{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T05:17:29Z","timestamp":1789708649226,"version":"4.0.1"},"reference-count":123,"publisher":"eLife Sciences Publications, Ltd","license":[{"start":{"date-parts":[[2022,11,14]],"date-time":"2022-11-14T00:00:00Z","timestamp":1668384000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2022,11,14]],"date-time":"2022-11-14T00:00:00Z","timestamp":1668384000000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2022,11,14]],"date-time":"2022-11-14T00:00:00Z","timestamp":1668384000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["elifesciences.org"],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2022,10,26]]},"abstract":"<jats:p>How dynamic interactions between nervous system regions in mammals performs online motor control remains an unsolved problem. In this paper, we show that feedback control is a simple, yet powerful way to understand the neural dynamics of sensorimotor control. We make our case using a minimal model comprising spinal cord, sensory and motor cortex, coupled by long connections that are plastic. It succeeds in learning how to perform reaching movements of a planar arm with 6 muscles in several directions from scratch. The model satisfies biological plausibility constraints, like neural implementation, transmission delays, local synaptic learning and continuous online learning. Using differential Hebbian plasticity the model can go from motor babbling to reaching arbitrary targets in less than 10 min of in silico time. Moreover, independently of the learning mechanism, properly configured feedback control has many emergent properties: neural populations in motor cortex show directional tuning and oscillatory dynamics, the spinal cord creates convergent force fields that add linearly, and movements are ataxic (as in a motor system without a cerebellum).<\/jats:p>","DOI":"10.7554\/elife.77216","type":"journal-article","created":{"date-parts":[[2022,11,14]],"date-time":"2022-11-14T04:15:12Z","timestamp":1668399312000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.77216","source":"Crossref","is-referenced-by-count":10,"title":["Self-configuring feedback loops for sensorimotor control"],"prefix":"10.7554","volume":"11","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-0712-145X","authenticated-orcid":true,"given":"Sergio Oscar","family":"Verduzco-Flores","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02qg15b79","id-type":"ROR","asserted-by":"publisher"}],"name":"Computational Neuroscience Unit, Okinawa Institute of Science and Technology","place":["Okinawa, Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8618-5138","authenticated-orcid":true,"given":"Erik","family":"De Schutter","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02qg15b79","id-type":"ROR","asserted-by":"publisher"}],"name":"Computational Neuroscience Unit, Okinawa Institute of Science and Technology","place":["Okinawa, Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"4374","published-online":{"date-parts":[[2022,11,14]]},"reference":[{"key":"bib1","doi-asserted-by":"publisher","first-page":"611","DOI":"10.1007\/s00429-012-0475-5","article-title":"Predictions not commands: active inference in the motor system","volume":"218","author":"Adams","year":"2013","journal-title":"Brain Structure & Function"},{"key":"bib2","doi-asserted-by":"publisher","first-page":"559","DOI":"10.1146\/annurev-neuro-062111-150527","article-title":"Circuits for skilled reaching and grasping","volume":"35","author":"Alstermark","year":"2012","journal-title":"Annual Review of Neuroscience"},{"key":"bib3","doi-asserted-by":"publisher","first-page":"975","DOI":"10.1016\/j.neuron.2012.05.011","article-title":"Motor circuits in action: specification, connectivity, and function","volume":"74","author":"Arber","year":"2012","journal-title":"Neuron"},{"key":"bib4","doi-asserted-by":"publisher","first-page":"1403","DOI":"10.1126\/science.aat5994","article-title":"Connecting neuronal circuits for movement","volume":"360","author":"Arber","year":"2018","journal-title":"Science"},{"key":"bib5","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0074454","article-title":"Differential joint-specific corticospinal tract projections within the cervical enlargement","volume":"8","author":"Asante","year":"2013","journal-title":"PLOS ONE"},{"key":"bib6","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1111\/tops.12527","article-title":"The hierarchical evolution in human vision modeling","volume":"13","author":"Ballard","year":"2021","journal-title":"Topics in Cognitive Science"},{"key":"bib7","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1007\/s004220050421","article-title":"A large-scale model of some spinal reflex circuits","volume":"78","author":"Bashor","year":"1998","journal-title":"Biol