{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,24]],"date-time":"2026-04-24T12:49:13Z","timestamp":1777034953584,"version":"3.51.4"},"reference-count":72,"publisher":"Public Library of Science (PLoS)","issue":"11","content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"DOI":"10.1371\/journal.pcbi.1000209","type":"journal-article","created":{"date-parts":[[2008,11,13]],"date-time":"2008-11-13T22:38:21Z","timestamp":1226615901000},"page":"e1000209","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":607,"title":["A Hierarchy of Time-Scales and the Brain"],"prefix":"10.1371","volume":"4","author":[{"given":"Stefan J.","family":"Kiebel","sequence":"first","affiliation":[]},{"given":"Jean","family":"Daunizeau","sequence":"additional","affiliation":[]},{"given":"Karl J.","family":"Friston","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2008,11,14]]},"reference":[{"key":"ref1","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1162\/1064546053278973","article-title":"The development of embodied cognition: six lessons from babies.","volume":"11","author":"L Smith","year":"2005","journal-title":"Artif Life"},{"key":"ref2","doi-asserted-by":"crossref","first-page":"553","DOI":"10.1016\/S0166-2236(97)01149-1","article-title":"The brain has a body: adaptive behavior emerges from interactions of nervous system, body and environment.","volume":"20","author":"HJ Chiel","year":"1997","journal-title":"Trends Neurosci"},{"key":"ref3","doi-asserted-by":"crossref","first-page":"1344","DOI":"10.1126\/science.1145463","article-title":"Evolution in the social brain.","volume":"317","author":"RI Dunbar","year":"2007","journal-title":"Science"},{"key":"ref4","doi-asserted-by":"crossref","first-page":"233","DOI":"10.1159\/000076784","article-title":"Brains, innovations and evolution in birds and primates.","volume":"63","author":"L Lefebvre","year":"2004","journal-title":"Brain Behav Evol"},{"key":"ref5","doi-asserted-by":"crossref","first-page":"4436","DOI":"10.1073\/pnas.062041299","article-title":"Social intelligence, innovation, and enhanced brain size in primates.","volume":"99","author":"SM Reader","year":"2002","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref6","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1016\/j.jphysparis.2006.10.001","article-title":"A free energy principle for the brain.","volume":"100","author":"K Friston","year":"2006","journal-title":"J Physiol Paris"},{"key":"ref7","first-page":"1595","article-title":"Perception.","author":"H Teuber","year":"1960"},{"key":"ref8","doi-asserted-by":"crossref","first-page":"815","DOI":"10.1098\/rstb.2005.1622","article-title":"A theory of cortical responses.","volume":"360","author":"K Friston","year":"2005","journal-title":"Philos Trans R Soc B Biol Sci"},{"key":"ref9","doi-asserted-by":"crossref","first-page":"79","DOI":"10.1038\/4580","article-title":"Predictive coding in the visual cortex: a functional interpretation of some extra-classical receptive-field effects.","volume":"2","author":"RP Rao","year":"1999","journal-title":"Nat Neurosci"},{"key":"ref10","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1016\/S1364-6613(00)01537-0","article-title":"Forward modeling allows feedback control for fast reaching movements.","volume":"4","author":"M Desmurget","year":"2000","journal-title":"Trends Cogn Sci"},{"key":"ref11","doi-asserted-by":"crossref","first-page":"1880","DOI":"10.1126\/science.7569931","article-title":"An internal model for sensorimotor integration.","volume":"269","author":"DM Wolpert","year":"1995","journal-title":"Science"},{"key":"ref12","doi-asserted-by":"crossref","first-page":"319","DOI":"10.1016\/S0896-6273(01)00285-9","article-title":"The prefrontal cortex\u2014an update: time is of the essence.","volume":"30","author":"JM Fuster","year":"2001","journal-title":"Neuron"},{"key":"ref13","doi-asserted-by":"crossref","first-page":"284","DOI":"10.1016\/j.pneurobio.2007.11.004","article-title":"Is a bird in the hand worth two in the future? The