{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,20]],"date-time":"2026-06-20T15:24:24Z","timestamp":1781969064057,"version":"3.54.5"},"reference-count":67,"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.1000220","type":"journal-article","created":{"date-parts":[[2008,11,6]],"date-time":"2008-11-06T22:54:56Z","timestamp":1226012096000},"page":"e1000220","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":400,"title":["Emergence of Functional Hierarchy in a Multiple Timescale Neural Network Model: A Humanoid Robot Experiment"],"prefix":"10.1371","volume":"4","author":[{"given":"Yuichi","family":"Yamashita","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jun","family":"Tani","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2008,11,7]]},"reference":[{"key":"ref1","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":"ref2","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1098\/rstb.2000.0550","article-title":"Hierarchical organization of macaque and cat cortical sensory systems explored with a novel network processor.","volume":"355","author":"CC Hilgetag","year":"2000","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"ref3","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":"ref4","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1038\/nn1409","article-title":"Hierarchical and asymmetric temporal sensitivity in human auditory cortices.","volume":"8","author":"A Boemio","year":"2005","journal-title":"Nat Neurosci"},{"key":"ref5","article-title":"Neural Organization: Structure, Function, and Dynamics","author":"MA Arbib","year":"1998"},{"key":"ref6","doi-asserted-by":"crossref","first-page":"1755","DOI":"10.1098\/rstb.2000.0733","article-title":"Motor learning through the combination of primitives.","volume":"355","author":"FA Mussa-Ivaldi","year":"2000","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"ref7","doi-asserted-by":"crossref","first-page":"589","DOI":"10.1007\/s00422-006-0127-z","article-title":"Early motor development from partially ordered neural-body dynamics\u2014experiments with a cortico-spinal-musculo-skeletal model.","volume":"95","author":"Y Kuniyoshi","year":"2006","journal-title":"Biol Cybern"},{"key":"ref8","first-page":"742","article-title":"Learning of action through adaptive combination of motor primitives.","volume":"407","author":"KA Thoroughman","year":"2000","journal-title":"Science"},{"key":"ref9","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1007\/s00221-003-1548-8","article-title":"Chunking during human visuomotor sequence learning.","volume":"152","author":"K Sakai","year":"2003","journal-title":"Exp Brain Res"},{"key":"ref10","doi-asserted-by":"crossref","first-page":"467","DOI":"10.1523\/JNEUROSCI.13-02-00467.1993","article-title":"Convergent force fields organized in the frog&apos;s spinal cord.","volume":"13","author":"SF Giszter","year":"1993","journal-title":"J Neurosci"},{"key":"ref11","doi-asserted-by":"crossref","first-page":"349","DOI":"10.1016\/S0896-6273(02)01003-6","article-title":"The cortical control of movement revisited.","volume":"36","author":"MS Graziano","year":"2002","journal-title":"Neuron"},{"key":"ref12","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/6730.001.0001","article-title":"The Embodied Mind","author":"FJ Varela","year":"1991"},{"key":"ref13","doi-asserted-by":"crossref","first-page":"1088","DOI":"10.1126\/science.1145803","article-title":"Self-organization, embodiment, and biologically inspired robotics.","volume":"318","author":"R Pfeifer","year":"2007","journal-title":"Science"},{"key":"ref14","first-page":"S16","article-title":"Robotic neuroscience: a synthetic approach to the brain.","volume":"24","author":"K Doya","year":"2001","journal-title":"Neurosci Res Suppl"},{"key":"ref15","doi-asserted-by":"crossref","first-page":"584","DOI":"10.1016\/j.neunet.2008.03.008","article-title":"Achieving \u201corganic compositionality\u201d through self-organization: reviews on brain-inspired robotics experiments.","volume":"21","author":"J Tani","year":"2008","journal-title":"Neural Netw"},{"key":"ref16","doi-asserted-by":"crossref","first-page":"1131","DOI":"10.1016\/S0893-6080(99)00060-X","article-title":"Learning to perceive the world as articulated: an approach for hierarchical learning in sensory-motor systems.","volume":"12","author":"J Tani","year":"1999","journal-title":"Neural Netw"},{"key":"ref17","doi-asserted-by":"crossref","first-page":"2201","DOI":"10.1162\/089976601750541778","article-title":"Mosaic