{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T19:14:09Z","timestamp":1785525249345,"version":"3.56.0"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1014521","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T00:00:00Z","timestamp":1785456000000}}],"reference-count":92,"publisher":"Public Library of Science (PLoS)","issue":"7","license":[{"start":{"date-parts":[[2026,7,24]],"date-time":"2026-07-24T00:00:00Z","timestamp":1784851200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Ministry of University and Research","award":["PE0000006"],"award-info":[{"award-number":["PE0000006"]}]},{"DOI":"10.13039\/501100013168","name":"Istituto Nazionale di Fisica Nucleare","doi-asserted-by":"publisher","award":["LINCOLN CSN4"],"award-info":[{"award-number":["LINCOLN CSN4"]}],"id":[{"id":"10.13039\/501100013168","id-type":"DOI","asserted-by":"publisher"}]},{"name":"University of Padova, Dipartimento di Fisica e Astronomia","award":["PARD 2023"],"award-info":[{"award-number":["PARD 2023"]}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>The cerebral cortex operates in a state of restless activity, even in the absence of external stimuli. Collective neuronal activities, such as neural avalanches and synchronized oscillations, are also found under rest conditions, and these features have been suggested to support sensory processing, brain readiness for rapid responses, and computational efficiency. The rat barrel cortex and thalamus circuit, with its somatotopic organization for processing sensory inputs from the whiskers, provides a powerful system to explore such interplay. To characterize these resting state circuits, we perform simultaneous multi-electrode recordings in rats\u2019 barrel cortex and thalamus. During spontaneous activity, oscillations with frequencies centered around 11 Hz are detected concomitantly with slow oscillations below 4 Hz, as well as power-law distributed avalanches. The phase of the lower-frequency oscillation appears to modulate the higher-frequency amplitude, and it has a role in gating avalanche occurrences. We then record neural activity during controlled whisker movements and observe that the 11 Hz barrel circuit active at rest is indeed the one involved in response to whisker stimulation. We finally show how a thalamic-driven firing-rate model can describe the entire phenomenology observed at resting state and predict the response of the barrel cortex to controlled whisker movement, suggesting that the same intrinsic dynamics underlying resting-state activity also shape sensory responses.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1014521","type":"journal-article","created":{"date-parts":[[2026,7,24]],"date-time":"2026-07-24T17:46:46Z","timestamp":1784915206000},"page":"e1014521","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Whisker stimulation reinforces a resting-state network in the barrel cortex: Nested oscillations and avalanches"],"prefix":"10.1371","volume":"22","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7113-8806","authenticated-orcid":true,"given":"Benedetta","family":"Mariani","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ram\u00f3n","family":"Guevara","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mattia","family":"Tambaro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marta","family":"Maschietto","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alessandro","family":"Leparulo","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stefano","family":"Vassanelli","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1603-8375","authenticated-orcid":true,"given":"Samir","family":"Suweis","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,7,24]]},"reference":[{"issue":"1","key":"pcbi.1014521.ref001","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1089\/brain.2011.0019","article-title":"The restless brain","volume":"1","author":"ME Raichle","year":"2011","journal-title":"Brain