/j.celrep.2023.112254","article-title":"Time-resolved correlation of distributed brain activity tracks E-I balance and accounts for diverse scale-free phenomena","volume":"42","author":"A Nanda","year":"2023","journal-title":"Cell Reports"},{"issue":"5","key":"pcbi.1012723.ref030","doi-asserted-by":"crossref","first-page":"497","DOI":"10.1016\/j.neunet.2005.06.014","article-title":"Metastability, instability, and state transition in neocortex","volume":"18","author":"WJ Freeman","year":"2005","journal-title":"Neural Networks"},{"key":"pcbi.1012723.ref031","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1016\/j.neuroimage.2013.11.047","article-title":"Exploring mechanisms of spontaneous functional connectivity in MEG: how delayed network interactions lead to structured amplitude envelopes of band-pass filtered oscillations","volume":"90","author":"J Cabral","year":"2014","journal-title":"NeuroImage"},{"issue":"1","key":"pcbi.1012723.ref032","doi-asserted-by":"crossref","first-page":"3095","DOI":"10.1038\/s41598-017-03073-5","article-title":"The dynamics of resting fluctuations in the brain: metastability and its dynamical cortical core","volume":"7","author":"G Deco","year":"2017","journal-title":"Scientific Reports"},{"key":"pcbi.1012723.ref033","doi-asserted-by":"crossref","unstructured":"Arsiwalla XD, Verschure P. Why the Brain Might Operate Near the Edge of Criticality. In: Lintas A, Rovetta S, Verschure PFMJ, Villa AEP, editors. Artificial Neural Networks and Machine Learning\u2014ICANN 2017. Cham: Springer International Publishing; 2017. p. 326\u2013333.","DOI":"10.1007\/978-3-319-68600-4_38"},{"key":"pcbi.1012723.ref034","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1016\/j.neubiorev.2015.04.014","article-title":"The brain\u2019s code and its canonical computational motifs. From sensory cortex to the default mode network: A multi-scale model of brain function in health and disease","volume":"55","author":"FE Turkheimer","year":"2015","journal-title":"Neuroscience and Biobehavioral Reviews"},{"key":"pcbi.1012723.ref035","doi-asserted-by":"crossref","unstructured":"Harris B, Gollo LL, Fulcher BD. Tracking the distance to criticality in systems with unknown noise; 2024. Available from: http:\/\/arxiv.org\/abs\/2310.14791.","DOI":"10.1103\/PhysRevX.14.031021"},{"issue":"Pt A","key":"pcbi.1012723.ref036","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/j.neuroimage.2015.08.069","article-title":"Local inhibitory plasticity tunes macroscopic brain dynamics and allows the emergence of functional brain networks","volume":"124","author":"PJ Hellyer","year":"2016","journal-title":"NeuroImage"},{"issue":"3","key":"pcbi.1012723.ref037","first-page":"757","article-title":"Multiscale dynamic mean field (MDMF) model relates resting-state brain dynamics with local cortical excitatory-inhibitory neurotransmitter homeostasis","volume":"5","author":"A Naskar","year":"2021","journal-title":"Network Neuroscience"},{"issue":"23","key":"pcbi.1012723.ref038","doi-asserted-by":"crossref","first-page":"7886","DOI":"10.1523\/JNEUROSCI.5068-13.2014","article-title":"How Local Excitation-Inhibition Ratio Impacts the Whole Brain Dynamics","volume":"34","author":"G Deco","year":"2014","journal-title":"Journal of Neuroscience"},{"issue":"2","key":"pcbi.1012723.ref039","doi-asserted-by":"crossref","first-page":"e1006007","DOI":"10.1371\/journal.pcbi.1006007","article-title":"A biophysical model of dynamic balancing of excitation and inhibition in fast oscillatory large-scale networks","volume":"14","author":"RG Abeysuriya","year":"2018","journal-title":"PLoS computational biology"},{"key":"pcbi.1012723.ref040","doi-asserted-by":"crossref","first-page":"120236","DOI":"10.1016\/j.neuroimage.2023.120236","article-title":"Multi-modal and multi-model interrogation of large-scale functional brain networks","volume":"277","author":"F Castaldo","year":"2023","journal-title":"NeuroImage"},{"issue":"7","key":"pcbi.1012723.ref041","doi-asserted-by":"crossref","first-page":"e1011279","DOI":"10.1371\/journal.pcbi.1011279","article-title":"Multiscale effects of excitatory-inhibitory homeostasis in lesioned cortical networks: A computational study","volume":"19","author":"F P\u00e1scoa dos Santos","year":"2023","journal-title":"PLOS Computational