model for sensorimotor learning and control.","volume":"13","author":"M Haruno","year":"2001","journal-title":"Neural Comput"},{"key":"ref18","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1109\/TSMCB.2007.907738","article-title":"Codevelopmental learning between human and humanoid robot using a dynamic neural-network model.","volume":"38","author":"J Tani","year":"2008","journal-title":"IEEE Trans Syst Man Cybern B Cybern"},{"key":"ref19","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1037\/0033-295X.108.1.57","article-title":"Time scales in motor learning and development.","volume":"108","author":"KM Newell","year":"2001","journal-title":"Psychol Rev"},{"key":"ref20","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1123\/mcj.8.2.188","article-title":"Multiple time scales and multiform dynamics in learning to juggle.","volume":"8","author":"R Huys","year":"2004","journal-title":"Motor Control"},{"key":"ref21","doi-asserted-by":"crossref","first-page":"e179","DOI":"10.1371\/journal.pbio.0040179","article-title":"Interacting adaptive processes with different timescales underlie short-term motor learning.","volume":"4","author":"MA Smith","year":"2006","journal-title":"PLoS Biol"},{"key":"ref22","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1038\/nn1901","article-title":"The dynamics of memory as a consequence of optimal adaptation to a changing body.","volume":"10","author":"KP Kording","year":"2007","journal-title":"Nat Neurosci"},{"key":"ref23","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1038\/35067550","article-title":"The brainweb: phase synchronization and large-scale integration.","volume":"2","author":"F Varela","year":"2001","journal-title":"Nat Rev Neurosci"},{"key":"ref24","doi-asserted-by":"crossref","first-page":"10240","DOI":"10.1073\/pnas.0701519104","article-title":"Network structure of cerebral cortex shapes functional connectivity on multiple time scales.","volume":"104","author":"CJ Honey","year":"2007","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref25","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":"ref26","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":"ref27","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":"PW Paine","year":"2005","journal-title":"Adapt Behav"},{"key":"ref28","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1017\/S0140525X01003910","article-title":"The dynamics of embodiment: a field theory of infant perseverative reaching.","volume":"24","author":"E Thelen","year":"2001","journal-title":"Behav Brain Sci"},{"key":"ref29","doi-asserted-by":"crossref","first-page":"815","DOI":"10.1098\/rstb.2005.1622","article-title":"A theory of cortical responses.","volume":"360","author":"KJ Friston","year":"2005","journal-title":"Philos Trans R Soc Lond B Biol Sci"},{"key":"ref30","first-page":"521","article-title":"Learning nonparametric models for probabilistic imitation.","volume":"19","author":"DB Grimes","year":"2006","journal-title":"Adv Neural Inf Process Syst"},{"key":"ref31","article-title":"Detecting neural state transitions using hidden Markov models for motor cortical prostheses.","author":"C Kemere","year":"2008","journal-title":"J Neurophysiol"},{"key":"ref32","doi-asserted-by":"crossref","first-page":"011918","DOI":"10.1103\/PhysRevE.74.011918","article-title":"Stable propagation of a burst through a one-dimensional homogeneous excitatory chain model of songbird nucleus HVC.","volume":"74","author":"M Li","year":"1996","journal-title":"Phys Rev E"},{"key":"ref33","doi-asserted-by":"crossref","first-page":"2038","DOI":"10.1152\/jn.01311.2006","article-title":"Model of birdsong learning based on gradient estimation by dynamic perturbation of neural conductances.","volume":"98","author":"IR Fiete","year":"2007","journal-title":"J Neurophysiol"},{"key":"ref34","doi-asserted-by":"crossref","first-page":"e1000022","DOI":"10.1371\/journal.pcbi.1000022","article-title":"Structure of spontaneous UP and DOWN transitions self-organizing in a cortical network model.","volume":"4","author":"S Kang","year":"2008","journal-title":"PLoS Comput Biol"},{"key":"ref35","doi-asserted-by":"crossref","first-page":"179","DOI":"10.1207\/s15516709cog1402_1","article-title":"Finding structure in time.","volume":"14","author":"J Elman","year":"1990","journal-title":"Cogn Sci"},{"key":"ref36","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1207\/s15516709cog1603_1","article-title":"Forward