Connect"},{"issue":"1","key":"pcbi.1014521.ref002","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1038\/nrn2961","article-title":"Emerging concepts for the dynamical organization of resting-state activity in the brain","volume":"12","author":"G Deco","year":"2011","journal-title":"Nature Reviews Neuroscience"},{"key":"pcbi.1014521.ref003","doi-asserted-by":"crossref","first-page":"307","DOI":"10.3389\/fphys.2012.00307","article-title":"What kind of noise is brain noise: anomalous scaling behavior of the resting brain activity fluctuations","volume":"3","author":"D Fraiman","year":"2012","journal-title":"Front Physiol"},{"issue":"2","key":"pcbi.1014521.ref004","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1004100","article-title":"Resting-state temporal synchronization networks emerge from connectivity topology and heterogeneity","volume":"11","author":"A Ponce-Alvarez","year":"2015","journal-title":"PLoS Comput Biol"},{"issue":"6437","key":"pcbi.1014521.ref005","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1126\/science.aav7893","article-title":"Spontaneous behaviors drive multidimensional, brainwide activity","volume":"364","author":"C Stringer","year":"2019","journal-title":"Science"},{"key":"pcbi.1014521.ref006","article-title":"The brain\u2019s \u201cdark energy\u201d puzzle: How strongly is glucose metabolism linked to resting-state brain activity?","author":"T Volpi","year":"2024","journal-title":"Journal of Cerebral Blood Flow & Metabolism"},{"issue":"5","key":"pcbi.1014521.ref007","doi-asserted-by":"crossref","first-page":"762","DOI":"10.1016\/j.neuron.2012.08.019","article-title":"Synaptic energy use and supply","volume":"75","author":"JJ Harris","year":"2012","journal-title":"Neuron"},{"issue":"31","key":"pcbi.1014521.ref008","doi-asserted-by":"crossref","first-page":"13040","DOI":"10.1073\/pnas.0905267106","article-title":"Correspondence of the brain\u2019s functional architecture during activation and rest","volume":"106","author":"SM Smith","year":"2009","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"32","key":"pcbi.1014521.ref009","doi-asserted-by":"crossref","first-page":"13170","DOI":"10.1073\/pnas.0700668104","article-title":"Electrophysiological signatures of resting state networks in the human brain","volume":"104","author":"D Mantini","year":"2007","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"28","key":"pcbi.1014521.ref010","doi-asserted-by":"crossref","first-page":"10095","DOI":"10.1073\/pnas.1319166111","article-title":"Information-based fitness and the emergence of criticality in living systems","volume":"111","author":"J Hidalgo","year":"2014","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"9","key":"pcbi.1014521.ref011","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1016\/j.tics.2021.05.007","article-title":"The secret life of predictive brains: what\u2019s spontaneous activity for?","volume":"25","author":"G Pezzulo","year":"2021","journal-title":"Trends in Cognitive Sciences"},{"issue":"7140","key":"pcbi.1014521.ref012","doi-asserted-by":"crossref","first-page":"83","DOI":"10.1038\/nature05758","article-title":"Intrinsic functional architecture in the anaesthetized monkey brain","volume":"447","author":"JL Vincent","year":"2007","journal-title":"Nature"},{"issue":"10","key":"pcbi.1014521.ref013","doi-asserted-by":"crossref","first-page":"744","DOI":"10.1038\/nphys1803","article-title":"Emergent complex neural dynamics","volume":"6","author":"DR Chialvo","year":"2010","journal-title":"Nature Phys"},{"key":"pcbi.1014521.ref014","first-page":"20","article-title":"The entropic brain: a theory of conscious states informed by neuroimaging research with psychedelic drugs","volume":"8","author":"RL Carhart-Harris","year":"2014","journal-title":"Front Hum Neurosci"},{"issue":"3","key":"pcbi.1014521.ref015","doi-asserted-by":"crossref","first-page":"031001","DOI":"10.1103\/RevModPhys.90.031001","article-title":"Colloquium: