{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,27]],"date-time":"2026-10-27T15:51:12Z","timestamp":1793116272574,"version":"4.5.0"},"reference-count":98,"publisher":"eLife Sciences Publications, Ltd","license":[{"start":{"date-parts":[[2026,2,17]],"date-time":"2026-02-17T00:00:00Z","timestamp":1771286400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,2,17]],"date-time":"2026-02-17T00:00:00Z","timestamp":1771286400000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,2,17]],"date-time":"2026-02-17T00:00:00Z","timestamp":1771286400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["R01 NS112304"],"award-info":[{"award-number":["R01 NS112304"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["R01 NS112304"],"award-info":[{"award-number":["R01 NS112304"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["R01 NS115900"],"award-info":[{"award-number":["R01 NS115900"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["R01 NS115900"],"award-info":[{"award-number":["R01 NS115900"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["R01 NS130799"],"award-info":[{"award-number":["R01 NS130799"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["R01 NS130799"],"award-info":[{"award-number":["R01 NS130799"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["T32 NS121768"],"award-info":[{"award-number":["T32 NS121768"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["T32 NS121768"],"award-info":[{"award-number":["T32 NS121768"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"name":"Kentucky Spinal Cord Injury Research Trust"}],"content-domain":{"domain":["elifesciences.org"],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:p>Rats exhibit significant recovery of locomotor function following incomplete spinal cord injuries, albeit with altered gait expression and reduced speed and stepping frequency. These changes likely result from and give insight into the reorganization within spared and injured spinal circuitry. Previously, we developed computational models of the mouse spinal locomotor circuitry controlling speed-dependent gait expression (Danner et al., 2017; Zhang et al., 2022). Here, we adapted these models to the rat and used the adapted model to explore potential circuit-level changes underlying altered gait expression observed after recovery from two different thoracic spinal cord injuries (lateral hemisection and contusion) that have roughly comparable levels of locomotor recovery (Danner et al., 2023). The model reproduced experimentally observed gait expression before injury and after recovery from lateral hemisection and contusion and suggests two distinct, injury-specific routes to restored function. First, recovery after lateral hemisection required substantial functional restoration of damaged descending drive and long propriospinal connections, suggesting compensatory plasticity through formation of detour pathways. Second, recovery after a moderate midline contusion predominantly relied on reorganization of spared sublesional networks and altered control of supralesional cervical circuits, compensating for weakened propriospinal and descending pathways. Despite these differences, sensitivity analysis revealed that restored activation of sublesional lumbar rhythm-generating circuits and appropriately balanced lumbar commissural connectivity are the key determinants of post-injury gait expression, suggesting that injury symmetry shapes how the cord reorganizes, but effective recovery in both cases depends on re-engaging these lumbar networks, which makes them prime targets for therapeutic intervention.<\/jats:p>","DOI":"10.7554\/elife.107480","type":"journal-article","created":{"date-parts":[[2025,7,8]],"date-time":"2025-07-08T10:30:55Z","timestamp":1751970655000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.107480","source":"Crossref","is-referenced-by-count":4,"title":["Linking spinal circuit reorganization to recovery after thoracic spinal cord injury"],"prefix":"10.7554","volume":"14","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1971-9707","authenticated-orcid":true,"given":"Natalia A","family":"Shevtsova","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04bdffz58","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Neurobiology and Anatomy, Drexel University College of Medicine","place":["Philadelphia, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7141-1347","authenticated-orcid":true,"given":"Andrew B","family":"Lockhart","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04bdffz58","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Neurobiology and Anatomy, Drexel University College of Medicine","place":["Philadelphia, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"validation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3461-349X","authenticated-orcid":true,"given":"Ilya A","family":"Rybak","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04bdffz58","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Neurobiology