Cybern"},{"key":"bib8","doi-asserted-by":"publisher","first-page":"492","DOI":"10.1152\/jn.1996.76.1.492","article-title":"Cerebellar ataxia: abnormal control of interaction torques across multiple joints","volume":"76","author":"Bastian","year":"1996","journal-title":"Journal of Neurophysiology"},{"key":"bib9","doi-asserted-by":"publisher","first-page":"476","DOI":"10.1017\/S0317167100032194","article-title":"Multi-joint reaching movements and eye-hand tracking in cerebellar incoordination: investigation of a patient with complete loss of Purkinje cells","volume":"18","author":"Becker","year":"1991","journal-title":"Canadian Journal of Neurological Sciences \/ Journal Canadien Des Sciences Neurologiques"},{"key":"bib10","doi-asserted-by":"publisher","DOI":"10.3389\/fninf.2013.00048","article-title":"Nengo: a python tool for building large-scale functional brain models","volume":"7","author":"Bekolay","year":"2014","journal-title":"Frontiers in Neuroinformatics"},{"key":"bib11","doi-asserted-by":"publisher","first-page":"390","DOI":"10.1126\/science.1134960","article-title":"Balanced inhibition and excitation drive spike activity in spinal half-centers","volume":"315","author":"Berg","year":"2007","journal-title":"Science"},{"key":"bib12","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1016\/j.cophys.2019.01.006","article-title":"When networks walk a fine line: balance of excitation and inhibition in spinal motor circuits","volume":"8","author":"Berg","year":"2019","journal-title":"Current Opinion in Physiology"},{"key":"bib13","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1126\/science.1857964","article-title":"Computations underlying the execution of movement: a biological perspective","volume":"253","author":"Bizzi","year":"1991","journal-title":"Science"},{"key":"bib14","doi-asserted-by":"publisher","first-page":"101","DOI":"10.1038\/35039000","article-title":"New perspectives on spinal motor systems","volume":"1","author":"Bizzi","year":"2000","journal-title":"Nature Reviews. Neuroscience"},{"key":"bib15","doi-asserted-by":"publisher","DOI":"10.3389\/fncom.2013.00051","article-title":"The neural origin of muscle synergies","volume":"7","author":"Bizzi","year":"2013","journal-title":"Frontiers in Computational Neuroscience"},{"key":"bib16","doi-asserted-by":"publisher","first-page":"1815","DOI":"10.1152\/jn.00715.2019","article-title":"From motor planning to execution: a sensorimotor loop perspective","volume":"124","author":"Bizzi","year":"2020","journal-title":"Journal of Neurophysiology"},{"key":"bib17","doi-asserted-by":"publisher","DOI":"10.3389\/fninf.2011.00020","article-title":"Modeling the connectome of a simple spinal cord","volume":"5","author":"Borisyuk","year":"2011","journal-title":"Frontiers in Neuroinformatics"},{"key":"bib18","doi-asserted-by":"publisher","first-page":"2761","DOI":"10.1152\/jn.00052.2018","article-title":"Synergies in coordination: a comprehensive overview of neural, computational, and behavioral approaches","volume":"120","author":"Bruton","year":"2018","journal-title":"Journal of Neurophysiology"},{"key":"bib19","doi-asserted-by":"publisher","first-page":"389","DOI":"10.1037\/a0037016","article-title":"Integrating reinforcement learning, equilibrium points, and minimum variance to understand the development of reaching: a computational model","volume":"121","author":"Caligiore","year":"2014","journal-title":"Psychological Review"},{"key":"bib20","doi-asserted-by":"publisher","first-page":"923","DOI":"10.1523\/JNEUROSCI.2301-04.2005","article-title":"Mechanisms of rhythm generation in a spinal locomotor network deprived of crossed connections: the lamprey hemicord","volume":"25","author":"Cangiano","year":"2005","journal-title":"The Journal of Neuroscience"},{"key":"bib21","doi-asserted-by":"publisher","first-page":"172","DOI":"10.1016\/j.neuroscience.2017.06.005","article-title":"A stable brain from unstable components: emerging concepts and implications for neural computation","volume":"357","author":"Chambers","year":"2017","journal-title":"Neuroscience"},{"key":"bib22","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1177\/0278364906063830","article-title":"Adaptive tracking control for robots with unknown kinematic and dynamic properties","volume":"25","author":"Cheah","year":"2006","journal-title":"The International Journal of Robotics Research"},{"key":"bib23","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.neuron.2015.09.021","article-title":"The cognitive architecture of spatial navigation: hippocampal and striatal contributions","volume":"88","author":"Chersi","year":"2015","journal-title":"Neuron"},{"key":"bib24","doi-asserted-by":"publisher","first-page":"51","DOI":"10.1038\/nature11129","article-title":"Neural