models: supervised learning with a distal teacher.","volume":"16","author":"MI Jordan","year":"1992","journal-title":"Cogn Sci"},{"key":"ref37","article-title":"Recurrent network: neurophysiological modeling.","author":"EE Fetz","year":"2002"},{"key":"ref38","doi-asserted-by":"crossref","first-page":"375","DOI":"10.1016\/0893-6080(89)90022-1","article-title":"Adaptive neural oscillator using continuous-time back-propagation learning.","volume":"2","author":"K Doya","year":"1989","journal-title":"Neural Netw"},{"key":"ref39","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1177\/1059712308089185","article-title":"Learning multiple goal-directed actions through self-organization of a dynamic neural network model: a humanoid robot experiment.","volume":"16","author":"R Nishimoto","year":"2008","journal-title":"Adapt Behav"},{"key":"ref40","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":"ref41","doi-asserted-by":"crossref","first-page":"1317","DOI":"10.1016\/S0893-6080(98)00066-5","article-title":"Multiple paired forward and inverse models for motor control.","volume":"11","author":"D Wolpert","year":"1998","journal-title":"Neural Netw"},{"key":"ref42","doi-asserted-by":"crossref","first-page":"213","DOI":"10.1016\/j.neunet.2005.01.001","article-title":"A model of smooth pursuit in primates based on learning the target dynamics.","volume":"18","author":"T Shibata","year":"2005","journal-title":"Neural Netw"},{"key":"ref43","doi-asserted-by":"crossref","first-page":"8170","DOI":"10.1073\/pnas.0802602105","article-title":"Forward estimation of movement state in posterior parietal cortex.","volume":"105","author":"GH Mulliken","year":"2008","journal-title":"Proc Natl Acad Sci U S A"},{"key":"ref44","doi-asserted-by":"crossref","first-page":"925","DOI":"10.1016\/j.neunet.2004.02.003","article-title":"Learning to generate combinatorial action sequences utilizing the initial sensitivity of deterministic dynamical systems.","volume":"17","author":"R Nishimoto","year":"2004","journal-title":"Neural Netw"},{"key":"ref45","doi-asserted-by":"crossref","DOI":"10.7551\/mitpress\/5236.001.0001","article-title":"Parallel distributed processing","author":"DE Rumelhart","year":"1986"},{"key":"ref46","doi-asserted-by":"crossref","first-page":"1063","DOI":"10.1016\/S0896-6273(03)00761-X","article-title":"Learning in spiking neural networks by reinforcement of stochastic synaptic transmission.","volume":"40","author":"HS Seung","year":"2003","journal-title":"Neuron"},{"key":"ref47","doi-asserted-by":"crossref","first-page":"041909","DOI":"10.1103\/PhysRevE.69.041909","article-title":"Learning in neural networks by reinforcement of irregular spiking.","volume":"69","author":"X Xie","year":"2004","journal-title":"Phys Rev E Stat Nonlin Soft Matter Phys"},{"key":"ref48","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1017\/S0140525X00034026","article-title":"The representing brain: neural correlates of motor imitation and imaginary.","volume":"17","author":"M Jeannerod","year":"1994","journal-title":"Behav Brain Sci"},{"key":"ref49","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1109\/3477.499793","article-title":"Model-based learning for mobile robot navigation from the dynamical systems perspective.","volume":"26","author":"J Tani","year":"1996","journal-title":"IEEE Trans Syst Man Cybern B Cybern"},{"key":"ref50","doi-asserted-by":"crossref","first-page":"e1049","DOI":"10.1371\/journal.pone.0001049","article-title":"Identification and classification of hubs in brain networks.","volume":"2","author":"O Sporns","year":"2007","journal-title":"PLoS ONE"},{"key":"ref51","doi-asserted-by":"crossref","first-page":"802","DOI":"10.1002\/hbm.20579","article-title":"Dynamical consequences of lesions in cortical networks.","volume":"29","author":"CJ Honey","year":"2008","journal-title":"Hum Brain Mapp"},{"key":"ref52","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1016\/S0896-6273(01)00423-8","article-title":"The cortical motor system.","volume":"31","author":"G Rizzolatti","year":"2001","journal-title":"Neuron"},{"key":"ref53","doi-asserted-by":"crossref","first-page":"677","DOI":"10.1016\/S0031-9384(02)00929-0","article-title":"Motor areas in the frontal lobe of the primate.","volume":"77","author":"RP Dum","year":"2002","journal-title":"Physiol Behav"},{"key":"ref54","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1007\/BF00816161","article-title":"Topographical organization of cortical efferent zones