and Anatomy, Drexel University College of Medicine","place":["Philadelphia, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3816-3676","authenticated-orcid":true,"given":"David SK","family":"Magnuson","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/01ckdn478","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Neurological Surgery, University of Louisville School of Medicine, Health Sciences Campus","place":["Louisville, United States"]},{"id":[{"id":"https:\/\/ror.org\/01ckdn478","id-type":"ROR","asserted-by":"publisher"}],"name":"Kentucky Spinal Cord Injury Research Center, University of Louisville School of Medicine, Health Sciences Campus","place":["Louisville, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4642-7064","authenticated-orcid":true,"given":"Simon M","family":"Danner","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/04bdffz58","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Neurobiology and Anatomy, Drexel University College of Medicine","place":["Philadelphia, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"corresponding-author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]}],"member":"4374","published-online":{"date-parts":[[2026,2,17]]},"reference":[{"key":"bib1","doi-asserted-by":"publisher","first-page":"2212","DOI":"10.1152\/jn.00768.2011","type":"journal-article","article-title":"Serotonin modulates multiple calcium current subtypes in commissural interneurons of the neonatal mouse","volume":"107","author":"Abbinanti","year":"2012","journal-title":"Journal of Neurophysiology"},{"key":"bib2","doi-asserted-by":"publisher","first-page":"2191","DOI":"10.1152\/jn.00336.2012","type":"journal-article","article-title":"Postnatal emergence of serotonin-induced plateau potentials in commissural interneurons of the mouse spinal cord","volume":"108","author":"Abbinanti","year":"2012","journal-title":"Journal of Neurophysiology"},{"key":"bib3","doi-asserted-by":"publisher","first-page":"1586","DOI":"10.1063\/1.3637934","type":"conference-paper","article-title":"Odeint \u2013 Solving Ordinary Differential Equations in C++","author":"Ahnert","year":"2011","unstructured":"Ahnert K, Mulansky M, Simos TE, Psihoyios G, Tsitouras C, Anastassi Z. 2011. Odeint \u2013 Solving Ordinary Differential Equations in C++. Numerical analysis and applied mathematics icnaam 2011."},{"key":"bib4","doi-asserted-by":"publisher","first-page":"16877","DOI":"10.1073\/pnas.1419045111","type":"journal-article","article-title":"Degradation of mouse locomotor pattern in the absence of proprioceptive sensory feedback","volume":"111","author":"Akay","year":"2014","journal-title":"PNAS"},{"key":"bib5","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1007\/s00422-013-0546-6","type":"journal-article","article-title":"Contributions of phase resetting and interlimb coordination to the adaptive control of hindlimb obstacle avoidance during locomotion in rats: a simulation study","volume":"107","author":"Aoi","year":"2013","journal-title":"Biological Cybernetics"},{"key":"bib6","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.43587","type":"journal-article","article-title":"Computational modeling of brainstem circuits controlling locomotor frequency and gait","volume":"8","author":"Ausborn","year":"2019","journal-title":"eLife"},{"key":"bib7","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22136835","type":"journal-article","article-title":"Computational modeling of spinal locomotor circuitry in the age of molecular genetics","volume":"22","author":"Ausborn","year":"2021","journal-title":"International Journal of Molecular Sciences"},{"key":"bib8","doi-asserted-by":"publisher","first-page":"1988","DOI":"10.1111\/j.1460-9568.2006.04726.x","type":"journal-article","article-title":"Spontaneous locomotor recovery in spinal cord injured rats is accompanied by anatomical plasticity of reticulospinal fibers","volume":"23","author":"Ballermann","year":"2006","journal-title":"The European Journal of Neuroscience"},{"key":"bib9","doi-asserted-by":"publisher","first-page":"1727","DOI":"10.1046\/j.0953-816x.2001.01794.x","type":"journal-article","article-title":"Forelimb locomotor generators and quadrupedal locomotion in the neonatal rat","volume":"14","author":"Ballion","year":"2001","journal-title":"The European Journal of Neuroscience"},{"key":"bib10","doi-asserted-by":"publisher","first-page":"84","DOI":"10.1016\/0006-8993(87)91442-9","type":"journal-article","article-title":"Recovery of locomotion after chronic spinalization in the adult cat","volume":"412","author":"Barbeau","year":"1987","journal-title":"Brain Research"},{"key":"bib11","doi-asserted-by":"publisher","first-page":"3976","DOI":"10.1523\/JNEUROSCI.5692-07.2008","type":"journal-article","article-title":"Prominent role of the spinal central pattern generator in the recovery of locomotion after partial spinal cord injuries","volume":"28","author":"Barri\u00e8re","year":"2008","journal-title":"The