{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,11]],"date-time":"2026-07-11T03:07:08Z","timestamp":1783739228756,"version":"3.55.0"},"reference-count":48,"publisher":"MDPI AG","issue":"2","license":[{"start":{"date-parts":[[2026,1,29]],"date-time":"2026-01-29T00:00:00Z","timestamp":1769644800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Pavlov Institute of Physiology, Russian Academy of Sciences","award":["1021062411782-5-3.1.8"],"award-info":[{"award-number":["1021062411782-5-3.1.8"]}]},{"name":"Pavlov Institute of Physiology, Russian Academy of Sciences","award":["2025-10-01"],"award-info":[{"award-number":["2025-10-01"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Life"],"abstract":"<jats:p>(1) Background: We developed a novel technology that regulates human locomotion using transcutaneous electrical spinal cord stimulation to activate spinal locomotor networks and posterior root stimulation to activate leg flexor and extensor motor pools during swing and stance phases, respectively. This technology effectively restores walking in post-stroke individuals while forward propulsion in the stance phase and foot dorsiflexion in the swing phase are insufficient. In this study the effectiveness of regulating the stance and swing phases while healthy volunteers walked on a treadmill with transcutaneous electrical stimulation of the posterior roots, leg muscles, and their combined effects has been examined. (2) Methods: We analyzed the kinematic characteristics of stepping movements in healthy participants with spinal stimulation of the posterior roots and flexor\/extensor leg muscles. (3) Results: Our findings clearly show that posterior root stimulation at T12 combined with tibialis anterior muscle stimulation during the swing phase effectively regulates foot dorsiflexion, whereas posterior root stimulation at L2 combined with hamstrings and medial gastrocnemius stimulation during the stance phase effectively regulates forward propulsion. (4) Conclusions: Combined stimulation in the stance and swing phases within the same gait cycle resulted in the most coordinated stepping, and effective control of forward propulsion and foot dorsiflexion.<\/jats:p>","DOI":"10.3390\/life16020226","type":"journal-article","created":{"date-parts":[[2026,1,29]],"date-time":"2026-01-29T10:56:34Z","timestamp":1769684194000},"page":"226","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Modulation of Forward Propulsion and Foot Dorsiflexion by Spinal and Muscular Stimulation During Human Stepping"],"prefix":"10.3390","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9757-7946","authenticated-orcid":false,"given":"Sergey","family":"Ananyev","sequence":"first","affiliation":[{"name":"Pavlov Institute of Physiology, Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0003-7810-8396","authenticated-orcid":false,"given":"Ivan","family":"Sakun","sequence":"additional","affiliation":[{"name":"Pavlov Institute of Physiology, Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5948-0991","authenticated-orcid":false,"given":"Vsevolod","family":"Lyakhovetskii","sequence":"additional","affiliation":[{"name":"Pavlov Institute of Physiology, Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexander","family":"Grishin","sequence":"additional","affiliation":[{"name":"Pavlov Institute of Physiology, Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8934-5253","authenticated-orcid":false,"given":"Tatiana","family":"Moshonkina","sequence":"additional","affiliation":[{"name":"Pavlov Institute of Physiology, Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yury","family":"Gerasimenko","sequence":"additional","affiliation":[{"name":"Pavlov Institute of Physiology, Russian Academy of Sciences, 199034 St. Petersburg, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,1,29]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1016\/bs.irn.2019.08.003","article-title":"The Effects and Potential Mechanisms of Locomotor Training on Improvements of Functional Recovery after Spinal Cord Injury","volume":"Volume 147","author":"Yu","year":"2019","journal-title":"International Review of Neurobiology"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"405","DOI":"10.4103\/1673-5374.245951","article-title":"Rehabilitation Following Spinal Cord Injury: How Animal Models Can Help Our Understanding of Exercise-Induced Neuroplasticity","volume":"14","author":"Loy","year":"2019","journal-title":"Neural Regen. Res."