population dynamics during reaching","volume":"487","author":"Churchland","year":"2012","journal-title":"Nature"},{"key":"bib25","doi-asserted-by":"publisher","first-page":"520","DOI":"10.1007\/s10827-008-0092-8","article-title":"Simulation system of spinal cord motor nuclei and associated nerves and muscles, in a web-based architecture","volume":"25","author":"Cisi","year":"2008","journal-title":"Journal of Computational Neuroscience"},{"key":"bib26","doi-asserted-by":"publisher","DOI":"10.1186\/s13408-015-0034-5","article-title":"Wilson-cowan equations for neocortical dynamics","volume":"6","author":"Cowan","year":"2016","journal-title":"Journal of Mathematical Neuroscience"},{"key":"bib27","doi-asserted-by":"publisher","first-page":"319","DOI":"10.1142\/S0129065707001160","article-title":"Does abnormal spinal reciprocal inhibition lead to co-contraction of antagonist motor units? A modeling study","volume":"17","author":"Cutsuridis","year":"2007","journal-title":"International Journal of Neural Systems"},{"key":"bib28","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.31050","article-title":"Computational modeling of spinal circuits controlling limb coordination and gaits in quadrupeds","volume":"6","author":"Danner","year":"2017","journal-title":"eLife"},{"key":"bib29","doi-asserted-by":"publisher","first-page":"357","DOI":"10.1093\/brain\/121.2.357","article-title":"Influence of vision on upper limb reaching movements in patients with cerebellar ataxia","volume":"121 (Pt 2)","author":"Day","year":"1998","journal-title":"Brain"},{"key":"bib30","doi-asserted-by":"publisher","DOI":"10.3389\/fncom.2013.00019","article-title":"Are muscle synergies useful for neural control?","volume":"7","author":"de Rugy","year":"2013","journal-title":"Frontiers in Computational Neuroscience"},{"key":"bib31","doi-asserted-by":"publisher","DOI":"10.1098\/rspb.2016.2134","article-title":"A spiking neural model of adaptive arm control","volume":"283","author":"DeWolf","year":"2016","journal-title":"Proceedings. Biological Sciences"},{"key":"bib32","doi-asserted-by":"publisher","DOI":"10.3389\/fnbot.2015.00013","article-title":"Cortical spiking network interfaced with virtual musculoskeletal arm and robotic arm","volume":"9","author":"Dura-Bernal","year":"2015","journal-title":"Frontiers in Neurorobotics"},{"key":"bib33","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1159\/000102178","article-title":"Physiology of motor control in man","volume":"44","author":"Eccles","year":"1981","journal-title":"Applied Neurophysiology"},{"key":"bib34","doi-asserted-by":"publisher","first-page":"13763","DOI":"10.1073\/pnas.231499798","article-title":"Degeneracy and complexity in biological systems","volume":"98","author":"Edelman","year":"2001","journal-title":"PNAS"},{"key":"bib35","doi-asserted-by":"publisher","first-page":"3427","DOI":"10.1113\/jphysiol.2014.273581","article-title":"The effective neural drive to muscles is the common synaptic input to motor neurons","volume":"592","author":"Farina","year":"2014","journal-title":"The Journal of Physiology"},{"key":"bib36","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1080\/00222895.1986.10735369","article-title":"Once more on the equilibrium-point hypothesis (lambda model) for motor control","volume":"18","author":"Feldman","year":"1986","journal-title":"Journal of Motor Behavior"},{"key":"bib37","doi-asserted-by":"publisher","first-page":"563","DOI":"10.1016\/j.neuron.2010.02.003","article-title":"Spike-time-dependent plasticity and heterosynaptic competition organize networks to produce long scale-free sequences of neural activity","volume":"65","author":"Fiete","year":"2010","journal-title":"Neuron"},{"key":"bib38","doi-asserted-by":"publisher","first-page":"1527","DOI":"10.1523\/JNEUROSCI.02-11-01527.1982","article-title":"On the relations between the direction of two-dimensional arm movements and cell discharge in primate motor cortex","volume":"2","author":"Georgopoulos","year":"1982","journal-title":"The Journal of Neuroscience"},{"key":"bib39","doi-asserted-by":"publisher","first-page":"1416","DOI":"10.1126\/science.3749885","article-title":"Neuronal population coding of movement direction","volume":"233","author":"Georgopoulos","year":"1986","journal-title":"Science"},{"key":"bib40","doi-asserted-by":"publisher","first-page":"383","DOI":"10.1016\/j.brainresrev.2007.05.001","article-title":"Local shaping of function in the motor cortex: motor contrast, directional tuning","volume":"55","author":"Georgopoulos","year":"2007","journal-title":"Brain Research Reviews"},{"key":"bib41","doi-asserted-by":"publisher","first-page":"311","DOI":"10.1093\/brain\/99.2.311","article-title":"Kinematic effects of deafferentation and