Journal of Neuroscience"},{"key":"bib12","doi-asserted-by":"publisher","first-page":"244","DOI":"10.1006\/exnr.1996.0098","type":"journal-article","article-title":"Graded histological and locomotor outcomes after spinal cord contusion using the NYU weight-drop device versus transection","volume":"139","author":"Basso","year":"1996","journal-title":"Experimental Neurology"},{"key":"bib13","doi-asserted-by":"publisher","first-page":"1426","DOI":"10.1016\/j.cub.2015.04.005","type":"journal-article","article-title":"Phenotypic characterization of speed-associated gait changes in mice reveals modular organization of locomotor networks","volume":"25","author":"Bellardita","year":"2015","journal-title":"Current Biology"},{"key":"bib14","doi-asserted-by":"crossref","DOI":"10.1002\/9781119113713","type":"book","volume-title":"Understanding Mammalian Locomotion: Concepts and Applications","author":"Bertram","year":"2016"},{"key":"bib15","doi-asserted-by":"publisher","first-page":"1047","DOI":"10.1016\/j.neuron.2013.01.026","type":"journal-article","article-title":"Activity-dependent changes in extracellular Ca2+ and K+ reveal pacemakers in the spinal locomotor-related network","volume":"77","author":"Brocard","year":"2013","journal-title":"Neuron"},{"key":"bib16","doi-asserted-by":"publisher","DOI":"10.1016\/j.expneurol.2023.114631","type":"journal-article","article-title":"Long ascending propriospinal neurons are heterogenous and subject to spinal cord injury induced anatomic plasticity","volume":"373","author":"Brown","year":"2024","journal-title":"Experimental Neurology"},{"key":"bib17","doi-asserted-by":"publisher","first-page":"S137","DOI":"10.1097\/00007632-200112151-00023","type":"journal-article","article-title":"Establishing prognosis and maximizing functional outcomes after spinal cord injury","volume":"26","author":"Burns","year":"2001","journal-title":"Spine"},{"key":"bib18","doi-asserted-by":"publisher","first-page":"953","DOI":"10.1016\/s0896-6273(03)00353-2","type":"journal-article","article-title":"Functional identification of interneurons responsible for left-right coordination of hindlimbs in mammals","volume":"38","author":"Butt","year":"2003","journal-title":"Neuron"},{"key":"bib19","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1038\/nature25448","type":"journal-article","article-title":"Midbrain circuits that set locomotor speed and gait selection","volume":"553","author":"Caggiano","year":"2018","journal-title":"Nature"},{"key":"bib20","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1038\/nature24064","type":"journal-article","article-title":"Locomotor speed control circuits in the caudal brainstem","volume":"551","author":"Capelli","year":"2017","journal-title":"Nature"},{"key":"bib21","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1145\/79505.79507","type":"journal-article","article-title":"A variable order Runge-Kutta method for initial value problems with rapidly varying right-hand sides","volume":"16","author":"Cash","year":"1990","journal-title":"ACM Transactions on Mathematical Software"},{"key":"bib22","doi-asserted-by":"publisher","first-page":"785","DOI":"10.1145\/2939672.2939785","type":"conference-paper","article-title":"XGBoost: a scalable tree boosting system","author":"Chen","year":"2016","unstructured":"Chen T, Guestrin C. 2016. XGBoost: a scalable tree boosting system. Proceedings of the 22nd ACM SIGKDD International Conference on Knowledge Discovery and Data Mining."},{"key":"bib23","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1002\/jmor.1051460202","type":"journal-article","article-title":"Muscle activity in rat locomotion: movement analysis and electromyography of the flexors and extensors of the elbow","volume":"146","author":"Cohen","year":"1975","journal-title":"Journal of Morphology"},{"key":"bib24","doi-asserted-by":"publisher","first-page":"142","DOI":"10.55782\/ane-2014-1981","type":"journal-article","article-title":"Recovery of locomotion after partial spinal cord lesions in cats: assessment using behavioral, electrophysiological and imaging techniques","volume":"74","author":"Cohen-Adad","year":"2014","journal-title":"Acta Neurobiologiae Experimentalis"},{"key":"bib25","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1038\/nm1682","type":"journal-article","article-title":"Recovery of supraspinal control of stepping via indirect propriospinal relay connections after spinal cord injury","volume":"14","author":"Courtine","year":"2008","journal-title":"Nature Medicine"},{"key":"bib26","doi-asserted-by":"publisher","first-page":"7098","DOI":"10.1523\/JNEUROSCI.1206-09.2009","type":"journal-article","article-title":"In mice lacking V2a interneurons, gait depends on speed of locomotion","volume":"29","author":"Crone","year":"2009","journal-title":"The Journal of Neuroscience"},{"key":"bib27","doi-asserted-by":"publisher","first-page":"577","DOI":"10.1093\/brain\/awu372","type":"journal-article","article-title":"Human