Criticality and dynamical scaling in living systems","volume":"90","author":"MA Munoz","year":"2018","journal-title":"Reviews of Modern Physics"},{"issue":"35","key":"pcbi.1014521.ref016","doi-asserted-by":"crossref","first-page":"11167","DOI":"10.1523\/JNEUROSCI.23-35-11167.2003","article-title":"Neuronal avalanches in neocortical circuits","volume":"23","author":"JM Beggs","year":"2003","journal-title":"J Neurosci"},{"issue":"29","key":"pcbi.1014521.ref017","doi-asserted-by":"crossref","first-page":"9817","DOI":"10.1523\/JNEUROSCI.5990-11.2012","article-title":"Critical-state dynamics of avalanches and oscillations jointly emerge from balanced excitation\/inhibition in neuronal networks","volume":"32","author":"S-S Poil","year":"2012","journal-title":"J Neurosci"},{"issue":"21","key":"pcbi.1014521.ref018","doi-asserted-by":"crossref","first-page":"3373","DOI":"10.1016\/j.neuron.2021.07.031","article-title":"What is the dynamical regime of cerebral cortex?","volume":"109","author":"Y Ahmadian","year":"2021","journal-title":"Neuron"},{"issue":"2","key":"pcbi.1014521.ref019","doi-asserted-by":"crossref","first-page":"025010","DOI":"10.1088\/2632-072X\/ac7a83","article-title":"Criticality and network structure drive emergent oscillations in a stochastic whole-brain model","volume":"3","author":"G Barzon","year":"2022","journal-title":"J Phys Complex"},{"issue":"21","key":"pcbi.1014521.ref020","doi-asserted-by":"crossref","first-page":"7576","DOI":"10.1073\/pnas.0800537105","article-title":"Neuronal avalanches organize as nested theta- and beta\/gamma-oscillations during development of cortical layer 2\/3","volume":"105","author":"ED Gireesh","year":"2008","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"37","key":"pcbi.1014521.ref021","doi-asserted-by":"crossref","first-page":"15921","DOI":"10.1073\/pnas.0904089106","article-title":"Spontaneous cortical activity in awake monkeys composed of neuronal avalanches","volume":"106","author":"T Petermann","year":"2009","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1014521.ref022","doi-asserted-by":"crossref","first-page":"204","DOI":"10.3389\/fnsys.2014.00204","article-title":"On the temporal organization of neuronal avalanches","volume":"8","author":"F Lombardi","year":"2014","journal-title":"Front Syst Neurosci"},{"issue":"20","key":"pcbi.1014521.ref023","doi-asserted-by":"crossref","first-page":"208102","DOI":"10.1103\/PhysRevLett.108.208102","article-title":"Universal critical dynamics in high resolution neuronal avalanche data","volume":"108","author":"N Friedman","year":"2012","journal-title":"Phys Rev Lett"},{"issue":"20","key":"pcbi.1014521.ref024","doi-asserted-by":"crossref","first-page":"208101","DOI":"10.1103\/PhysRevLett.122.208101","article-title":"Criticality between Cortical States","volume":"122","author":"AJ Fontenele","year":"2019","journal-title":"Phys Rev Lett"},{"key":"pcbi.1014521.ref025","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.79950","article-title":"Scale-free behavioral dynamics directly linked with scale-free cortical dynamics","volume":"12","author":"SA Jones","year":"2023","journal-title":"Elife"},{"issue":"1","key":"pcbi.1014521.ref026","doi-asserted-by":"crossref","first-page":"2555","DOI":"10.1038\/s41467-023-37976-x","article-title":"Parabolic avalanche scaling in the synchronization of cortical cell assemblies","volume":"14","author":"E Capek","year":"2023","journal-title":"Nat Commun"},{"issue":"6","key":"pcbi.1014521.ref027","doi-asserted-by":"crossref","DOI":"10.1016\/j.neuron.2018.10.045","article-title":"Whole-Brain Neuronal Activity Displays Crackling Noise Dynamics","volume":"100","author":"A Ponce-Alvarez","year":"2018","journal-title":"Neuron"},{"issue":"11","key":"pcbi.1014521.ref028","doi-asserted-by":"crossref","first-page":"820","DOI":"10.1016\/j.tins.2022.08.007","article-title":"How critical is brain criticality?","volume":"45","author":"J