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"114178","DOI":"10.1016\/j.expneurol.2022.114178","article-title":"Activity-Dependent Plasticity and Spinal Cord Stimulation for Motor Recovery Following Spinal Cord Injury","volume":"357","author":"Samejima","year":"2022","journal-title":"Exp. Neurol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1016\/S0966-6362(97)00042-8","article-title":"Neural Control of Locomotion; Part 1: The Central Pattern Generator from Cats to Humans","volume":"7","author":"Duysens","year":"1998","journal-title":"Gait Posture"},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"360","DOI":"10.1111\/j.1749-6632.1998.tb09062.x","article-title":"Evidence for a Spinal Central Pattern Generator in Humans","volume":"860","author":"Dimitrijevic","year":"1998","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"1379","DOI":"10.1016\/S1388-2457(03)00120-2","article-title":"Spinal Cord Pattern Generators for Locomotion","volume":"114","author":"Dietz","year":"2003","journal-title":"Clin. Neurophysiol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"158","DOI":"10.1134\/S0362119712020065","article-title":"Transcutaneous Electrical Stimulation of the Spinal Cord: A Noninvasive Tool for the Activation of Stepping Pattern Generators in Humans","volume":"38","author":"Gorodnichev","year":"2012","journal-title":"Hum. Physiol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"225","DOI":"10.1016\/j.rehab.2015.05.003","article-title":"Transcutaneous Electrical Spinal-Cord Stimulation in Humans","volume":"58","author":"Gerasimenko","year":"2015","journal-title":"Ann. Phys. Rehabil. Med."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"7802180220","DOI":"10.5014\/ajot.2024.050533","article-title":"Spinal Cord Stimulation for Poststroke Hemiparesis: A Scoping Review","volume":"78","author":"Allen","year":"2024","journal-title":"Am. J. Occup. Ther."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1016\/j.apmr.2022.01.161","article-title":"Spinal Cord Stimulation Research in the Restoration of Motor, Sensory, and Autonomic Function for Individuals Living With Spinal Cord Injuries: A Scoping Review","volume":"103","author":"Laskin","year":"2022","journal-title":"Arch. Phys. Med. Rehabil."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1016\/j.neurom.2023.03.014","article-title":"Targeted Selection of Stimulation Parameters for Restoration of Motor and Autonomic Function in Individuals With Spinal Cord Injury","volume":"27","author":"Angeli","year":"2024","journal-title":"Neuromodul. Technol. Neural Interface"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1677","DOI":"10.1038\/s41591-018-0175-7","article-title":"Neuromodulation of Lumbosacral Spinal Networks Enables Independent Stepping after Complete Paraplegia","volume":"24","author":"Gill","year":"2018","journal-title":"Nat. Med."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1038\/s41586-018-0649-2","article-title":"Targeted Neurotechnology Restores Walking in Humans with Spinal Cord Injury","volume":"563","author":"Wagner","year":"2018","journal-title":"Nature"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"429","DOI":"10.3727\/096368915X686904","article-title":"Electrical Stimulation and Motor Recovery","volume":"24","author":"Young","year":"2015","journal-title":"Cell Transpl."},{"key":"ref_15","doi-asserted-by":"crossref","unstructured":"Moshonkina, T., Grishin, A., Bogacheva, I., Gorodnichev, R., Ovechkin, A., Siu, R., Edgerton, V.R., and Gerasimenko, Y. (2021). Novel Non-Invasive Strategy for Spinal Neuromodulation to Control Human Locomotion. Front. Hum. Neurosci., 14.","DOI":"10.3389\/fnhum.2020.622533"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1038\/sc.2010.62","article-title":"Muscle Fatigue Characteristics in Paralyzed Muscle after Spinal Cord Injury","volume":"49","author":"Pelletier","year":"2011","journal-title":"Spinal Cord"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"454","DOI":"10.1152\/jn.00367.2019","article-title":"Changes in Motoneuron Excitability during Voluntary Muscle Activity in Humans with Spinal Cord Injury","volume":"123","author":"Vastano","year":"2020","journal-title":"J. Neurophysiol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"783","DOI":"10.1177\/1545968310372091","article-title":"Changes in Gait Symmetry and Velocity After Stroke: A Cross-Sectional Study From Weeks to Years After Stroke","volume":"24","author":"Patterson","year":"2010","journal-title":"Neurorehabilit. Neural Repair"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"409","DOI":"10.3109\/02699052.2014.995231","article-title":"Associations between Lower Limb Strength and Gait Velocity Following Stroke: A Systematic Review","volume":"29","author":"Mentiplay","year":"2015","journal-title":"Brain Inj."