{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T23:57:09Z","timestamp":1789171029783,"version":"build-2803163510"},"reference-count":93,"publisher":"Wiley","issue":"10","license":[{"start":{"date-parts":[[2025,5,5]],"date-time":"2025-05-05T00:00:00Z","timestamp":1746403200000},"content-version":"vor","delay-in-days":4,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc\/4.0\/"}],"content-domain":{"domain":["physoc.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Journal of Physiology"],"published-print":{"date-parts":[[2025,5]]},"abstract":"<jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      It has been suggested that during locomotion, the nervous system controls movement by activating groups of muscles, or muscle synergies. Analysis of muscle synergies can reveal the organization of spinal locomotor networks and how it depends on the state of the nervous system, such as before and after spinal cord injury, and on different locomotor conditions, including a change in speed. The goal of this study was to investigate the effects of spinal transection and locomotor speed on hindlimb muscle synergies and their time\u2010dependent activity patterns in adult cats. EMG activities of 15\u00a0hindlimb muscles were recorded in nine adult cats of either sex during tied\u2010belt treadmill locomotion at speeds of 0.4, 0.7 and 1.0\u00a0m\/s before and after recovery from a low thoracic spinal transection. We determined EMG burst groups using cluster analysis of EMG burst onset and offset times and muscle synergies using non\u2010negative matrix factorization (NNMF). We found five major EMG burst groups and five muscle synergies in each of six experimental conditions (2\u00a0states \u00d7 3\u00a0speeds). In each case, the synergies accounted for at least 90% of muscle EMG variance. Both spinal transection and locomotion speed modified subgroups of EMG burst groups and the composition and activation patterns of selected synergies. However, these changes did not modify the general organization of muscle synergies. Based on the obtained results, we propose an organization for a pattern formation network of a two\u2010level central pattern generator that can be tested in neuromechanical simulations of spinal circuits controlling cat locomotion.\n                      <jats:boxed-text content-type=\"graphic\" position=\"anchor\">\n                        <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" mimetype=\"image\/png\" position=\"anchor\" specific-use=\"enlarged-web-image\" xlink:href=\"graphic\/tjp16685-gra-0001-m.png\">\n                          <jats:alt-text>image<\/jats:alt-text>\n                        <\/jats:graphic>\n                      <\/jats:boxed-text>\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Key points<\/jats:title>\n                    <jats:p>\n                      <jats:list list-type=\"bullet\">\n                        <jats:list-item>\n                          <jats:p>Analysis of muscle synergies during locomotion can reveal the organization of spinal locomotor networks.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>We recorded EMG activity of 15\u00a0hindlimb muscles in cats locomoting on a treadmill at speeds 0.4, 0.7 and 1.0\u00a0m\/s before and after recovery from spinal cord transection at low thoracic level.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>We found five muscle synergies in all six experimental conditions (2\u00a0spinal states x 3\u00a0speeds) that include two flexor synergies operating in the swing phase and three extensor synergies operating in the stance phase.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>Major features of found synergies (the number, muscle composition and activation patterns) were not substantially affected by spinal transection and locomotion speed, suggesting that spinal control mechanism operates muscle synergies.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>Based on the obtained results, we proposed an organization of a pattern formation network of a two\u2010level central pattern generator controlling locomotor activity of hindlimb muscles.<\/jats:p>\n                        <\/jats:list-item>\n                      <\/jats:list>\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1113\/jp288089","type":"journal-article","created":{"date-parts":[[2025,5,5]],"date-time":"2025-05-05T03:51:43Z","timestamp":1746417103000},"page":"3061-3088","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":9,"title":["Effects of spinal transection and locomotor speed on muscle synergies of the cat hindlimb"],"prefix":"10.1113","volume":"603","author":[{"given":"Alexander N.","family":"Klishko","sequence":"first","affiliation":[{"name":"School of Biological Sciences Georgia Institute of Technology  Atlanta GA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jonathan","family":"Harnie","sequence":"additional","affiliation":[{"name":"Department of Pharmacology\u2010Physiology Universit\u00e9 de Sherbrooke  Sherbrooke QC Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Claire E.","family":"Hanson","sequence":"additional","affiliation":[{"name":"School of Biological Sciences Georgia Institute of Technology  Atlanta GA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"S. Mohammadali","family":"Rahmati","sequence":"additional","affiliation":[{"name":"School of Biological Sciences Georgia Institute of Technology  Atlanta GA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ilya A.","family":"Rybak","sequence":"additional","affiliation":[{"name":"Department of Neurobiology and Anatomy Drexel University  Philadelphia PA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9259-2706","authenticated-orcid":false,"given":"Alain","family":"Frigon","sequence":"additional","affiliation":[{"name":"Department