O\u2019Byrne","year":"2022","journal-title":"Trends in Neurosciences"},{"issue":"16","key":"pcbi.1014521.ref029","doi-asserted-by":"crossref","DOI":"10.1016\/j.neuron.2025.05.020","article-title":"Is criticality a unified setpoint of brain function?","volume":"113","author":"KB Hengen","year":"2025","journal-title":"Neuron"},{"issue":"17","key":"pcbi.1014521.ref030","doi-asserted-by":"crossref","DOI":"10.1126\/sciadv.adj9303","article-title":"Low-dimensional criticality embedded in high-dimensional awake brain dynamics","volume":"10","author":"AJ Fontenele","year":"2024","journal-title":"Sci Adv"},{"issue":"1","key":"pcbi.1014521.ref031","first-page":"6","article-title":"We got rhythm: Dynamical systems of the nervous system","volume":"47","author":"N Kopell","year":"2000","journal-title":"Notices of the AMS"},{"issue":"7","key":"pcbi.1014521.ref032","doi-asserted-by":"crossref","first-page":"2389","DOI":"10.1111\/ejn.13796","article-title":"Neuronal oscillations: unavoidable and useful?","volume":"48","author":"W Singer","year":"2018","journal-title":"Eur J Neurosci"},{"key":"pcbi.1014521.ref033","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780195301069.001.0001","volume-title":"Rhythms of the Brain","author":"G Buzsaki","year":"2006"},{"issue":"3","key":"pcbi.1014521.ref034","doi-asserted-by":"crossref","first-page":"751","DOI":"10.1016\/j.neuron.2013.10.002","article-title":"Scaling brain size, keeping timing: evolutionary preservation of brain rhythms","volume":"80","author":"G Buzs\u00e1ki","year":"2013","journal-title":"Neuron"},{"issue":"11","key":"pcbi.1014521.ref035","doi-asserted-by":"crossref","first-page":"725","DOI":"10.1016\/j.tins.2015.09.001","article-title":"Neural Cross-Frequency Coupling: Connecting Architectures, Mechanisms, and Functions","volume":"38","author":"A Hyafil","year":"2015","journal-title":"Trends Neurosci"},{"issue":"1","key":"pcbi.1014521.ref036","doi-asserted-by":"crossref","first-page":"4694","DOI":"10.1038\/ncomms5694","article-title":"The contribution of frequency-specific activity to hierarchical information processing in the human auditory cortex","volume":"5","author":"L Fontolan","year":"2014","journal-title":"Nature Communications"},{"issue":"10","key":"pcbi.1014521.ref037","doi-asserted-by":"crossref","first-page":"474","DOI":"10.1016\/j.tics.2005.08.011","article-title":"A mechanism for cognitive dynamics: neuronal communication through neuronal coherence","volume":"9","author":"P Fries","year":"2005","journal-title":"Trends Cogn Sci"},{"issue":"3","key":"pcbi.1014521.ref038","doi-asserted-by":"crossref","first-page":"1904","DOI":"10.1152\/jn.00263.2005","article-title":"An oscillatory hierarchy controlling neuronal excitability and stimulus processing in the auditory cortex","volume":"94","author":"P Lakatos","year":"2005","journal-title":"J Neurophysiol"},{"issue":"2","key":"pcbi.1014521.ref039","doi-asserted-by":"crossref","DOI":"10.1523\/ENEURO.0153-16.2017","article-title":"Communication between Brain Areas Based on Nested Oscillations","volume":"4","author":"M Bonnefond","year":"2017","journal-title":"eNeuro"},{"issue":"1","key":"pcbi.1014521.ref040","doi-asserted-by":"crossref","first-page":"16403","DOI":"10.1038\/s41598-019-52326-y","article-title":"The scale-invariant, temporal profile of neuronal avalanches in relation to cortical \u03b3-oscillations","volume":"9","author":"SR Miller","year":"2019","journal-title":"Sci Rep"},{"issue":"3","key":"pcbi.1014521.ref041","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1038\/s43588-023-00410-9","article-title":"Statistical modeling of adaptive neural networks explains co-existence of avalanches and oscillations in resting human brain","volume":"3","author":"F Lombardi","year":"2023","journal-title":"Nat Comput Sci"},{"issue":"1","key":"pcbi.1014521.ref042","doi-asserted-by":"crossref","first-page":"1849","DOI":"10.1038\/s41467-024-46012-5","article-title":"Gamma