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"401","DOI":"10.1038\/sj.sc.3101615","article-title":"Stepping-like Movements in Humans with Complete Spinal Cord Injury Induced by Epidural Stimulation of the Lumbar Cord: Electromyographic Study of Compound Muscle Action Potentials","volume":"42","author":"Minassian","year":"2004","journal-title":"Spinal Cord"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"E176","DOI":"10.1111\/aor.12615","article-title":"Augmentation of Voluntary Locomotor Activity by Transcutaneous Spinal Cord Stimulation in Motor-Incomplete Spinal Cord-Injured Individuals","volume":"39","author":"Hofstoetter","year":"2015","journal-title":"Artif. Organs"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1805","DOI":"10.2174\/1381612822666161214144655","article-title":"Probing the Human Spinal Locomotor Circuits by Phasic Step-Induced Feedback and by Tonic Electrical and Pharmacological Neuromodulation","volume":"23","author":"Hofstoetter","year":"2017","journal-title":"Curr. Pharm. Des."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"1435","DOI":"10.1089\/neu.2018.5956","article-title":"Self-Assisted Standing Enabled by Non-Invasive Spinal Stimulation after Spinal Cord Injury","volume":"36","author":"Sayenko","year":"2019","journal-title":"J. Neurotrauma"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"5990","DOI":"10.1038\/s41598-024-56579-0","article-title":"Tapping into the Human Spinal Locomotor Centres with Transspinal Stimulation","volume":"14","author":"Skiadopoulos","year":"2024","journal-title":"Sci. Rep."},{"key":"ref_25","unstructured":"Ib\u00e1\u00f1ez, J., Gonz\u00e1lez-Vargas, J., Azor\u00edn, J.M., Akay, M., and Pons, J.L. (2017). A Preliminary Test of a Portable Prototype System of FES Foot Drop Correction and Gait Measurements with a Hemiplegic Subject. Proceedings of the Converging Clinical and Engineering Research on Neurorehabilitation II, Springer International Publishing."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"223","DOI":"10.1007\/s00421-010-1502-y","article-title":"Physiological and Methodological Considerations for the Use of Neuromuscular Electrical Stimulation","volume":"110","author":"Maffiuletti","year":"2010","journal-title":"Eur. J. Appl. Physiol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1007\/s10527-021-10029-7","article-title":"A System for Detecting Stepping Cycle Phases and Spinal Cord Stimulation as a Tool for Controlling Human Locomotion","volume":"54","author":"Grishin","year":"2021","journal-title":"Biomed. Eng."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s10462-016-9514-6","article-title":"Recent Developments in Human Gait Research: Parameters, Approaches, Applications, Machine Learning Techniques, Datasets and Challenges","volume":"49","author":"Prakash","year":"2018","journal-title":"Artif. Intell. Rev."},{"key":"ref_29","doi-asserted-by":"crossref","unstructured":"Anwary, A.R., Yu, H., and Vassallo, M. (2018). An Automatic Gait Feature Extraction Method for Identifying Gait Asymmetry Using Wearable Sensors. Sensors, 18.","DOI":"10.3390\/s18020676"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"426","DOI":"10.1044\/2014_JSLHR-S-13-0007","article-title":"Accuracy and Precision of a Custom Camera-Based System for 2-D and 3-D Motion Tracking During Speech and Nonspeech Motor Tasks","volume":"57","author":"Feng","year":"2014","journal-title":"J. Speech Lang. Hear. Res."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"560","DOI":"10.1016\/j.gaitpost.2006.12.018","article-title":"A New Anatomically Based Protocol for Gait Analysis in Children","volume":"26","author":"Leardini","year":"2007","journal-title":"Gait Posture"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"536","DOI":"10.1097\/WCO.0000000000001080","article-title":"Historical Development and Contemporary Use of Neuromodulation in Human Spinal Cord Injury","volume":"35","author":"Harkema","year":"2022","journal-title":"Curr. Opin. Neurol."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1968","DOI":"10.1089\/neu.2015.4008","article-title":"Noninvasive Reactivation of Motor Descending Control after Paralysis","volume":"32","author":"Gerasimenko","year":"2015","journal-title":"J. Neurotrauma"},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"60","DOI":"10.1134\/S0362119721010059","article-title":"Regulation of Gait Cycle Phases during Noninvasive Electrical Stimulation of the Spinal Cord","volume":"47","author":"Gorodnichev","year":"2021","journal-title":"Hum. Physiol."