Biology"},{"issue":"1715","key":"pcbi.1012723.ref042","doi-asserted-by":"crossref","first-page":"20160157","DOI":"10.1098\/rstb.2016.0157","article-title":"Multiple shared mechanisms for homeostatic plasticity in rodent somatosensory and visual cortex","volume":"372","author":"MA Gainey","year":"2017","journal-title":"Philosophical Transactions of the Royal Society of London Series B, Biological Sciences"},{"issue":"3","key":"pcbi.1012723.ref043","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1016\/S0896-6273(00)80563-2","article-title":"BDNF has opposite effects on the quantal amplitude of pyramidal neuron and interneuron excitatory synapses","volume":"21","author":"LC Rutherford","year":"1998","journal-title":"Neuron"},{"issue":"1","key":"pcbi.1012723.ref044","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0006-3495(72)86068-5","article-title":"Excitatory and inhibitory interactions in localized populations of model neurons","volume":"12","author":"HR Wilson","year":"1972","journal-title":"Biophysical Journal"},{"issue":"1","key":"pcbi.1012723.ref045","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1186\/s13408-015-0034-5","article-title":"Wilson-Cowan Equations for Neocortical Dynamics","volume":"6","author":"JD Cowan","year":"2016","journal-title":"The Journal of Mathematical Neuroscience"},{"key":"pcbi.1012723.ref046","article-title":"Bidirectionally Regulating Gamma Oscillations in Wilson-Cowan Model by Self-Feedback Loops: A Computational Study","volume":"16","author":"X Li","year":"2022","journal-title":"Frontiers in Systems Neuroscience"},{"issue":"6","key":"pcbi.1012723.ref047","doi-asserted-by":"crossref","first-page":"819","DOI":"10.1016\/j.neuron.2008.02.031","article-title":"Rapid synaptic scaling induced by changes in postsynaptic firing","volume":"57","author":"K Ibata","year":"2008","journal-title":"Neuron"},{"issue":"2","key":"pcbi.1012723.ref048","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1016\/j.neuron.2013.08.038","article-title":"Firing rate homeostasis in visual cortex of freely behaving rodents","volume":"80","author":"KB Hengen","year":"2013","journal-title":"Neuron"},{"issue":"1","key":"pcbi.1012723.ref049","first-page":"1","article-title":"Distributed hierarchical processing in the primate cerebral cortex","volume":"1","author":"DJ Felleman","year":"1991","journal-title":"Cerebral Cortex (New York, NY: 1991)"},{"issue":"5","key":"pcbi.1012723.ref050","first-page":"430","article-title":"Intrinsic synaptic organization of the motor cortex","volume":"3","author":"A Keller","year":"1993","journal-title":"Cerebral Cortex (New York, NY: 1991)"},{"issue":"2","key":"pcbi.1012723.ref051","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":"20","key":"pcbi.1012723.ref052","doi-asserted-by":"crossref","first-page":"6402","DOI":"10.1523\/JNEUROSCI.16-20-06402.1996","article-title":"Gamma oscillation by synaptic inhibition in a hippocampal interneuronal network model","volume":"16","author":"XJ Wang","year":"1996","journal-title":"The Journal of Neuroscience: The Official Journal of the Society for Neuroscience"},{"key":"pcbi.1012723.ref053","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780195301069.001.0001","volume-title":"Rhythms of the Brain","author":"G Buzsaki","year":"2006"},{"issue":"6","key":"pcbi.1012723.ref054","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1016\/j.neuron.2009.09.009","article-title":"Cortical enlightenment: are attentional gamma oscillations driven by ING or PING?","volume":"63","author":"P Tiesinga","year":"2009","journal-title":"Neuron"},{"key":"pcbi.1012723.ref055","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1146\/annurev-neuro-062111-150444","article-title":"Mechanisms of gamma oscillations","volume":"35","author":"G Buzsaki","year":"2012","journal-title":"Annual Review of Neuroscience"},{"issue":"25","key":"pcbi.1012723.ref056","doi-asserted-by":"crossref","first-page":"10302","DOI":"10.1073\/pnas.0901831106","article-title":"Key role of coupling, delay, and noise in resting brain fluctuations","volume":"106","author":"G Deco","year":"2009","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"1","key":"pcbi.1012723.ref057","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/s42005-022-00950-y","article-title":"Metastable oscillatory modes emerge from