cerebellar ablation","volume":"99","author":"Gilman","year":"1976","journal-title":"Brain"},{"key":"bib42","doi-asserted-by":"publisher","first-page":"467","DOI":"10.1523\/JNEUROSCI.13-02-00467.1993","article-title":"Convergent force fields organized in the frog\u2019s spinal cord","volume":"13","author":"Giszter","year":"1993","journal-title":"The Journal of Neuroscience"},{"key":"bib43","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1016\/j.conb.2015.04.004","article-title":"Motor primitives--new data and future questions","volume":"33","author":"Giszter","year":"2015","journal-title":"Current Opinion in Neurobiology"},{"key":"bib44","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1016\/j.conb.2014.03.006","article-title":"Inhibition downunder: an update from the spinal cord","volume":"26","author":"Goulding","year":"2014","journal-title":"Current Opinion in Neurobiology"},{"key":"bib45","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1016\/j.nlm.2013.08.003","article-title":"Learning from the spinal cord: how the study of spinal cord plasticity informs our view of learning","volume":"108","author":"Grau","year":"2014","journal-title":"Neurobiology of Learning and Memory"},{"key":"bib46","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1016\/j.neubiorev.2018.05.009","article-title":"Early human motor development: from variation to the ability to vary and adapt","volume":"90","author":"Hadders-Algra","year":"2018","journal-title":"Neuroscience and Biobehavioral Reviews"},{"key":"bib47","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2012.00127","article-title":"Towards a circuit mechanism for movement tuning in motor cortex","volume":"6","author":"Harrison","year":"2012","journal-title":"Frontiers in Neural Circuits"},{"key":"bib48","volume-title":"Dendrites as Biochemical Compartments","author":"Helmchen","year":"1999"},{"key":"bib49","doi-asserted-by":"publisher","first-page":"10","DOI":"10.1007\/s00422-004-0485-3","article-title":"Biological arm motion through reinforcement learning","volume":"91","author":"Izawa","year":"2004","journal-title":"Biol Cybern"},{"key":"bib50","volume-title":"Neuroscience in the 21st Century: From Basic to Clinical","author":"Jankowska","year":"2013"},{"key":"bib51","doi-asserted-by":"publisher","DOI":"10.1201\/9780203507599","volume-title":"The Primate Visual System","author":"Kaas","year":"2003"},{"key":"bib52","doi-asserted-by":"publisher","first-page":"2136","DOI":"10.1126\/science.285.5436.2136","article-title":"Muscle and movement representations in the primary motor cortex","volume":"285","author":"Kakei","year":"1999","journal-title":"Science"},{"key":"bib53","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1007\/978-0-387-77064-2_8","article-title":"From intention to action: motor cortex and the control of reaching movements","volume":"629","author":"Kalaska","year":"2009","journal-title":"Advances in Experimental Medicine and Biology"},{"key":"bib54","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2021.109577","article-title":"Network-centered homeostasis through inhibition maintains hippocampal spatial map and cortical circuit function","volume":"36","author":"Kaleb","year":"2021","journal-title":"Cell Reports"},{"key":"bib55","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.67256","article-title":"Rotational dynamics in motor cortex are consistent with a feedback controller","volume":"10","author":"Kalidindi","year":"2021","journal-title":"eLife"},{"key":"bib56","doi-asserted-by":"publisher","first-page":"440","DOI":"10.1038\/nn.3643","article-title":"Cortical activity in the null space: permitting preparation without movement","volume":"17","author":"Kaufman","year":"2014","journal-title":"Nature Neuroscience"},{"key":"bib57","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1016\/j.neuroscience.2020.06.019","article-title":"50 years since the MarR, Ito, and albus models of the cerebellum","volume":"462","author":"Kawato","year":"2021","journal-title":"Neuroscience"},{"key":"bib58","volume-title":"Principles Of Applied Mathematics: Transformation And Approximation","author":"Keener","year":"1995"},{"key":"bib59","volume-title":"Progress in Motor Control: A Multidisciplinary Perspective, Advances in Experimental Medicine and Biology","author":"Kelso","year":"2009"},{"key":"bib60","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1152\/jn.00706.2005","article-title":"Primate upper limb muscles exhibit activity patterns that differ from their anatomical action during a postural task","volume":"95","author":"Kurtzer","year":"2006","journal-title":"Journal of Neurophysiology"},{"key":"bib61","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1146\/annurev.neuro.31.060407.125547","article-title":"Descending