},{"key":"ref_35","doi-asserted-by":"crossref","unstructured":"Siu, R., Brown, E.H., Mesbah, S., Gonnelli, F., Pisolkar, T., Edgerton, V.R., Ovechkin, A.V., and Gerasimenko, Y.P. (2022). Novel Noninvasive Spinal Neuromodulation Strategy Facilitates Recovery of Stepping after Motor Complete Paraplegia. J. Clin. Med., 11.","DOI":"10.3390\/jcm11133670"},{"key":"ref_36","first-page":"46","article-title":"Control of Walking Cycle Using Noninvasive Electrical Stimulation of the Spinal Cord and Muscles","volume":"51","author":"Moshonkina","year":"2025","journal-title":"Hum. Physiol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"21351","DOI":"10.1038\/s41598-021-00658-z","article-title":"Predictive Simulation of Post-Stroke Gait with Functional Electrical Stimulation","volume":"11","author":"Santos","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"101388","DOI":"10.1016\/j.rehab.2020.03.012","article-title":"Functional Electrical Stimulation of the Peroneal Nerve Improves Post-Stroke Gait Speed When Combined with Physiotherapy. A Systematic Review and Meta-Analysis","volume":"64","author":"Rech","year":"2021","journal-title":"Ann. Phys. Rehabil. Med."},{"key":"ref_39","first-page":"417","article-title":"Functional Electrical Stimulation for Gait Correction in the Early Recovery Phase after Ischemic Stroke","volume":"27","author":"Skvortsov","year":"2025","journal-title":"Extreme Med."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1105","DOI":"10.1080\/17434440.2025.2546476","article-title":"Advancing Gait Rehabilitation through Wearable Technologies: Current Landscape and Future Directions","volume":"22","author":"Bartloff","year":"2025","journal-title":"Expert Rev. Med. Devices"},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"372","DOI":"10.1159\/000101161","article-title":"Electrical Stimulation of the Cervical Spinal Cord Increases Cerebral Blood Flow in Humans","volume":"48","author":"Hosobuchi","year":"1985","journal-title":"Stereotact. Funct. Neurosurg."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"262","DOI":"10.1159\/000066729","article-title":"Spinal Cord Stimulation and Cerebral Blood Flow in Stroke: Personal Experience","volume":"76","author":"Visocchi","year":"2002","journal-title":"Stereotact. Funct. Neurosurg."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"893","DOI":"10.1007\/s11910-023-01319-6","article-title":"Neuromodulation for Post-Stroke Motor Recovery: A Narrative Review of Invasive and Non-Invasive Tools","volume":"23","author":"Keser","year":"2023","journal-title":"Curr. Neurol. Neurosci. Rep."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"100566","DOI":"10.1016\/j.cobme.2024.100566","article-title":"Rehabilitation of Motor and Sensory Function Using Spinal Cord Stimulation: Recent Advances","volume":"32","author":"Iversen","year":"2024","journal-title":"Curr. Opin. Biomed. Eng."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1134\/S1607672922340087","article-title":"A New Technology for Recovery of Locomotion in Patients after a Stroke","volume":"507","author":"Moshonkina","year":"2022","journal-title":"Dokl. Biochem. Biophys."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"3524","DOI":"10.1038\/s41467-019-11244-3","article-title":"Brainstem and Spinal Cord MRI Identifies Altered Sensorimotor Pathways Post-Stroke","volume":"10","author":"Karbasforoushan","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1111\/aor.14660","article-title":"The Effects of Transcutaneous Spinal Cord Stimulation Delivered with and without High-frequency Modulation on Spinal and Corticospinal Excitability","volume":"48","author":"Massey","year":"2024","journal-title":"Artif. Organs"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"100090","DOI":"10.1016\/j.ynirp.2022.100090","article-title":"Transcutaneous Spinal Stimulation Alters Cortical and Subcortical Activation Patterns during Mimicked-Standing: A Proof-of-Concept fMRI Study","volume":"2","author":"Manson","year":"2022","journal-title":"Neuroimage Rep."}],"container-title":["Life"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2075-1729\/16\/2\/226\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,29]],"date-time":"2026-01-29T11:05:27Z","timestamp":1769684727000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2075-1729\/16\/2\/226"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,1,29]]},"references-count":48,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2026,2]]}},"alternative-id":["life16020226"],"URL":"https:\/\/doi.org\/10.3390\/life16020226","relation":{},"ISSN":["2075-1729"],"issn-type":[{"value":"2075-1729","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,1,29]]}}}