spinal locomotor control is based on flexibly organized burst generators","volume":"138","author":"Danner","year":"2015","journal-title":"Brain"},{"key":"bib28","doi-asserted-by":"publisher","first-page":"6947","DOI":"10.1113\/JP272787","type":"journal-article","article-title":"Central control of interlimb coordination and speed-dependent gait expression in quadrupeds","volume":"594","author":"Danner","year":"2016","journal-title":"The Journal of Physiology"},{"key":"bib29","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.31050","type":"journal-article","article-title":"Computational modeling of spinal circuits controlling limb coordination and gaits in quadrupeds","volume":"6","author":"Danner","year":"2017","journal-title":"eLife"},{"key":"bib30","doi-asserted-by":"publisher","DOI":"10.3389\/fncel.2019.00516","type":"journal-article","article-title":"Spinal V3 interneurons and left-right coordination in mammalian locomotion","volume":"13","author":"Danner","year":"2019","journal-title":"Frontiers in Cellular Neuroscience"},{"key":"bib31","doi-asserted-by":"publisher","DOI":"10.1016\/j.expneurol.2023.114496","type":"journal-article","article-title":"Spinal control of locomotion before and after spinal cord injury","volume":"368","author":"Danner","year":"2023","journal-title":"Experimental Neurology"},{"key":"bib32","type":"software","article-title":"CPGNetworkSimulator","volume-title":"Software Heritage","author":"Danner","year":"2025","unstructured":"Danner SM. 2025. CPGNetworkSimulator. Software Heritage. swh:1:rev:bbfb1d8ba71efb398e62aaa78a2cf08178e1bca3. https:\/\/archive.softwareheritage.org\/swh:1:dir:89038effd5c5767eee5283127d5baf8f75c679b6;origin=https:\/\/github.com\/SimonDanner\/CPGNetworkSimulator;visit=swh:1:snp:bab9b0e72113743222c48843a4d7cbec3ba5c396;anchor=swh:1:rev:bbfb1d8ba71efb398e62aaa78a2cf08178e1bca3."},{"key":"bib33","type":"software","article-title":"Rat-sci-locomotion-model","volume-title":"Software Heritage","author":"Danner","year":"2025","unstructured":"Danner SM, Lockhart AB, Shevtsova NA. 2025. Rat-sci-locomotion-model. Software Heritage. swh:1:rev:44cb548014ca88659a2adbfdb45c581cb7c0ab84. https:\/\/archive.softwareheritage.org\/swh:1:dir:1c24ae00e2b0a59ee0b48cabe48c88e6a9e6fddd;origin=https:\/\/github.com\/dannerlab\/rat-sci-locomotion-model;visit=swh:1:snp:fc5f0cb393aef6bcb4353d4e3377cdd627695cd2;anchor=swh:1:rev:44cb548014ca88659a2adbfdb45c581cb7c0ab84."},{"key":"bib34","doi-asserted-by":"publisher","first-page":"76","DOI":"10.1016\/j.brainres.2006.08.063","type":"journal-article","article-title":"Long ascending propriospinal projections from lumbosacral to upper cervical spinal cord in the rat","volume":"1119","author":"Dutton","year":"2006","journal-title":"Brain Research"},{"key":"bib35","doi-asserted-by":"publisher","first-page":"961","DOI":"10.1113\/jphysiol.2003.048009","type":"journal-article","article-title":"Both dorsal horn and lamina VIII interneurones contribute to crossed reflexes from feline group II muscle afferents","volume":"552","author":"Edgley","year":"2003","journal-title":"The Journal of Physiology"},{"key":"bib36","doi-asserted-by":"publisher","DOI":"10.5962\/bhl.title.1850","type":"book","volume-title":"Vorlesungen \u00dcber Den Bau Der Nerv\u00f6sen Centralorgane Des Menschen Und Der Thiere","author":"Edinger","year":"1896"},{"key":"bib37","doi-asserted-by":"publisher","first-page":"8292","DOI":"10.1523\/JNEUROSCI.0474-20.2020","type":"journal-article","article-title":"The gigantocellular reticular nucleus plays a significant role in locomotor recovery after incomplete spinal cord injury","volume":"40","author":"Engmann","year":"2020","journal-title":"The Journal of Neuroscience"},{"key":"bib38","doi-asserted-by":"publisher","first-page":"679","DOI":"10.1162\/neco.1994.6.4.679","type":"journal-article","article-title":"Reduction of conductance-based models with slow synapses to neural nets","volume":"6","author":"Ermentrout","year":"1994","journal-title":"Neural Computation"},{"key":"bib39","doi-asserted-by":"publisher","first-page":"13399","DOI":"10.1523\/JNEUROSCI.0701-14.2014","type":"journal-article","article-title":"Bridging the gap: a reticulo-propriospinal detour bypassing an incomplete spinal cord injury","volume":"34","author":"Filli","year":"2014","journal-title":"The Journal of Neuroscience"},{"key":"bib40","doi-asserted-by":"publisher","first-page":"509","DOI":"10.4103\/1673-5374.155425","type":"journal-article","article-title":"Structural and functional reorganization of propriospinal connections promotes functional recovery after spinal cord injury","volume":"10","author":"Filli","year":"2015","journal-title":"Neural Regeneration Research"},{"key":"bib41","doi-asserted-by":"publisher","DOI":"10.3389\/fnana.2017.00005","type":"journal-article","article-title":"Anatomical and molecular properties of long descending propriospinal neurons in mice","volume":"11","author":"Flynn","year":"2017","journal-title":"Frontiers