pathways in motor control","volume":"31","author":"Lemon","year":"2008","journal-title":"Annual Review of Neuroscience"},{"key":"bib62","doi-asserted-by":"publisher","first-page":"586","DOI":"10.1038\/nn.3675","article-title":"Identification of a cellular node for motor control pathways","volume":"17","author":"Levine","year":"2014","journal-title":"Nature Neuroscience"},{"key":"bib63","doi-asserted-by":"publisher","first-page":"4400","DOI":"10.1109\/IEMBS.2005.1615441","article-title":"Hierarchical Feedback and Learning for Multi-joint Arm Movement Control","author":"Li","year":"2005","unstructured":"Li W, Todorov E, Pan X. 2005. Hierarchical Feedback and Learning for Multi-joint Arm Movement Control. In 2005 IEEE Engineering in Medicine and Biology 27th Annual Conference."},{"key":"bib64","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1016\/j.neuron.2012.10.041","article-title":"Preference distributions of primary motor cortex neurons reflect control solutions optimized for limb biomechanics","volume":"77","author":"Lillicrap","year":"2013","journal-title":"Neuron"},{"key":"bib65","doi-asserted-by":"publisher","first-page":"1306","DOI":"10.1038\/nn.3492","article-title":"Balanced cortical microcircuitry for maintaining information in working memory","volume":"16","author":"Lim","year":"2013","journal-title":"Nature Neuroscience"},{"key":"bib66","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1114\/1.1432692","article-title":"Neural and mechanical contributions to the stretch reflex: a model synthesis","volume":"30","author":"Lin","year":"2002","journal-title":"Annals of Biomedical Engineering"},{"key":"bib67","doi-asserted-by":"publisher","DOI":"10.3389\/fncom.2015.00070","article-title":"Major remaining gaps in models of sensorimotor systems","volume":"9","author":"Loeb","year":"2015","journal-title":"Frontiers in Computational Neuroscience"},{"key":"bib68","doi-asserted-by":"publisher","first-page":"1371","DOI":"10.1162\/neco.2008.01-08-698","article-title":"Redundancy, self-motion, and motor control","volume":"21","author":"Martin","year":"2009","journal-title":"Neural Computation"},{"key":"bib69","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1007\/BF00340475","article-title":"Dominance of the short-latency component in perturbation induced electromyographic responses of long-trained monkeys","volume":"64","author":"Meyer-Lohmann","year":"1986","journal-title":"Experimental Brain Research"},{"key":"bib70","doi-asserted-by":"publisher","first-page":"918","DOI":"10.1109\/TCDS.2018.2832844","article-title":"An incremental self-organizing architecture for sensorimotor learning and prediction","volume":"10","author":"Mici","year":"2018","journal-title":"IEEE Transactions on Cognitive and Developmental Systems"},{"key":"bib71","doi-asserted-by":"publisher","first-page":"1772","DOI":"10.1152\/jn.00868.2005","article-title":"Mathematical models of proprioceptors. I. control and transduction in the muscle spindle","volume":"96","author":"Mileusnic","year":"2006","journal-title":"Journal of Neurophysiology"},{"key":"bib72","doi-asserted-by":"publisher","first-page":"2279","DOI":"10.1152\/jn.00632.2002","article-title":"Prediction of muscle activity by populations of sequentially recorded primary motor cortex neurons","volume":"89","author":"Morrow","year":"2003","journal-title":"Journal of Neurophysiology"},{"key":"bib73","doi-asserted-by":"publisher","first-page":"1448","DOI":"10.1038\/nn.4653","article-title":"Spatial representation in the hippocampal formation: a history","volume":"20","author":"Moser","year":"2017","journal-title":"Nature Neuroscience"},{"key":"bib74","doi-asserted-by":"publisher","first-page":"701","DOI":"10.1093\/brain\/120.4.701","article-title":"The columnar organization of the neocortex","volume":"120","author":"Mountcastle","year":"1997","journal-title":"Brain"},{"key":"bib75","doi-asserted-by":"publisher","first-page":"7534","DOI":"10.1073\/pnas.91.16.7534","article-title":"Linear combinations of primitives in vertebrate motor control","volume":"91","author":"Mussa-Ivaldi","year":"1994","journal-title":"PNAS"},{"key":"bib76","doi-asserted-by":"publisher","first-page":"1755","DOI":"10.1098\/rstb.2000.0733","article-title":"Motor learning through the combination of primitives","volume":"355","author":"Mussa\u2013Ivaldi","year":"2000","journal-title":"Philosophical Transactions of the Royal Society of London. Series B"},{"key":"bib77","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1016\/j.neuron.2019.09.045","article-title":"Excitatory and inhibitory subnetworks are equally selective during decision-making and emerge simultaneously during