oscillatory complexity conveys behavioral information in hippocampal networks","volume":"15","author":"V Douchamps","year":"2024","journal-title":"Nat Commun"},{"issue":"10","key":"pcbi.1014521.ref043","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1002176","article-title":"Synchronous chaos and broad band gamma rhythm in a minimal multi-layer model of primary visual cortex","volume":"7","author":"D Battaglia","year":"2011","journal-title":"PLoS Comput Biol"},{"issue":"8","key":"pcbi.1014521.ref044","doi-asserted-by":"crossref","first-page":"601","DOI":"10.1038\/nrn2411","article-title":"\u201cWhere\u201d and \u201cwhat\u201d in the whisker sensorimotor system","volume":"9","author":"ME Diamond","year":"2008","journal-title":"Nat Rev Neurosci"},{"key":"pcbi.1014521.ref045","volume-title":"The Functions of the Vibrissae in the Behavior of the White Rat\u2026","author":"SB Vincent","year":"1912"},{"issue":"4","key":"pcbi.1014521.ref046","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1016\/j.conb.2006.06.009","article-title":"Active sensation: insights from the rodent vibrissa sensorimotor system","volume":"16","author":"D Kleinfeld","year":"2006","journal-title":"Curr Opin Neurobiol"},{"issue":"2","key":"pcbi.1014521.ref047","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1016\/j.neuron.2007.09.017","article-title":"The functional organization of the barrel cortex","volume":"56","author":"CCH Petersen","year":"2007","journal-title":"Neuron"},{"issue":"2","key":"pcbi.1014521.ref048","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/0006-8993(70)90079-X","article-title":"The structural organization of layer IV in the somatosensory region (SI) of mouse cerebral cortex. The description of a cortical field composed of discrete cytoarchitectonic units","volume":"17","author":"TA Woolsey","year":"1970","journal-title":"Brain Res"},{"issue":"2","key":"pcbi.1014521.ref049","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/0006-8993(80)90065-7","article-title":"Synchrony among rhythmical facial tremor, neocortical \u201calpha\u201d waves, and thalamic non-sensory neuronal bursts in intact awake rats","volume":"195","author":"K Semba","year":"1980","journal-title":"Brain Res"},{"issue":"7018","key":"pcbi.1014521.ref050","doi-asserted-by":"crossref","first-page":"758","DOI":"10.1038\/nature03132","article-title":"Early motor activity drives spindle bursts in the developing somatosensory cortex","volume":"432","author":"R Khazipov","year":"2004","journal-title":"Nature"},{"issue":"12","key":"pcbi.1014521.ref051","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pcbi.1004576","article-title":"Maximizing Sensory Dynamic Range by Tuning the Cortical State to Criticality","volume":"11","author":"SH Gautam","year":"2015","journal-title":"PLoS Comput Biol"},{"issue":"9","key":"pcbi.1014521.ref052","doi-asserted-by":"crossref","first-page":"1310","DOI":"10.1016\/j.neuron.2025.02.009","article-title":"The predictive nature of spontaneous brain activity across scales and species","volume":"113","author":"A Dimakou","year":"2025","journal-title":"Neuron"},{"issue":"1971","key":"pcbi.1014521.ref053","doi-asserted-by":"crossref","first-page":"903","DOI":"10.1098\/rspa.1998.0193","article-title":"The empirical mode decomposition and the Hilbert spectrum for nonlinear and non-stationary time series analysis","volume":"454","author":"NE Huang","year":"1998","journal-title":"Proc R Soc Lond A"},{"issue":"8","key":"pcbi.1014521.ref054","doi-asserted-by":"crossref","first-page":"3100","DOI":"10.1523\/JNEUROSCI.23-08-03100.2003","article-title":"Imaging spatiotemporal dynamics of surround inhibition in the barrels somatosensory cortex","volume":"23","author":"DD Derdikman","year":"2003","journal-title":"Journal of Neuroscience"},{"issue":"17","key":"pcbi.1014521.ref055","doi-asserted-by":"crossref","first-page":"7603","DOI":"10.1523\/JNEUROSCI.19-17-07603.1999","article-title":"Behavioral