in Neuroanatomy"},{"key":"bib42","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1111\/j.1748-1716.1980.tb06534.x","type":"journal-article","article-title":"The locomotion of the low spinal cat","volume":"108","author":"Forssberg","year":"1980","journal-title":"II. Interlimb Coordination. Acta Physiol Scand"},{"key":"bib43","doi-asserted-by":"publisher","first-page":"17","DOI":"10.1179\/016164107X251781","type":"journal-article","article-title":"Adaptive changes in the injured spinal cord and their role in promoting functional recovery","volume":"30","author":"Fouad","year":"2008","journal-title":"Neurological Research"},{"key":"bib44","doi-asserted-by":"publisher","first-page":"1189","DOI":"10.1214\/aos\/1013203451","type":"journal-article","article-title":"Greedy function approximation: a gradient boosting machine","volume":"29","author":"Friedman","year":"2001","journal-title":"The Annals of Statistics"},{"key":"bib45","doi-asserted-by":"publisher","first-page":"4617","DOI":"10.1113\/jphysiol.2009.176669","type":"journal-article","article-title":"Asymmetric control of cycle period by the spinal locomotor rhythm generator in the adult cat","volume":"587","author":"Frigon","year":"2009","journal-title":"The Journal of Physiology"},{"key":"bib46","doi-asserted-by":"publisher","first-page":"2224","DOI":"10.1152\/jn.00978.2016","type":"journal-article","article-title":"The neural control of interlimb coordination during mammalian locomotion","volume":"117","author":"Frigon","year":"2017","journal-title":"Journal of Neurophysiology"},{"key":"bib47","doi-asserted-by":"publisher","first-page":"1179","DOI":"10.1002\/j.2040-4603.1981.tb00864.x","type":"book-chapter","article-title":"Control of locomotion in bipeds, tetrapods, and fish","volume-title":"Handb. Physiol. Nerv. Syst. II. Am. Physiol. Soc","author":"Grillner","year":"1981"},{"key":"bib48","first-page":"1","type":"journal-article","article-title":"The stride cycle of the cat: the modelling of locomotion by computerized analysis of automatic recordings","volume":"Suppl 521","author":"Halbertsma","year":"1983","journal-title":"Acta Physiologica Scandinavica"},{"key":"bib49","doi-asserted-by":"publisher","first-page":"766","DOI":"10.2307\/1311182","type":"journal-article","article-title":"The Quadrupedal Gaits of Vertebrates: The timing of leg movements relates to balance, body shape, agility, speed, and energy expenditure","volume":"39","author":"Hildebrand","year":"1989","journal-title":"BioScience"},{"key":"bib50","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-36587-w","type":"journal-article","article-title":"Deconstructing the modular organization and real-time dynamics of mammalian spinal locomotor networks","volume":"14","author":"Hsu","year":"2023","journal-title":"Nature Communications"},{"key":"bib51","doi-asserted-by":"publisher","first-page":"46","DOI":"10.1016\/j.brainresrev.2007.06.022","type":"journal-article","article-title":"Spinal interneuronal networks in the cat: elementary components","volume":"57","author":"Jankowska","year":"2008","journal-title":"Brain Research Reviews"},{"key":"bib52","doi-asserted-by":"publisher","first-page":"401","DOI":"10.1113\/jphysiol.2008.159236","type":"journal-article","article-title":"Commissural interneurons with input from group I and II muscle afferents in feline lumbar segments: neurotransmitters, projections and target cells","volume":"587","author":"Jankowska","year":"2009","journal-title":"The Journal of Physiology"},{"key":"bib53","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1016\/j.brainresrev.2007.07.019","type":"journal-article","article-title":"Descending command systems for the initiation of locomotion in mammals","volume":"57","author":"Jordan","year":"2008","journal-title":"Brain Research Reviews"},{"key":"bib54","doi-asserted-by":"publisher","first-page":"884","DOI":"10.1016\/j.cub.2018.02.007","type":"journal-article","article-title":"Distinct contributions of mesencephalic locomotor region nuclei to locomotor control in the freely behaving mouse","volume":"28","author":"Josset","year":"2018","journal-title":"Current Biology"},{"key":"bib55","doi-asserted-by":"publisher","first-page":"953","DOI":"10.1523\/JNEUROSCI.4640-11.2012","type":"journal-article","article-title":"Cervicolumbar coordination in mammalian quadrupedal locomotion: role of spinal thoracic circuitry and limb sensory inputs","volume":"32","author":"Juvin","year":"2012","journal-title":"The Journal of Neuroscience"},{"key":"bib56","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1016\/0168-0102(90)90042-d","type":"journal-article","article-title":"Chronically isolated lumbar half spinal cord generates locomotor activities in the ipsilateral hindlimb of the cat","volume":"9","author":"Kato","year":"1990","journal-title":"Neuroscience Research"},{"key":"bib57","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1146\/annurev.neuro.29.051605.112910","type":"journal-article","article-title":"Locomotor