learning","volume":"105","author":"Najafi","year":"2020","journal-title":"Neuron"},{"key":"bib78","doi-asserted-by":"publisher","first-page":"1301","DOI":"10.1016\/j.neuron.2013.08.028","article-title":"Spike-timing-dependent plasticity in primate corticospinal connections induced during free behavior","volume":"80","author":"Nishimura","year":"2013","journal-title":"Neuron"},{"key":"bib79","doi-asserted-by":"publisher","first-page":"138","DOI":"10.1016\/j.cobeha.2017.12.021","article-title":"Acquisition, maintenance, and therapeutic use of a simple motor skill","volume":"20","author":"Norton","year":"2018","journal-title":"Current Opinion in Behavioral Sciences"},{"key":"bib80","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1016\/s1364-6613(98)01241-8","article-title":"Six principles for biologically based computational models of cortical cognition","volume":"2","author":"O\u2019Reilly","year":"1998","journal-title":"Trends in Cognitive Sciences"},{"key":"bib81","doi-asserted-by":"publisher","first-page":"449","DOI":"10.1152\/jn.00876.2002","article-title":"Neuronal activity in the supplementary motor area of monkeys adapting to a new dynamic environment","volume":"91","author":"Padoa-Schioppa","year":"2004","journal-title":"Journal of Neurophysiology"},{"key":"bib82","doi-asserted-by":"publisher","first-page":"2774","DOI":"10.1523\/JNEUROSCI.3349-13.2014","article-title":"Premotor spinal network with balanced excitation and inhibition during motor patterns has high resilience to structural division","volume":"34","author":"Petersen","year":"2014","journal-title":"The Journal of Neuroscience"},{"key":"bib83","doi-asserted-by":"publisher","DOI":"10.1017\/CBO9780511545047","volume-title":"The Circuitry of the Human Spinal Cord","author":"Pierrot-Deseilligny","year":"2005"},{"key":"bib84","doi-asserted-by":"publisher","first-page":"1380","DOI":"10.1162\/neco.2006.18.6.1380","article-title":"Strongly improved stability and faster convergence of temporal sequence learning by using input correlations only","volume":"18","author":"Porr","year":"2006","journal-title":"Neural Computation"},{"key":"bib85","doi-asserted-by":"publisher","first-page":"351","DOI":"10.1126\/science.179.4071.351","article-title":"Feedback: beyond behaviorism stimulus-response laws are wholly predictable within a control-system model of behavioral organization","volume":"179","author":"Powers","year":"1973","journal-title":"Science"},{"key":"bib86","volume-title":"Behavior: The Control of Perception","author":"Powers","year":"2005"},{"key":"bib87","doi-asserted-by":"publisher","first-page":"488","DOI":"10.1038\/s41583-021-00473-5","article-title":"Biological constraints on neural network models of cognitive function","volume":"22","author":"Pulverm\u00fcller","year":"2021","journal-title":"Nature Reviews. Neuroscience"},{"key":"bib88","doi-asserted-by":"publisher","first-page":"1761","DOI":"10.1038\/s41593-019-0520-2","article-title":"A deep learning framework for neuroscience","volume":"22","author":"Richards","year":"2019","journal-title":"Nature Neuroscience"},{"key":"bib89","doi-asserted-by":"publisher","first-page":"28","DOI":"10.3200\/JMBR.36.1.28-38","article-title":"Adaptive motor behavior of cerebellar patients during exposure to unfamiliar external forces","volume":"36","author":"Richter","year":"2004","journal-title":"Journal of Motor Behavior"},{"key":"bib90","doi-asserted-by":"publisher","first-page":"653","DOI":"10.1016\/j.neuron.2007.04.030","article-title":"Motor learning with unstable neural representations","volume":"54","author":"Rokni","year":"2007","journal-title":"Neuron"},{"key":"bib91","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1016\/s0167-9457(02)00159-8","article-title":"Cerebellar ataxia: quantitative assessment and cybernetic interpretation","volume":"22","author":"Sanguineti","year":"2003","journal-title":"Human Movement Science"},{"key":"bib92","doi-asserted-by":"publisher","first-page":"837","DOI":"10.1109\/72.165588","article-title":"Gaussian networks for direct adaptive control","volume":"3","author":"Sanner","year":"1992","journal-title":"IEEE Transactions on Neural Networks"},{"key":"bib93","doi-asserted-by":"crossref","DOI":"10.4324\/9780429467875-12","volume-title":"Reaching for Objects: A Neural Process Account in A Developmental Perspective","author":"Sch\u00f6ner","year":"2018"},{"key":"bib94","doi-asserted-by":"publisher","first-page":"161","DOI":"10.1038\/35093102","article-title":"Dissociation between hand motion and population vectors from neural activity in motor cortex","volume":"413","author":"Scott","year":"2001","journal-title":"Nature"},{"key":"bib95","volume-title":"The Computational Neurobiology of Reaching and Pointing: A Foundation for Motor Learning","author":"Shadmehr","year":"2005"},{"key":"bib96","doi-asserted-by":"publisher","first-page":"1082","DOI":"10.1016\/j.neuroscience.2006.12.072","article-title":"Variability