synchronization in the brain spacetime connectome","volume":"5","author":"J Cabral","year":"2022","journal-title":"Communications Physics"},{"issue":"3","key":"pcbi.1012723.ref058","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1177\/003591573502800330","article-title":"Pharmacology and Nerve-Endings","volume":"28","author":"H Dale","year":"1935","journal-title":"Proceedings of the Royal Society of Medicine"},{"issue":"5","key":"pcbi.1012723.ref059","doi-asserted-by":"crossref","first-page":"1290","DOI":"10.1016\/j.neuron.2015.05.010","article-title":"Subnetwork-Specific Homeostatic Plasticity in Mouse Visual Cortex In Vivo","volume":"86","author":"SJ Barnes","year":"2015","journal-title":"Neuron"},{"issue":"9","key":"pcbi.1012723.ref060","doi-asserted-by":"crossref","first-page":"1090","DOI":"10.1038\/nn.2621","article-title":"Narp regulates homeostatic scaling of excitatory synapses on parvalbumin-expressing interneurons","volume":"13","author":"MC Chang","year":"2010","journal-title":"Nature Neuroscience"},{"key":"pcbi.1012723.ref061","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"},{"key":"pcbi.1012723.ref062","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/2526.001.0001","volume-title":"Dynamical Systems in Neuroscience: The Geometry of Excitability and Bursting","author":"EM Izhikevich","year":"2006"},{"key":"pcbi.1012723.ref063","doi-asserted-by":"crossref","first-page":"16347","DOI":"10.1038\/srep16347","article-title":"Gamma oscillations in V1 are correlated with GABA(A) receptor density: A multi-modal MEG and Flumazenil-PET study","volume":"5","author":"J Kujala","year":"2015","journal-title":"Scientific Reports"},{"issue":"2","key":"pcbi.1012723.ref064","doi-asserted-by":"crossref","first-page":"bhad522","DOI":"10.1093\/cercor\/bhad522","article-title":"GABAergic inhibition shapes behavior and neural dynamics in human visual working memory","volume":"34","author":"J Kujala","year":"2024","journal-title":"Cerebral Cortex"},{"issue":"2","key":"pcbi.1012723.ref065","doi-asserted-by":"crossref","first-page":"203","DOI":"10.1113\/jphysiol.1994.sp020124","article-title":"Separate activation of fast and slow inhibitory postsynaptic potentials in rat neocortex in vitro","volume":"476","author":"LS Benardo","year":"1994","journal-title":"The Journal of Physiology"},{"issue":"5614","key":"pcbi.1012723.ref066","doi-asserted-by":"crossref","first-page":"1902","DOI":"10.1126\/science.1082053","article-title":"Identified Sources and Targets of Slow Inhibition in the Neocortex","volume":"299","author":"G Tam\u00e1s","year":"2003","journal-title":"Science"},{"key":"pcbi.1012723.ref067","article-title":"Distinct Superficial and Deep Laminar Domains of Activity in the Visual Cortex during Rest and Stimulation","volume":"4","author":"A Maier","year":"2010","journal-title":"Frontiers in Systems Neuroscience"},{"issue":"40","key":"pcbi.1012723.ref068","doi-asserted-by":"crossref","first-page":"14332","DOI":"10.1073\/pnas.1402773111","article-title":"Alpha and gamma oscillations characterize feedback and feedforward processing in monkey visual cortex","volume":"111","author":"T van Kerkoerle","year":"2014","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"11","key":"pcbi.1012723.ref069","doi-asserted-by":"crossref","first-page":"e1601335","DOI":"10.1126\/sciadv.1601335","article-title":"Feedforward and feedback frequency-dependent interactions in a large-scale laminar network of the primate cortex","volume":"2","author":"JF Mejias","year":"2016","journal-title":"Science Advances"},{"key":"pcbi.1012723.ref070","doi-asserted-by":"crossref","first-page":"119938","DOI":"10.1016\/j.neuroimage.2023.119938","article-title":"A physical neural mass model framework for the analysis of oscillatory generators from laminar electrophysiological recordings","volume":"270","author":"R Sanchez-Todo","year":"2023","journal-title":"NeuroImage"},{"issue":"10","key":"pcbi.1012723.ref071","doi-asserted-by":"crossref","first-page":"3366","DOI":"10.1523\/JNEUROSCI.2523-11.2012","article-title":"Ongoing Cortical Activity at Rest: Criticality, Multistability, and Ghost Attractors","volume":"32","author":"G Deco","year":"2012","journal-title":"Journal of Neuroscience"},{"key":"pcbi.1012723.ref072","doi-asserted-by":"crossref","first-page":"118928","DOI":"10.1016\/j.neuroimage.2022.118928","article-title":"Dynamic