circuits in the mammalian spinal cord","volume":"29","author":"Kiehn","year":"2006","journal-title":"Annual Review of Neuroscience"},{"key":"bib58","doi-asserted-by":"publisher","first-page":"224","DOI":"10.1038\/nrn.2016.9","type":"journal-article","article-title":"Decoding the organization of spinal circuits that control locomotion","volume":"17","author":"Kiehn","year":"2016","journal-title":"Nature Reviews. Neuroscience"},{"key":"bib59","doi-asserted-by":"publisher","DOI":"10.3389\/fbioe.2022.825149","type":"journal-article","article-title":"Contribution of afferent feedback to adaptive hindlimb walking in cats: a neuromusculoskeletal modeling study","volume":"10","author":"Kim","year":"2022","journal-title":"Frontiers in Bioengineering and Biotechnology"},{"key":"bib60","doi-asserted-by":"publisher","first-page":"9433","DOI":"10.1523\/JNEUROSCI.17-24-09433.1997","type":"journal-article","article-title":"Crossed rhythmic synaptic input to motoneurons during selective activation of the contralateral spinal locomotor network","volume":"17","author":"Kjaerulff","year":"1997","journal-title":"The Journal of Neuroscience"},{"key":"bib61","doi-asserted-by":"publisher","first-page":"178","DOI":"10.1016\/j.expneurol.2010.03.008","type":"journal-article","article-title":"Task-specificity vs. ceiling effect: step-training in shallow water after spinal cord injury","volume":"224","author":"Kuerzi","year":"2010","journal-title":"Experimental Neurology"},{"key":"bib62","doi-asserted-by":"publisher","first-page":"3452","DOI":"10.1016\/j.cub.2023.07.014","type":"journal-article","article-title":"Distinct roles of spinal commissural interneurons in transmission of contralateral sensory information","volume":"33","author":"Laflamme","year":"2023","journal-title":"Current Biology"},{"key":"bib63","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2020.598888","type":"journal-article","article-title":"On the organization of the locomotor CPG: Insights from split-belt locomotion and mathematical modeling","volume":"14","author":"Latash","year":"2020","journal-title":"Frontiers in Neuroscience"},{"key":"bib64","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2016.00042","type":"journal-article","article-title":"Speed-dependent modulation of the locomotor behavior in adult mice reveals attractor and transitional gaits","volume":"10","author":"Lemieux","year":"2016","journal-title":"Frontiers in Neuroscience"},{"key":"bib65","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-45300-4","type":"journal-article","article-title":"Functional plasticity of glutamatergic neurons of medullary reticular nuclei after spinal cord injury in mice","volume":"15","author":"Lemieux","year":"2024","journal-title":"Nature Communications"},{"key":"bib66","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1152\/jn.1948.11.5.455","type":"journal-article","article-title":"Analysis of forelimb-hindlimb reflex activity in acutely decapitate cats","volume":"11","author":"Lloyd","year":"1948","journal-title":"Journal of Neurophysiology"},{"key":"bib67","doi-asserted-by":"publisher","first-page":"421","DOI":"10.1016\/0014-4886(86)90094-4","type":"journal-article","article-title":"Effects of training on the recovery of full-weight-bearing stepping in the adult spinal cat","volume":"92","author":"Lovely","year":"1986","journal-title":"Experimental Neurology"},{"key":"bib68","doi-asserted-by":"publisher","first-page":"235","DOI":"10.1016\/S0079-6123(06)65015-2","type":"journal-article","article-title":"Modeling the mammalian locomotor CPG: insights from mistakes and perturbations","volume":"165","author":"McCrea","year":"2007","journal-title":"Progress in Brain Research"},{"key":"bib69","doi-asserted-by":"publisher","first-page":"134","DOI":"10.1016\/j.brainresrev.2007.08.006","type":"journal-article","article-title":"Organization of mammalian locomotor rhythm and pattern generation","volume":"57","author":"McCrea","year":"2008","journal-title":"Brain Research Reviews"},{"key":"bib70","doi-asserted-by":"publisher","DOI":"10.1098\/rsos.240207","type":"journal-article","article-title":"Sensory feedback and central neuronal interactions in mouse locomotion","volume":"11","author":"Molkov","year":"2024","journal-title":"Royal Society Open Science"},{"key":"bib71","doi-asserted-by":"publisher","first-page":"610","DOI":"10.1093\/brain\/82.4.610","type":"journal-article","article-title":"Fasciculi proprii of the spinal cord in man","volume":"82","author":"Nathan","year":"1959","journal-title":"Brain"},{"key":"bib72","doi-asserted-by":"publisher","first-page":"16","DOI":"10.1093\/icb\/icm024","type":"journal-article","article-title":"Neuromechanics: an integrative approach for understanding motor control","volume":"47","author":"Nishikawa","year":"2007","journal-title":"Integrative and Comparative Biology"},{"key":"bib73","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780198524052.001.0001","type":"book","volume-title":"Neuronal