neuroeconomics of intertemporal decision-making.","volume":"84","author":"T Kalenscher","year":"2008","journal-title":"Prog Neurobiol"},{"key":"ref14","doi-asserted-by":"crossref","first-page":"3581","DOI":"10.1121\/1.2934687","article-title":"The temporal analysis of spoken language.","volume":"123","author":"D Poeppel","year":"2008","journal-title":"J Acoust Soc Am"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"1123","DOI":"10.1098\/rstb.2002.1099","article-title":"The prefrontal cortex: categories, concepts and cognition.","volume":"357","author":"EK Miller","year":"2002","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"746","DOI":"10.1038\/264746a0","article-title":"Hearing lips and seeing voices.","volume":"264","author":"H McGurk","year":"1976","journal-title":"Nature"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/j.conb.2007.03.004","article-title":"Efficient reinforcement learning: computational theories, neuroscience and robotics.","volume":"17","author":"M Kawato","year":"2007","journal-title":"Curr Opin Neurobiol"},{"key":"ref18","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1016\/S0921-8890(01)00113-0","article-title":"Acquisition of stand-up behavior by a real robot using hierarchical reinforcement learning.","volume":"36","author":"J Morimoto","year":"2001","journal-title":"Rob Auton Syst"},{"key":"ref19","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1016\/S0531-5131(03)00190-0","article-title":"Hierarchical MOSAIC for movement generation.","volume":"1250","author":"M Haruno","year":"2003","journal-title":"Int Congr Ser"},{"key":"ref20","doi-asserted-by":"crossref","first-page":"1226","DOI":"10.1038\/nn963","article-title":"Optimal feedback control as a theory of motor coordination.","volume":"5","author":"E Todorov","year":"2002","journal-title":"Nat Neurosci"},{"key":"ref21","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1080\/09540090208559329","article-title":"Evolving robots able to self-localize in the environment: the importance of viewing cognition as the result of processes occurring at different time-scales.","volume":"14","author":"S Nolfi","year":"2002","journal-title":"Connect Sci"},{"key":"ref22","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/S0893-6080(02)00214-9","article-title":"Learning to generate articulated behavior through the bottom-up and the top-down interaction processes.","volume":"16","author":"J Tani","year":"2003","journal-title":"Neural Netw"},{"key":"ref23","doi-asserted-by":"crossref","first-page":"e112","DOI":"10.1371\/journal.pcbi.0030112","article-title":"Slowness: an objective for spike-timing-dependent plasticity?","volume":"3","author":"H Sprekeler","year":"2007","journal-title":"PLoS Comput Biol"},{"key":"ref24","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1162\/089976602317318938","article-title":"Slow feature analysis: unsupervised learning of invariances.","volume":"14","author":"L Wiskott","year":"2002","journal-title":"Neural Comput"},{"key":"ref25","doi-asserted-by":"crossref","first-page":"1626","DOI":"10.1126\/science.1128115","article-title":"High gamma power is phase-locked to theta oscillations in human neocortex.","volume":"313","author":"RT Canolty","year":"2006","journal-title":"Science"},{"key":"ref26","doi-asserted-by":"crossref","first-page":"1867","DOI":"10.1073\/pnas.97.4.1867","article-title":"Gamma rhythms and beta rhythms have different synchronization properties.","volume":"97","author":"N Kopell","year":"2000","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref27","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"},{"key":"ref28","doi-asserted-by":"crossref","first-page":"991","DOI":"10.1523\/JNEUROSCI.3387-05.2006","article-title":"Temporal structure in zebra finch song: implications for motor coding.","volume":"26","author":"CM Glaze","year":"2006","journal-title":"J Neurosci"},{"key":"ref29","doi-asserted-by":"crossref","first-page":"1445","DOI":"10.1152\/jn.2001.86.3.1445","article-title":"Feature analysis of