projecting to distal forelimb muscles in the monkey.","volume":"14","author":"H Asanuma","year":"1972","journal-title":"Exp Brain Res"},{"key":"ref55","doi-asserted-by":"crossref","first-page":"2136","DOI":"10.1126\/science.285.5436.2136","article-title":"Muscle and movement representations in the primary motor cortex.","volume":"285","author":"S Kakei","year":"1999","journal-title":"Science"},{"key":"ref56","doi-asserted-by":"crossref","first-page":"1329","DOI":"10.1523\/JNEUROSCI.02-09-01329.1982","article-title":"The premotor cortex of the monkey.","volume":"2","author":"M Weinrich","year":"1982","journal-title":"J Neurosci"},{"key":"ref57","doi-asserted-by":"crossref","first-page":"385","DOI":"10.1093\/brain\/107.2.385","article-title":"A neurophysiological study of the premotor cortex in the rhesus monkey.","volume":"107","author":"M Weinrich","year":"1984","journal-title":"Brain"},{"key":"ref58","doi-asserted-by":"crossref","first-page":"491","DOI":"10.1007\/BF00248742","article-title":"Functional organization of inferior area 6 in macaque monkey: II. Area F5 and the control of distal movements.","volume":"71","author":"G Rizzolatti","year":"1988","journal-title":"Exp Brain Res"},{"key":"ref59","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1016\/j.neuron.2005.01.027","article-title":"Neural correlates of reaching decisions in dorsal premotor cortex: specification of multiple direction choices and final selection of action.","volume":"45","author":"P Cisek","year":"2005","journal-title":"Neuron"},{"key":"ref60","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1016\/j.conb.2007.02.003","article-title":"Distinctions between dorsal and ventral premotor areas: anatomical connectivity and functional properties.","volume":"17","author":"E Hoshi","year":"2007","journal-title":"Curr Opin Neurobiol"},{"key":"ref61","first-page":"208","article-title":"Selective coding of motor sequence in the supplementary motor area of the monkey cerebral cortex.","volume":"82","author":"H Mushiake","year":"190","journal-title":"Exp Brain Res"},{"key":"ref62","doi-asserted-by":"crossref","first-page":"239","DOI":"10.1016\/j.neuron.2007.09.013","article-title":"Mapping behavioral repertoire onto the cortex.","volume":"56","author":"MS Graziano","year":"2007","journal-title":"Neuron"},{"key":"ref63","doi-asserted-by":"crossref","first-page":"813","DOI":"10.1126\/science.7292014","article-title":"Spatial frequency columns in primary visual cortex.","volume":"214","author":"RB Tootell","year":"1981","journal-title":"Science"},{"key":"ref64","doi-asserted-by":"crossref","first-page":"9270","DOI":"10.1523\/JNEUROSCI.17-23-09270.1997","article-title":"Spatial relationships among three columnar systems in cat area 17.","volume":"17","author":"M Hubener","year":"1997","journal-title":"J Neurosci"},{"key":"ref65","doi-asserted-by":"crossref","first-page":"624","DOI":"10.1038\/nn1057","article-title":"Distinct spatial frequency sensitivities for processing faces and emotional expressions.","volume":"6","author":"P Vuilleumier","year":"2003","journal-title":"Nat Neurosci"},{"key":"ref66","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1016\/0166-2236(90)90117-S","article-title":"The color-opponent and broad-band channels of the primate visual system.","volume":"13","author":"PH Schiller","year":"1990","journal-title":"Trends Neurosci"},{"key":"ref67","article-title":"Self-Organizing Maps","author":"T Kohonen","year":"1996"}],"updated-by":[{"DOI":"10.1371\/annotation\/c580e39c-00bc-43a2-9b15-af71350f9d43","type":"correction","label":"Correction","source":"publisher","updated":{"date-parts":[[2010,10,6]],"date-time":"2010-10-06T00:00:00Z","timestamp":1286323200000}}],"container-title":["PLoS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/dx.plos.org\/10.1371\/journal.pcbi.1000220","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2020,5,9]],"date-time":"2020-05-09T23:22:20Z","timestamp":1589066540000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1000220"}},"subtitle":[],"editor":[{"given":"Olaf","family":"Sporns","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2008,11,7]]},"references-count":67,"journal-issue":{"issue":"11","published-online":{"date-parts":[[2008,11,7]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1000220","relation":{"correction":[{"id-type":"doi","id":"10.1371\/annotation\/c580e39c-00bc-43a2-9b15-af71350f9d43","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2008,11,7]]}}}