in neuronal activity in primate cortex during working memory tasks","volume":"146","author":"Shafi","year":"2007","journal-title":"Neuroscience"},{"key":"bib97","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1146\/annurev-neuro-062111-150509","article-title":"Cortical control of arm movements: a dynamical systems perspective","volume":"36","author":"Shenoy","year":"2013","journal-title":"Annual Review of Neuroscience"},{"key":"bib98","doi-asserted-by":"publisher","first-page":"2403","DOI":"10.1113\/JP270121","article-title":"Organization of left-right coordination of neuronal activity in the mammalian spinal cord: insights from computational modelling","volume":"593","author":"Shevtsova","year":"2015","journal-title":"The Journal of Physiology"},{"key":"bib99","doi-asserted-by":"publisher","first-page":"6117","DOI":"10.1113\/JP272437","article-title":"Organization of flexor-extensor interactions in the mammalian spinal cord: insights from computational modelling","volume":"594","author":"Shevtsova","year":"2016","journal-title":"The Journal of Physiology"},{"key":"bib100","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22052667","article-title":"Spinal inhibitory interneurons: Gatekeepers of sensorimotor pathways","volume":"22","author":"Stachowski","year":"2021","journal-title":"International Journal of Molecular Sciences"},{"key":"bib101","doi-asserted-by":"publisher","first-page":"266","DOI":"10.1038\/s41586-019-1787-x","article-title":"Distributed coding of choice, action and engagement across the mouse brain","volume":"576","author":"Steinmetz","year":"2019","journal-title":"Nature"},{"key":"bib102","doi-asserted-by":"publisher","first-page":"333","DOI":"10.1007\/s10827-007-0037-7","article-title":"Analysis of reflex modulation with a biologically realistic neural network","volume":"23","author":"Stienen","year":"2007","journal-title":"Journal of Computational Neuroscience"},{"key":"bib103","doi-asserted-by":"publisher","first-page":"544","DOI":"10.1016\/j.neuron.2009.07.018","article-title":"Generating coherent patterns of activity from chaotic neural networks","volume":"63","author":"Sussillo","year":"2009","journal-title":"Neuron"},{"key":"bib104","doi-asserted-by":"publisher","first-page":"1025","DOI":"10.1038\/nn.4042","article-title":"A neural network that finds a naturalistic solution for the production of muscle activity","volume":"18","author":"Sussillo","year":"2015","journal-title":"Nature Neuroscience"},{"key":"bib105","volume-title":"Reinforcement Learning: An Introduction","author":"Sutton","year":"2018"},{"key":"bib106","doi-asserted-by":"publisher","first-page":"8643","DOI":"10.1073\/pnas.1704328114","article-title":"Neural basis for hand muscle synergies in the primate spinal cord","volume":"114","author":"Takei","year":"2017","journal-title":"PNAS"},{"key":"bib107","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.neures.2015.10.012","article-title":"Modeling the motor cortex: optimality, recurrent neural networks, and spatial dynamics","volume":"104","author":"Tanaka","year":"2016","journal-title":"Neuroscience Research"},{"key":"bib108","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1016\/j.neuron.2018.08.016","article-title":"Thalamocortical axonal activity in motor cortex exhibits layer-specific dynamics during motor learning","volume":"100","author":"Tanaka","year":"2018","journal-title":"Neuron"},{"key":"bib109","doi-asserted-by":"publisher","first-page":"391","DOI":"10.1038\/73964","article-title":"Direct cortical control of muscle activation in voluntary arm movements: a model","volume":"3","author":"Todorov","year":"2000","journal-title":"Nature Neuroscience"},{"key":"bib110","doi-asserted-by":"publisher","first-page":"601","DOI":"10.1016\/j.conb.2009.09.002","article-title":"The case for and against muscle synergies","volume":"19","author":"Tresch","year":"2009","journal-title":"Current Opinion in Neurobiology"},{"key":"bib111","doi-asserted-by":"publisher","first-page":"1163","DOI":"10.1523\/JNEUROSCI.4415-07.2008","article-title":"Primary motor cortex tuning to intended movement kinematics in humans with tetraplegia","volume":"28","author":"Truccolo","year":"2008","journal-title":"The Journal of Neuroscience"},{"key":"bib112","doi-asserted-by":"publisher","DOI":"10.1088\/1741-2560\/11\/5\/056006","article-title":"Useful properties of spinal circuits for learning and performing planar reaches","volume":"11","author":"Tsianos","year":"2014","journal-title":"Journal of Neural Engineering"},{"key":"bib113","doi-asserted-by":"publisher","first-page":"1286","DOI":"10.1016\/j.celrep.2018.03.137","article-title":"Corticospinal circuits from the sensory and motor cortices differentially