natural sounds in the songbird auditory forebrain.","volume":"86","author":"K Sen","year":"2001","journal-title":"J Neurophysiol"},{"key":"ref30","doi-asserted-by":"crossref","first-page":"6924","DOI":"10.1523\/JNEUROSCI.14-11-06924.1994","article-title":"Identification of a forebrain motor programming network for the learned song of zebra finches.","volume":"14","author":"ET Vu","year":"1994","journal-title":"J Neurosci"},{"key":"ref31","doi-asserted-by":"crossref","first-page":"1871","DOI":"10.1126\/science.273.5283.1871","article-title":"Temporal hierarchical control of singing in birds.","volume":"273","author":"AC Yu","year":"1996","journal-title":"Science"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"335","DOI":"10.1016\/j.tics.2006.05.006","article-title":"Probabilistic models of language processing and acquisition.","volume":"10","author":"N Chater","year":"2006","journal-title":"Trends Cogn Sci"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"1071","DOI":"10.1098\/rstb.2007.2160","article-title":"Speech perception at the interface of neurobiology and linguistics.","volume":"363","author":"D Poeppel","year":"2008","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"051921","DOI":"10.1103\/PhysRevE.65.051921","article-title":"Neuromuscular control of vocalizations in birdsong: a model.","volume":"65","author":"R Laje","year":"2002","journal-title":"Phys Rev E Stat Nonlin Soft Matter Phys"},{"key":"ref35","doi-asserted-by":"crossref","first-page":"288102","DOI":"10.1103\/PhysRevLett.89.288102","article-title":"Diversity within a birdsong.","volume":"89","author":"R Laje","year":"2002","journal-title":"Phys Rev Lett"},{"key":"ref36","doi-asserted-by":"crossref","first-page":"849","DOI":"10.1016\/j.neuroimage.2008.02.054","article-title":"DEM: a variational treatment of dynamic systems.","volume":"41","author":"KJ Friston","year":"2008","journal-title":"Neuroimage"},{"key":"ref37","article-title":"Variational algorithms for approximate Bayesian inference [dissertation]","author":"MJ Beal","year":"2003"},{"key":"ref38","doi-asserted-by":"crossref","first-page":"S209","DOI":"10.1088\/1741-2560\/2\/3\/S05","article-title":"Internal models and the construction of time: generalizing from state estimation to trajectory estimation to address temporal features of perception, including temporal illusions.","volume":"2","author":"R Grush","year":"2005","journal-title":"J Neural Eng"},{"key":"ref39","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1093\/cercor\/11.3.183","article-title":"Feature article: the structure and function of dynamic cortical and thalamic receptive fields.","volume":"11","author":"AA Ghazanfar","year":"2001","journal-title":"Cereb Cortex"},{"key":"ref40","doi-asserted-by":"crossref","first-page":"5105","DOI":"10.1523\/JNEUROSCI.3570-06.2007","article-title":"Encoding of movement fragments in the motor cortex.","volume":"27","author":"NG Hatsopoulos","year":"2007","journal-title":"J Neurosci"},{"key":"ref41","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1093\/cercor\/1.1.1","article-title":"Distributed hierarchical processing in the primate cerebral cortex.","volume":"1","author":"DJ Felleman","year":"1991","journal-title":"Cereb Cortex"},{"key":"ref42","doi-asserted-by":"crossref","first-page":"791","DOI":"10.1016\/S0896-6273(02)01091-7","article-title":"View from the top: hierarchies and reverse hierarchies in the visual system.","volume":"36","author":"S Hochstein","year":"2002","journal-title":"Neuron"},{"key":"ref43","doi-asserted-by":"crossref","first-page":"1013","DOI":"10.1093\/brain\/121.6.1013","article-title":"From sensation to cognition.","volume":"121","author":"MM Mesulam","year":"1998","journal-title":"Brain"},{"key":"ref44","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1038\/335311a0","article-title":"The functional logic of cortical connections.","volume":"335","author":"S Zeki","year":"1988","journal-title":"Nature"},{"key":"ref45","doi-asserted-by":"crossref","first-page":"373","DOI":"10.1016\/S0079-6123(02)36031-X","article-title":"Anatomical