primitives of brain network interaction","volume":"250","author":"M Schirner","year":"2022","journal-title":"NeuroImage"},{"issue":"1","key":"pcbi.1012723.ref073","doi-asserted-by":"crossref","first-page":"18001","DOI":"10.1209\/0295-5075\/132\/18001","article-title":"Critical slowing down indicators","volume":"132","author":"F Nazarimehr","year":"2020","journal-title":"Europhysics Letters"},{"issue":"2","key":"pcbi.1012723.ref074","doi-asserted-by":"crossref","first-page":"e1004097","DOI":"10.1371\/journal.pcbi.1004097","article-title":"Critical Slowing Down Governs the Transition to Neuron Spiking","volume":"11","author":"C Meisel","year":"2015","journal-title":"PLOS Computational Biology"},{"issue":"1","key":"pcbi.1012723.ref075","doi-asserted-by":"crossref","first-page":"5363","DOI":"10.1038\/s41467-020-18975-8","article-title":"Long-range phase synchronization of high-frequency oscillations in human cortex","volume":"11","author":"G Arnulfo","year":"2020","journal-title":"Nature Communications"},{"key":"pcbi.1012723.ref076","doi-asserted-by":"crossref","first-page":"171","DOI":"10.1146\/annurev-neuro-071714-034111","article-title":"Long-Range Neural Synchrony in Behavior","volume":"38","author":"AZ Harris","year":"2015","journal-title":"Annual Review of Neuroscience"},{"issue":"3","key":"pcbi.1012723.ref077","doi-asserted-by":"crossref","first-page":"1195","DOI":"10.1152\/physrev.00035.2008","article-title":"Neurophysiological and Computational Principles of Cortical Rhythms in Cognition","volume":"90","author":"XJ Wang","year":"2010","journal-title":"Physiological reviews"},{"issue":"1","key":"pcbi.1012723.ref078","doi-asserted-by":"crossref","first-page":"415","DOI":"10.1152\/jn.01095.2002","article-title":"What Determines the Frequency of Fast Network Oscillations With Irregular Neural Discharges? I. Synaptic Dynamics and Excitation-Inhibition Balance","volume":"90","author":"N Brunel","year":"2003","journal-title":"Journal of Neurophysiology"},{"issue":"4","key":"pcbi.1012723.ref079","doi-asserted-by":"crossref","DOI":"10.1016\/j.isci.2024.109401","article-title":"Emergence of complex oscillatory dynamics in the neuronal networks with long activity time of inhibitory synapses","volume":"27","author":"M Khanjanianpak","year":"2024","journal-title":"iScience"},{"issue":"3","key":"pcbi.1012723.ref080","doi-asserted-by":"crossref","first-page":"404","DOI":"10.1016\/j.neuron.2009.09.020","article-title":"Laminar Structure of Spontaneous and Sensory-Evoked Population Activity in Auditory Cortex","volume":"64","author":"S Sakata","year":"2009","journal-title":"Neuron"},{"issue":"3","key":"pcbi.1012723.ref081","first-page":"785","article-title":"The cell-type specific cortical microcircuit: relating structure and activity in a full-scale spiking network model","volume":"24","author":"TC Potjans","year":"2014","journal-title":"Cerebral Cortex (New York, NY: 1991)"},{"issue":"5","key":"pcbi.1012723.ref082","doi-asserted-by":"crossref","first-page":"1117","DOI":"10.1073\/pnas.1710323115","article-title":"Laminar recordings in frontal cortex suggest distinct layers for maintenance and control of working memory","volume":"115","author":"AM Bastos","year":"2018","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"3","key":"pcbi.1012723.ref083","doi-asserted-by":"crossref","first-page":"500","DOI":"10.1016\/j.neuron.2018.12.009","article-title":"Layer-Specific Physiological Features and Interlaminar Interactions in the Primary Visual Cortex of the Mouse","volume":"101","author":"Y Senzai","year":"2019","journal-title":"Neuron"},{"key":"pcbi.1012723.ref084","doi-asserted-by":"crossref","first-page":"806","DOI":"10.1016\/j.neuroimage.2017.06.045","article-title":"Benchmarking laminar fMRI: Neuronal spiking and synaptic activity during top-down and bottom-up processing in the different layers of cortex","volume":"197","author":"MW Self","year":"2019","journal-title":"NeuroImage"},{"issue":"6158","key":"pcbi.1012723.ref085","doi-asserted-by":"crossref","first-page":"1238406","DOI":"10.1126\/science.1238406","article-title":"Cortical high-density counterstream architectures","volume":"342","author":"NT Markov","year":"2013","journal-title":"Science (New