Control of Locomotion: From Mollusc to Man","author":"Orlovsky","year":"1999"},{"key":"bib74","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.conb.2014.12.008","type":"journal-article","article-title":"Serotonergic modulation of spinal motor control","volume":"33","author":"Perrier","year":"2015","journal-title":"Current Opinion in Neurobiology"},{"key":"bib75","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.53565","type":"journal-article","article-title":"Long ascending propriospinal neurons provide flexible, context-specific control of interlimb coordination","volume":"9","author":"Pocratsky","year":"2020","journal-title":"eLife"},{"key":"bib76","doi-asserted-by":"publisher","first-page":"263","DOI":"10.1038\/35067570","type":"journal-article","article-title":"Plasticity of motor systems after incomplete spinal cord injury","volume":"2","author":"Raineteau","year":"2001","journal-title":"Nature Reviews. Neuroscience"},{"key":"bib77","doi-asserted-by":"publisher","first-page":"1195","DOI":"10.1016\/j.neuroscience.2006.07.017","type":"journal-article","article-title":"Inter-enlargement pathways in the ventrolateral funiculus of the adult rat spinal cord","volume":"142","author":"Reed","year":"2006","journal-title":"Neuroscience"},{"key":"bib78","doi-asserted-by":"publisher","first-page":"1505","DOI":"10.1038\/nn.2691","type":"journal-article","article-title":"Extensive spontaneous plasticity of corticospinal projections after primate spinal cord injury","volume":"13","author":"Rosenzweig","year":"2010","journal-title":"Nature Neuroscience"},{"key":"bib79","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1152\/physrev.00028.2005","type":"journal-article","article-title":"Dynamic sensorimotor interactions in locomotion","volume":"86","author":"Rossignol","year":"2006","journal-title":"Physiological Reviews"},{"key":"bib80","doi-asserted-by":"publisher","first-page":"2146","DOI":"10.1152\/jn.90958.2008","type":"journal-article","article-title":"Multiple rhythmic states in a model of the respiratory central pattern generator","volume":"101","author":"Rubin","year":"2009","journal-title":"Journal of Neurophysiology"},{"key":"bib81","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1109\/ICCV.1998.710701","type":"conference-paper","article-title":"A metric for distributions with applications to image databases","author":"Rubner","year":"1998","unstructured":"Rubner Y, Tomasi C, Guibas LJ. 1998. A metric for distributions with applications to image databases. IEEE 6th International Conference on Computer Vision."},{"key":"bib82","doi-asserted-by":"publisher","first-page":"1063","DOI":"10.1016\/j.neuron.2016.10.032","type":"journal-article","article-title":"Long-distance descending spinal neurons ensure quadrupedal locomotor stability","volume":"92","author":"Ruder","year":"2016","journal-title":"Neuron"},{"key":"bib83","doi-asserted-by":"publisher","first-page":"617","DOI":"10.1113\/jphysiol.2006.118703","type":"journal-article","article-title":"Modelling spinal circuitry involved in locomotor pattern generation: insights from deletions during fictive locomotion","volume":"577","author":"Rybak","year":"2006","journal-title":"The Journal of Physiology"},{"key":"bib84","doi-asserted-by":"publisher","DOI":"10.1523\/ENEURO.0069-15.2015","type":"journal-article","article-title":"Organization of the mammalian locomotor CPG: review of computational model and circuit architectures based on genetically identified spinal interneurons(1,2,3)","volume":"2","author":"Rybak","year":"2015","journal-title":"eNeuro"},{"key":"bib85","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.98841.3","type":"journal-article","article-title":"Operation regimes of spinal circuits controlling locomotion and the role of supraspinal drives and sensory feedback","volume":"13","author":"Rybak","year":"2024","journal-title":"eLife"},{"key":"bib86","doi-asserted-by":"publisher","first-page":"259","DOI":"10.1016\/j.cpc.2009.09.018","type":"journal-article","article-title":"Variance based sensitivity analysis of model output. Design and estimator for the total sensitivity index","volume":"181","author":"Saltelli","year":"2010","journal-title":"Computer Physics Communications"},{"key":"bib87","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.70058","type":"journal-article","article-title":"Silencing long ascending propriospinal neurons after spinal cord injury improves hindlimb stepping in the adult rat","volume":"10","author":"Shepard","year":"2021","journal-title":"eLife"},{"key":"bib88","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.82944","type":"journal-article","article-title":"Silencing long-descending inter-enlargement propriospinal neurons improves hindlimb stepping after contusive spinal cord