origins of the classical receptive field and modulatory surround field of single neurons in macaque visual cortical area V1.","volume":"136","author":"A Angelucci","year":"2002","journal-title":"Prog Brain Res"},{"key":"ref46","doi-asserted-by":"crossref","first-page":"727","DOI":"10.1038\/329727a0","article-title":"Corticofugal feedback influences the generation of length tuning in the visual pathway.","volume":"329","author":"PC Murphy","year":"1987","journal-title":"Nature"},{"key":"ref47","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1016\/0006-8993(79)90485-2","article-title":"Laminar origins and terminations of cortical connections of the occipital lobe in the rhesus monkey.","volume":"179","author":"KS Rockland","year":"1979","journal-title":"Brain Res"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"7121","DOI":"10.1073\/pnas.95.12.7121","article-title":"On the actions that one nerve cell can have on another: distinguishing \u201cdrivers\u201d from \u201cmodulators\u201d.","volume":"95","author":"SM Sherman","year":"1998","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1016\/S0079-6123(02)36019-9","article-title":"Cortical interneurons: from Cajal to 2001.","volume":"136","author":"J DeFelipe","year":"2002","journal-title":"Prog Brain Res"},{"key":"ref50","doi-asserted-by":"crossref","first-page":"366","DOI":"10.1016\/j.neuron.2007.10.012","article-title":"Visual field maps in human cortex.","volume":"56","author":"BA Wandell","year":"2007","journal-title":"Neuron"},{"key":"ref51","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/j.conb.2007.07.003","article-title":"The thalamus is more than just a relay.","volume":"17","author":"SM Sherman","year":"2007","journal-title":"Curr Opin Neurobiol"},{"key":"ref52","doi-asserted-by":"crossref","first-page":"e78","DOI":"10.1371\/journal.pcbi.0010078","article-title":"Segregation of the brain into gray and white matter: a design minimizing conduction delays.","volume":"1","author":"Q Wen","year":"2005","journal-title":"PLoS Comput Biol"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/S0167-2789(00)00174-3","article-title":"Fractal time and 1\/f spectra in dynamic images and human vision.","volume":"148","author":"VA Billock","year":"2001","journal-title":"Physica D"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"1051","DOI":"10.1098\/rstb.2005.1643","article-title":"Dynamics of a neural system with a multiscale architecture.","volume":"360","author":"M Breakspear","year":"2005","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"ref55","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/S0167-2789(03)00046-0","article-title":"How fast elements can affect slow dynamics.","volume":"180","author":"K Fujimoto","year":"2003","journal-title":"Physica D"},{"key":"ref56","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-642-88338-5","article-title":"Synergetics","author":"H Haken","year":"1983"},{"key":"ref57","doi-asserted-by":"crossref","first-page":"395","DOI":"10.1016\/0893-6080(90)90022-D","article-title":"Dynamic pattern-recognition of coordinated biological motion.","volume":"3","author":"H Haken","year":"1990","journal-title":"Neural Netw"},{"key":"ref58","doi-asserted-by":"crossref","first-page":"960","DOI":"10.1103\/PhysRevLett.77.960","article-title":"Field theory of electromagnetic brain activity.","volume":"77","author":"VK Jirsa","year":"1996","journal-title":"Phys Rev Lett"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"2019","DOI":"10.1162\/089976698300016954","article-title":"Connecting cortical and behavioral dynamics: bimanual coordination.","volume":"10","author":"VK Jirsa","year":"1998","journal-title":"Neural Comput"},{"key":"ref60","doi-asserted-by":"crossref","first-page":"887","DOI":"10.1037\/0033-295X.113.4.887","article-title":"Hierarchical schemas and goals in the control of sequential behavior.","volume":"113","author":"RP Cooper","year":"2006","journal-title":"Psychol