modulation of tactile responses in the rat somatosensory system","volume":"19","author":"EE Fanselow","year":"1999","journal-title":"J Neurosci"},{"key":"pcbi.1014521.ref056","doi-asserted-by":"crossref","first-page":"10","DOI":"10.3389\/neuro.07.010.2009","article-title":"Behavioral modulation of stimulus-evoked oscillations in barrel cortex of alert rats","volume":"3","author":"S Venkatraman","year":"2009","journal-title":"Front Integr Neurosci"},{"issue":"3","key":"pcbi.1014521.ref057","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1016\/j.clinph.2010.08.006","article-title":"The 5-12 Hz oscillations in the barrel cortex of awake rats--sustained attention during behavioral idling?","volume":"122","author":"A Sobolewski","year":"2011","journal-title":"Clin Neurophysiol"},{"issue":"3","key":"pcbi.1014521.ref058","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1007\/BF00161134","article-title":"A quantitative population model of whisker barrels: re-examining the Wilson-Cowan equations","volume":"3","author":"DJ Pinto","year":"1996","journal-title":"J Comput Neurosci"},{"key":"pcbi.1014521.ref059","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/9609.001.0001","volume-title":"Analyzing neural time series data: theory and practice","author":"MX Cohen","year":"2014"},{"key":"pcbi.1014521.ref060","volume-title":"Introduction to the theory of Fourier integral","author":"EC Titchmarsh","year":"1937"},{"issue":"6","key":"pcbi.1014521.ref061","doi-asserted-by":"crossref","first-page":"1618","DOI":"10.1093\/cercor\/bht358","article-title":"Human thalamus regulates cortical activity via spatially specific and structurally constrained phase-amplitude coupling","volume":"25","author":"M Malekmohammadi","year":"2015","journal-title":"Cereb Cortex"},{"key":"pcbi.1014521.ref062","doi-asserted-by":"crossref","first-page":"709677","DOI":"10.3389\/fnsys.2021.709677","article-title":"Neuronal Avalanches Across the Rat Somatosensory Barrel Cortex and the Effect of Single Whisker Stimulation","volume":"15","author":"B Mariani","year":"2021","journal-title":"Front Syst Neurosci"},{"issue":"2","key":"pcbi.1014521.ref063","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0008982","article-title":"Can power-law scaling and neuronal avalanches arise from stochastic dynamics?","volume":"5","author":"J Touboul","year":"2010","journal-title":"PLoS One"},{"key":"pcbi.1014521.ref064","doi-asserted-by":"crossref","first-page":"012133","DOI":"10.1103\/PhysRevE.100.012133","article-title":"Time-series thresholding and the definition of avalanche size","volume":"100","author":"P Villegas","year":"2019","journal-title":"Phys Rev E"},{"issue":"1","key":"pcbi.1014521.ref065","doi-asserted-by":"crossref","first-page":"10770","DOI":"10.1038\/s41598-022-13686-0","article-title":"Disentangling the critical signatures of neural activity","volume":"12","author":"B Mariani","year":"2022","journal-title":"Sci Rep"},{"issue":"1","key":"pcbi.1014521.ref066","doi-asserted-by":"crossref","first-page":"33","DOI":"10.1093\/cercor\/13.1.33","article-title":"Cortical damping: analysis of thalamocortical response transformations in rodent barrel cortex","volume":"13","author":"DJ Pinto","year":"2003","journal-title":"Cereb Cortex"},{"issue":"4","key":"pcbi.1014521.ref067","doi-asserted-by":"crossref","first-page":"1601","DOI":"10.1523\/JNEUROSCI.13-04-01601.1993","article-title":"Thalamocortical response transformations in simulated whisker barrels","volume":"13","author":"HT Kyriazi","year":"1993","journal-title":"J Neurosci"},{"issue":"7","key":"pcbi.1014521.ref068","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.1712989115","article-title":"Landau-Ginzburg theory of cortex dynamics: Scale-free avalanches emerge at the edge of synchronization","volume":"115","author":"S di Santo","year":"2018","journal-title":"Proc Natl Acad Sci U S