York, NY)"},{"issue":"1","key":"pcbi.1012723.ref086","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1093\/cercor\/bhs270","article-title":"A Weighted and Directed Interareal Connectivity Matrix for Macaque Cerebral Cortex","volume":"24","author":"NT Markov","year":"2014","journal-title":"Cerebral Cortex"},{"issue":"4","key":"pcbi.1012723.ref087","doi-asserted-by":"crossref","first-page":"1370","DOI":"10.1523\/JNEUROSCI.21-04-01370.2001","article-title":"Long-Range Temporal Correlations and Scaling Behavior in Human Brain Oscillations","volume":"21","author":"K Linkenkaer-Hansen","year":"2001","journal-title":"Journal of Neuroscience"},{"key":"pcbi.1012723.ref088","doi-asserted-by":"crossref","DOI":"10.3389\/fphys.2012.00450","article-title":"Detrended Fluctuation Analysis: A Scale-Free View on Neuronal Oscillations","volume":"3","author":"R Hardstone","year":"2012","journal-title":"Frontiers in Physiology"},{"issue":"5","key":"pcbi.1012723.ref089","doi-asserted-by":"crossref","first-page":"2004","DOI":"10.1152\/jn.00253.2016","article-title":"Synaptic and intrinsic homeostasis cooperate to optimize single neuron response properties and tune integrator circuits","volume":"116","author":"J Cannon","year":"2016","journal-title":"Journal of Neurophysiology"},{"issue":"39","key":"pcbi.1012723.ref090","doi-asserted-by":"crossref","first-page":"24514","DOI":"10.1073\/pnas.1918368117","article-title":"Homeostatic mechanisms regulate distinct aspects of cortical circuit dynamics","volume":"117","author":"YK Wu","year":"2020","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"1","key":"pcbi.1012723.ref091","doi-asserted-by":"crossref","first-page":"5319","DOI":"10.1038\/ncomms6319","article-title":"Formation and maintenance of neuronal assemblies through synaptic plasticity","volume":"5","author":"A Litwin-Kumar","year":"2014","journal-title":"Nature Communications"},{"issue":"4","key":"pcbi.1012723.ref092","first-page":"045017","article-title":"Supercritical dynamics at the edge-of-chaos underlies optimal decision-making","volume":"2","author":"AF Amil","year":"2021","journal-title":"Journal of Physics: Complexity"},{"issue":"1","key":"pcbi.1012723.ref093","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1007\/s12035-017-0737-6","article-title":"Cholinergic Behavior State-Dependent Mechanisms of Neocortical Gain Control: a Neurocomputational Study","volume":"55","author":"JY Puigb\u00f2","year":"2018","journal-title":"Molecular Neurobiology"},{"issue":"1","key":"pcbi.1012723.ref094","doi-asserted-by":"crossref","first-page":"3754","DOI":"10.1038\/s41598-018-22077-3","article-title":"Associative properties of structural plasticity based on firing rate homeostasis in recurrent neuronal networks","volume":"8","author":"JV Gallinaro","year":"2018","journal-title":"Scientific Reports"},{"issue":"2","key":"pcbi.1012723.ref095","doi-asserted-by":"crossref","first-page":"e1009836","DOI":"10.1371\/journal.pcbi.1009836","article-title":"Homeostatic control of synaptic rewiring in recurrent networks induces the formation of stable memory engrams","volume":"18","author":"JV Gallinaro","year":"2022","journal-title":"PLOS Computational Biology"},{"issue":"13","key":"pcbi.1012723.ref096","doi-asserted-by":"crossref","first-page":"3722","DOI":"10.1523\/JNEUROSCI.2511-15.2016","article-title":"Homeostatic Activity-Dependent Tuning of Recurrent Networks for Robust Propagation of Activity","volume":"36","author":"J Gjorgjieva","year":"2016","journal-title":"Journal of Neuroscience"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1012723","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2025,1,16]],"date-time":"2025-01-16T00:00:00Z","timestamp":1736985600000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1012723","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,1,16]],"date-time":"2025-01-16T14:01:01Z","timestamp":1737036061000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1012723"}},"subtitle":[],"editor":[{"given":"Lorenzo","family":"Fontolan","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2025,1,6]]},"references-count":96,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,1,6]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1012723","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2024.06.05.597312","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,1,6]]}}}