regulate skilled movements through distinct spinal interneurons","volume":"23","author":"Ueno","year":"2018","journal-title":"Cell Reports"},{"key":"bib114","doi-asserted-by":"publisher","first-page":"619","DOI":"10.1007\/978-0-387-77064-2_33","article-title":"A mathematical approach to the mechanical capabilities of limbs and fingers","volume":"629","author":"Valero-Cuevas","year":"2009","journal-title":"Advances in Experimental Medicine and Biology"},{"key":"bib115","doi-asserted-by":"publisher","DOI":"10.3389\/fninf.2019.00018","article-title":"Draculab: a python simulator for firing rate neural networks with delayed adaptive connections","volume":"13","author":"Verduzco-Flores","year":"2019","journal-title":"Frontiers in Neuroinformatics"},{"key":"bib116","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1016\/j.neunet.2022.03.002","article-title":"A differential Hebbian framework for biologically-plausible motor control","volume":"150","author":"Verduzco-Flores","year":"2022","journal-title":"Neural Networks"},{"key":"bib117","doi-asserted-by":"publisher","first-page":"440","DOI":"10.1016\/j.cell.2017.08.014","article-title":"Deconstruction of corticospinal circuits for goal-directed motor skills","volume":"171","author":"Wang","year":"2017","journal-title":"Cell"},{"key":"bib118","doi-asserted-by":"publisher","first-page":"196","DOI":"10.1016\/0006-8993(83)91059-4","article-title":"Adaptive plasticity in the spinal stretch reflex","volume":"267","author":"Wolpaw","year":"1983","journal-title":"Brain Research"},{"key":"bib119","doi-asserted-by":"publisher","first-page":"588","DOI":"10.1016\/s0166-2236(97)01133-8","article-title":"The complex structure of a simple memory","volume":"20","author":"Wolpaw","year":"1997","journal-title":"Trends in Neurosciences"},{"key":"bib120","doi-asserted-by":"publisher","first-page":"388","DOI":"10.1016\/j.appet.2006.09.015","article-title":"Homeostasis: beyond curt Richter","volume":"49","author":"Woods","year":"2007","journal-title":"Appetite"},{"key":"bib121","doi-asserted-by":"publisher","first-page":"27655","DOI":"10.1073\/pnas.1919253117","article-title":"Forelimb force direction and magnitude independently controlled by spinal modules in the macaque","volume":"117","author":"Yaron","year":"2020","journal-title":"PNAS"},{"key":"bib122","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1006227","article-title":"General differential Hebbian learning: capturing temporal relations between events in neural networks and the brain","volume":"14","author":"Zappacosta","year":"2018","journal-title":"PLOS Computational Biology"},{"key":"bib123","doi-asserted-by":"publisher","first-page":"3432","DOI":"10.1523\/JNEUROSCI.1979-20.2021","article-title":"Differential contribution of V0 interneurons to execution of rhythmic and nonrhythmic motor behaviors","volume":"41","author":"Zelenin","year":"2021","journal-title":"The Journal of Neuroscience"}],"container-title":["eLife"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/cdn.elifesciences.org\/articles\/77216\/elife-77216-v3.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/cdn.elifesciences.org\/articles\/77216\/elife-77216-v3.xml","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/elifesciences.org\/articles\/77216","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,10,11]],"date-time":"2023-10-11T21:16:53Z","timestamp":1697059013000},"score":1,"resource":{"primary":{"URL":"https:\/\/elifesciences.org\/articles\/77216"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,11,14]]},"references-count":123,"alternative-id":["10.7554\/eLife.77216"],"URL":"https:\/\/doi.org\/10.7554\/elife.77216","archive":["CLOCKSS"],"relation":{"has-review":[{"id-type":"doi","id":"10.7554\/eLife.77216.sa2","asserted-by":"object"},{"id-type":"doi","id":"10.7554\/eLife.77216.sa1","asserted-by":"object"},{"id-type":"doi","id":"10.7554\/eLife.77216.sa0","asserted-by":"object"}]},"ISSN":["2050-084X"],"issn-type":[{"value":"2050-084X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,11,14]]},"assertion":[{"value":"single anonymised","name":"peer_review_transparency","label":"Peer review transparency","group":{"name":"peer_review_taxonomy"}},{"value":"other reviewer(s), editor","name":"peer_review_interaction","label":"Peer review interaction","group":{"name":"peer_review_taxonomy"}},{"value":"review summaries, review reports, author\/editor communication, reviewer identities reviewer opt in, editor identities","name":"peer_review_published","label":"Peer review published","group":{"name":"peer_review_taxonomy"}},{"value":"open (sign in with ORCID iD required)","name":"post_publication_commenting","label":"Post publication commenting","group":{"name":"post_publication_commenting"}}],"article-number":"e77216"}}