A"},{"key":"pcbi.1014521.ref069","doi-asserted-by":"crossref","first-page":"3572","DOI":"10.1038\/ncomms4572","article-title":"Whisker barrel cortex delta oscillations and gamma power in the awake mouse are linked to respiration","volume":"5","author":"J Ito","year":"2014","journal-title":"Nat Commun"},{"key":"pcbi.1014521.ref070","doi-asserted-by":"crossref","first-page":"023224","DOI":"10.1103\/PhysRevResearch.3.023224","article-title":"Hybrid-type synchronization transitions: Where incipient oscillations, scale-free avalanches, and bistability live together","volume":"3","author":"V Buend\u00eda","year":"2021","journal-title":"Phys Rev Res"},{"key":"pcbi.1014521.ref071","article-title":"The somatosensory barrel cortex controls the spindle thalamocortical oscillation by frequency locking","author":"M Tambaro","year":"2025","journal-title":"biorXiv"},{"issue":"1","key":"pcbi.1014521.ref072","doi-asserted-by":"crossref","DOI":"10.1016\/j.neuron.2017.11.020","article-title":"Old Brains Come Uncoupled in Sleep: Slow Wave-Spindle Synchrony, Brain Atrophy, and Forgetting","volume":"97","author":"RF Helfrich","year":"2018","journal-title":"Neuron"},{"issue":"37","key":"pcbi.1014521.ref073","doi-asserted-by":"crossref","first-page":"9538","DOI":"10.1523\/JNEUROSCI.0918-06.2006","article-title":"Layer-specific touch-dependent facilitation and depression in the somatosensory cortex during active whisking","volume":"26","author":"D Derdikman","year":"2006","journal-title":"J Neurosci"},{"issue":"26","key":"pcbi.1014521.ref074","doi-asserted-by":"crossref","first-page":"15330","DOI":"10.1073\/pnas.261273898","article-title":"Thalamic bursting in rats during different awake behavioral states","volume":"98","author":"EE Fanselow","year":"2001","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"1","key":"pcbi.1014521.ref075","doi-asserted-by":"crossref","first-page":"63","DOI":"10.1152\/jn.00487.2003","article-title":"Is spontaneous high-voltage rhythmic spike discharge in Long Evans rats an absence-like seizure activity?","volume":"91","author":"F-Z Shaw","year":"2004","journal-title":"J Neurophysiol"},{"key":"pcbi.1014521.ref076","doi-asserted-by":"crossref","first-page":"88","DOI":"10.3389\/fncir.2015.00088","article-title":"The Slow Oscillation in Cortical and Thalamic Networks: Mechanisms and Functions","volume":"9","author":"GT Neske","year":"2016","journal-title":"Front Neural Circuits"},{"issue":"3","key":"pcbi.1014521.ref077","doi-asserted-by":"crossref","first-page":"1286","DOI":"10.3390\/ijms22031286","article-title":"Understanding the Effects of Anesthesia on Cortical Electrophysiological Recordings: A Scoping Review","volume":"22","author":"V Sorrenti","year":"2021","journal-title":"Int J Mol Sci"},{"issue":"34","key":"pcbi.1014521.ref078","doi-asserted-by":"crossref","first-page":"10600","DOI":"10.1523\/JNEUROSCI.2053-09.2009","article-title":"A simple model of cortical dynamics explains variability and state dependence of sensory responses in urethane-anesthetized auditory cortex","volume":"29","author":"C Curto","year":"2009","journal-title":"J Neurosci"},{"key":"pcbi.1014521.ref079","doi-asserted-by":"crossref","unstructured":"Pikovsky A, Rosenblum M, Kurths J. Synchronization: a universal concept in nonlinear science. 2002.","DOI":"10.1017\/CBO9780511755743"},{"issue":"16","key":"pcbi.1014521.ref080","doi-asserted-by":"crossref","first-page":"6241","DOI":"10.1523\/JNEUROSCI.20-16-06241.2000","article-title":"Physiological and anatomical organization of multiwhisker response interactions in the barrel cortex of rats","volume":"20","author":"S Shimegi","year":"2000","journal-title":"J Neurosci"},{"issue":"6","key":"pcbi.1014521.ref081","doi-asserted-by":"crossref","first-page":"935","DOI":"10.1002\/cne.24381","article-title":"Brain maps 4.0-Structure of the rat brain: An open access atlas with global nervous system nomenclature ontology and flatmaps","volume":"526","author":"LW