Rev"},{"key":"ref61","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.tics.2008.02.004","article-title":"Cognitive control, hierarchy, and the rostro-caudal organization of the frontal lobes.","volume":"12","author":"D Badre","year":"2008","journal-title":"Trends Cogn Sci"},{"key":"ref62","doi-asserted-by":"crossref","first-page":"201","DOI":"10.1016\/j.tics.2008.02.009","article-title":"Hierarchical models of behavior and prefrontal function.","volume":"12","author":"MM Botvinick","year":"2008","journal-title":"Trends Cogn Sci"},{"key":"ref63","doi-asserted-by":"crossref","first-page":"1615","DOI":"10.1098\/rstb.2007.2056","article-title":"Multilevel structure in behaviour and in the brain: a model of Fuster's hierarchy.","volume":"362","author":"MM Botvinick","year":"2007","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"ref64","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1177\/1059712308089180","article-title":"On cognition as dynamical coupling: an analysis of behavioral attractor dynamics.","volume":"16","author":"A Montebelli","year":"2008","journal-title":"Adaptive Behav"},{"key":"ref65","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1177\/105971230501300303","article-title":"How hierarchical control self-organizes in artificial adaptive systems.","volume":"13","author":"RW Paine","year":"2005","journal-title":"Adaptive Behav"},{"key":"ref66","doi-asserted-by":"crossref","first-page":"143","DOI":"10.1016\/j.tics.2004.02.004","article-title":"Upper processing stages of the perception-action cycle.","volume":"8","author":"JM Fuster","year":"2004","journal-title":"Trends Cogn Sci"},{"key":"ref67","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1111\/j.1749-6632.1990.tb48901.x","article-title":"Prefrontal cortex and the bridging of temporal gaps in the perception-action cycle.","volume":"608","author":"JM Fuster","year":"1990","journal-title":"Ann N Y Acad Sci"},{"key":"ref68","doi-asserted-by":"crossref","first-page":"515","DOI":"10.1152\/jn.00587.2002","article-title":"Spatiotemporal tuning of motor cortical neurons for hand position and velocity.","volume":"91","author":"L Paninski","year":"2004","journal-title":"J Neurophysiol"},{"key":"ref69","doi-asserted-by":"crossref","first-page":"1242","DOI":"10.1016\/j.neunet.2006.06.007","article-title":"Heterarchical reinforcement-learning model for integration of multiple cortico-striatal loops: fMRI examination in stimulus-action-reward association learning.","volume":"19","author":"M Haruno","year":"2006","journal-title":"Neural Netw"},{"key":"ref70","doi-asserted-by":"crossref","first-page":"1127","DOI":"10.1016\/j.neuron.2007.09.038","article-title":"Endogenous cortical rhythms determine cerebral specialization for speech perception and production.","volume":"56","author":"AL Giraud","year":"2007","journal-title":"Neuron"},{"key":"ref71","doi-asserted-by":"crossref","first-page":"1001","DOI":"10.1016\/j.neuron.2007.06.004","article-title":"Phase patterns of neuronal responses reliably discriminate speech in human auditory cortex.","volume":"54","author":"H Luo","year":"2007","journal-title":"Neuron"},{"key":"ref72","doi-asserted-by":"crossref","first-page":"2539","DOI":"10.1523\/JNEUROSCI.5487-07.2008","article-title":"A hierarchy of temporal receptive windows in human cortex.","volume":"28","author":"U Hasson","year":"2008","journal-title":"J Neurosci"}],"container-title":["PLoS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pcbi.1000209","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,5,9]],"date-time":"2020-05-09T01:56:51Z","timestamp":1588989411000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1000209"}},"subtitle":[],"editor":[{"given":"Olaf","family":"Sporns","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2008,11,14]]},"references-count":72,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2008,11,14]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1000209","relation":{},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2008,11,14]]}}}