injuries","volume":"12","author":"Shepard","year":"2023","journal-title":"eLife"},{"key":"bib89","doi-asserted-by":"publisher","first-page":"2403","DOI":"10.1113\/JP270121","type":"journal-article","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":"bib90","doi-asserted-by":"publisher","first-page":"271","DOI":"10.1016\/S0378-4754(00)00270-6","type":"journal-article","article-title":"Global sensitivity indices for nonlinear mathematical models and their Monte Carlo estimates","volume":"55","author":"Sobol\u2032","year":"2001","journal-title":"Mathematics and Computers in Simulation"},{"key":"bib91","doi-asserted-by":"publisher","first-page":"1735","DOI":"10.1016\/j.ress.2009.05.007","type":"journal-article","article-title":"Implementation and evaluation of nonparametric regression procedures for sensitivity analysis of computationally demanding models","volume":"94","author":"Storlie","year":"2009","journal-title":"Reliability Engineering & System Safety"},{"key":"bib92","doi-asserted-by":"publisher","first-page":"1626","DOI":"10.1016\/j.cell.2014.11.019","type":"journal-article","article-title":"Muscle spindle feedback directs locomotor recovery and circuit reorganization after spinal cord injury","volume":"159","author":"Takeoka","year":"2014","journal-title":"Cell"},{"key":"bib93","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1016\/j.celrep.2019.03.010","type":"journal-article","article-title":"Functional local proprioceptive feedback circuits initiate and maintain locomotor recovery after spinal cord injury","volume":"27","author":"Takeoka","year":"2019","journal-title":"Cell Reports"},{"key":"bib94","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1038\/nature12286","type":"journal-article","article-title":"Dual-mode operation of neuronal networks involved in left-right alternation","volume":"500","author":"Talpalar","year":"2013","journal-title":"Nature"},{"key":"bib95","doi-asserted-by":"publisher","first-page":"613","DOI":"10.1152\/jn.00316.2007","type":"journal-article","article-title":"Contribution of persistent sodium current to locomotor pattern generation in neonatal rats","volume":"98","author":"Tazerart","year":"2007","journal-title":"Journal of Neurophysiology"},{"key":"bib96","doi-asserted-by":"publisher","first-page":"8577","DOI":"10.1523\/JNEUROSCI.1437-08.2008","type":"journal-article","article-title":"The persistent sodium current generates pacemaker activities in the central pattern generator for locomotion and regulates the locomotor rhythm","volume":"28","author":"Tazerart","year":"2008","journal-title":"The Journal of Neuroscience"},{"key":"bib97","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.73424","type":"journal-article","article-title":"The role of V3 neurons in speed-dependent interlimb coordination during locomotion in mice","volume":"11","author":"Zhang","year":"2022","journal-title":"eLife"},{"key":"bib98","doi-asserted-by":"publisher","first-page":"1716","DOI":"10.1093\/brain\/awu078","type":"journal-article","article-title":"Chasing central nervous system plasticity: the brainstem\u2019s contribution to locomotor recovery in rats with spinal cord injury","volume":"137","author":"Z\u00f6rner","year":"2014","journal-title":"Brain"}],"container-title":["eLife"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/cdn.elifesciences.org\/articles\/107480\/elife-107480-v1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/cdn.elifesciences.org\/articles\/107480\/elife-107480-v1.xml","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/elifesciences.org\/articles\/107480","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T09:11:25Z","timestamp":1787562685000},"score":1,"resource":{"primary":{"URL":"https:\/\/elifesciences.org\/articles\/107480"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,17]]},"references-count":98,"alternative-id":["10.7554\/eLife.107480"],"URL":"https:\/\/doi.org\/10.7554\/elife.107480","archive":["CLOCKSS"],"relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2025.05.17.654682","asserted-by":"subject"}],"is-same-as":[{"id-type":"doi","id":"10.7554\/eLife.107480.3","asserted-by":"subject"}],"is-version-of":[{"id-type":"doi","id":"10.7554\/eLife.107480.1","asserted-by":"subject"},{"id-type":"doi","id":"10.7554\/eLife.107480.2","asserted-by":"subject"}],"is-supplemented-by":[{"id-type":"uri","id":"https:\/\/archive.softwareheritage.org\/swh:1:dir:89038effd5c5767eee5283127d5baf8f75c679b6;origin=https:\/\/github.com\/SimonDanner\/CPGNetworkSimulator;visit=swh:1:snp:bab9b0e72113743222c48843a4d7cbec3ba5c396;anchor=swh:1:rev:bbfb1d8ba71efb398e62aaa78a2cf08178e1bca3","asserted-by":"subject"},{"id-type":"uri","id":"https:\/\/archive.softwareheritage.org\/swh:1:dir:1c24ae00e2b0a59ee0b48cabe48c88e6a9e6fddd;origin=https:\/\/github.com\/dannerlab\/rat-sci-locomotion-model;visit=swh:1:snp:fc5f0cb393aef6bcb4353d4e3377cdd627695cd2;anchor=swh:1:rev:44cb548014ca88659a2adbfdb45c581cb7c0ab84","asserted-by":"subject"}]},"ISSN":["2050-084X"],"issn-type":[{"value":"2050-084X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,17]]},"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":"RP107480"}}