Swanson","year":"2018","journal-title":"J Comp Neurol"},{"issue":"8","key":"pcbi.1014521.ref082","doi-asserted-by":"crossref","first-page":"1661","DOI":"10.1162\/089976604774201631","article-title":"Unsupervised spike detection and sorting with wavelets and superparamagnetic clustering","volume":"16","author":"RQ Quiroga","year":"2004","journal-title":"Neural Comput"},{"issue":"4","key":"pcbi.1014521.ref083","doi-asserted-by":"crossref","first-page":"2137","DOI":"10.1152\/jn.00582.2002","article-title":"Local field potentials and the encoding of whisker deflections by population firing synchrony in thalamic barreloids","volume":"89","author":"S Temereanca","year":"2003","journal-title":"J Neurophysiol"},{"issue":"47","key":"pcbi.1014521.ref084","doi-asserted-by":"crossref","first-page":"19113","DOI":"10.1073\/pnas.1312691110","article-title":"Cellular organization of cortical barrel columns is whisker-specific","volume":"110","author":"HS Meyer","year":"2013","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"59","key":"pcbi.1014521.ref085","doi-asserted-by":"crossref","first-page":"2977","DOI":"10.21105\/joss.02977","article-title":"EMD: Empirical Mode Decomposition and Hilbert-Huang Spectral Analyses in Python","volume":"6","author":"AJ Quinn","year":"2021","journal-title":"J Open Source Softw"},{"issue":"5","key":"pcbi.1014521.ref086","doi-asserted-by":"crossref","first-page":"1670","DOI":"10.1152\/jn.00315.2021","article-title":"Automatic decomposition of electrophysiological data into distinct nonsinusoidal oscillatory modes","volume":"126","author":"MS Fabus","year":"2021","journal-title":"J Neurophysiol"},{"key":"pcbi.1014521.ref087","doi-asserted-by":"crossref","unstructured":"Ryan Deering, James F. Kaiser. The Use of a Masking Signal to Improve Empirical Mode Decomposition. In: Proceedings. (ICASSP \u201905). IEEE International Conference on Acoustics, Speech, and Signal Processing, 2005. 485\u20138. https:\/\/doi.org\/10.1109\/icassp.2005.1416051","DOI":"10.1109\/ICASSP.2005.1416051"},{"key":"pcbi.1014521.ref088","article-title":"Method for mode mixing separation in empirical mode decomposition","author":"OB Fosso","year":"2017","journal-title":"arXiv preprint"},{"issue":"4","key":"pcbi.1014521.ref089","doi-asserted-by":"crossref","first-page":"661","DOI":"10.1137\/070710111","article-title":"Power-Law Distributions in Empirical Data","volume":"51","author":"A Clauset","year":"2009","journal-title":"SIAM Rev"},{"issue":"10","key":"pcbi.1014521.ref090","doi-asserted-by":"crossref","first-page":"107840","DOI":"10.1016\/j.isci.2023.107840","article-title":"Criticality of neuronal avalanches in human sleep and their relationship with sleep macro- and micro-architecture","volume":"26","author":"S Scarpetta","year":"2023","journal-title":"iScience"},{"issue":"1","key":"pcbi.1014521.ref091","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":"Biophys J"},{"issue":"3","key":"pcbi.1014521.ref092","doi-asserted-by":"crossref","first-page":"403","DOI":"10.1002\/nav.3800260304","article-title":"Simulation of nonhomogeneous poisson processes by thinning","volume":"26","author":"PAW Lewis","year":"1979","journal-title":"Naval Research Logistics"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1014521","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T00:00:00Z","timestamp":1785456000000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1014521","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,31]],"date-time":"2026-07-31T17:43:47Z","timestamp":1785519827000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1014521"}},"subtitle":[],"editor":[{"given":"Luisa Fernanda","family":"Ramirez","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2026,7,24]]},"references-count":92,"journal-issue":{"issue":"7","published-online":{"date-parts":[[2026,7,24]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1014521","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,24]]}}}