of Pharmacology\u2010Physiology Universit\u00e9 de Sherbrooke  Sherbrooke QC Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0499-3890","authenticated-orcid":false,"given":"Boris I.","family":"Prilutsky","sequence":"additional","affiliation":[{"name":"School of Biological Sciences Georgia Institute of Technology  Atlanta GA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2025,5,5]]},"reference":[{"key":"e_1_2_5_2_1","doi-asserted-by":"publisher","DOI":"10.1089\/neu.2022.0042"},{"key":"e_1_2_5_3_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.192054"},{"key":"e_1_2_5_4_1","volume-title":"The coordination and regulation of movements","author":"Bernstein N. A.","year":"1967"},{"key":"e_1_2_5_5_1","volume-title":"Bernstein's construction of movements: the original text and commentaries","author":"Bernstein N. A.","year":"2021"},{"key":"e_1_2_5_6_1","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22094858"},{"key":"e_1_2_5_7_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.brainresrev.2007.08.004"},{"key":"e_1_2_5_8_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4939-3267-2_12"},{"key":"e_1_2_5_9_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.2001.86.1.447"},{"key":"e_1_2_5_10_1","doi-asserted-by":"publisher","DOI":"10.1002\/jmor.10233"},{"key":"e_1_2_5_11_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature25448"},{"key":"e_1_2_5_12_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0147669"},{"key":"e_1_2_5_13_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4904-04.2005"},{"key":"e_1_2_5_14_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00625.2019"},{"key":"e_1_2_5_15_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00817.2003"},{"key":"e_1_2_5_16_1","volume-title":"Text\u2010atlas of cat anatomy","author":"Crouch J. E.","year":"1969"},{"key":"e_1_2_5_17_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0830-06.2006"},{"key":"e_1_2_5_18_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00419.2015"},{"key":"e_1_2_5_19_1","doi-asserted-by":"publisher","DOI":"10.1093\/brain\/awu372"},{"key":"e_1_2_5_20_1","doi-asserted-by":"publisher","DOI":"10.1016\/0166-2236(94)90064-7"},{"key":"e_1_2_5_21_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1265-22.2023"},{"key":"e_1_2_5_22_1","doi-asserted-by":"publisher","DOI":"10.1113\/JP277018"},{"key":"e_1_2_5_23_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00167.2019"},{"key":"e_1_2_5_24_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4343-15.2016"},{"key":"e_1_2_5_25_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0966-6362(97)00042-8"},{"key":"e_1_2_5_26_1","doi-asserted-by":"publisher","DOI":"10.1016\/0006-8993(75)90974-9"},{"key":"e_1_2_5_27_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1748-1716.1980.tb06534.x"},{"key":"e_1_2_5_28_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00676.2012"},{"key":"e_1_2_5_29_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00524.2013"},{"key":"e_1_2_5_30_1","doi-asserted-by":"publisher","DOI":"10.1113\/JP272740"},{"key":"e_1_2_5_31_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.2010.199125"},{"key":"e_1_2_5_32_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1995.sp020727"},{"key":"e_1_2_5_33_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2015.04.004"},{"key":"e_1_2_5_34_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.01300.2004"},{"key":"e_1_2_5_35_1","doi-asserted-by":"publisher","DOI":"10.1002\/cphy.cp010226"},{"key":"e_1_2_5_36_1","doi-asserted-by":"publisher","DOI":"10.3390\/ijms22115882"},{"key":"e_1_2_5_37_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jelekin.2016.07.010"},{"key":"e_1_2_5_38_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnhum.2015.00048"},{"key":"e_1_2_5_39_1","first-page":"1","article-title":"The stride cycle of the cat: The modelling of locomotion by computerized analysis of automatic recordings","volume":"521","author":"Halbertsma J. M.","year":"1983","journal-title":"Acta physiologica Scandinavica Supplementum"},{"key":"e_1_2_5_40_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00104.2024"},{"key":"e_1_2_5_41_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0816-20.2020"},{"key":"e_1_2_5_42_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnsys.2022.814028"},{"key":"e_1_2_5_43_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.50134"},{"key":"e_1_2_5_44_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.clinph.2014.02.001"},{"key":"e_1_2_5_45_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00018.2019"},{"key":"e_1_2_5_46_1","doi-asserted-by":"publisher","DOI":"10.2307\/1311182"},{"key":"e_1_2_5_47_1","first-page":"36","article-title":"When does a gait transition occur during human locomotion?","volume":"6","author":"Hreljac A.","year":"2007","journal-title":"Journal of Science and Medicine"},{"key":"e_1_2_5_48_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1327-05.2005"},{"key":"e_1_2_5_49_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460-9568.2008.06289.x"},{"key":"e_1_2_5_50_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00223.2003"},{"key":"e_1_2_5_51_1","doi-asserted-by":"publisher","DOI":"10.1038\/nrn.2016.9"},{"key":"e_1_2_5_52_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00310.2020"},{"key":"e_1_2_5_53_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00241.2006"},{"key":"e_1_2_5_54_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.2011.215137"},{"key":"e_1_2_5_55_1","doi-asserted-by":"publisher","DOI":"10.1113\/JP284503"},{"key":"e_1_2_5_56_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-022-05293-w"},{"key":"e_1_2_5_57_1","doi-asserted-by":"publisher","DOI":"10.1113\/JP280555"},{"key":"e_1_2_5_58_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnsys.2023.1199079"},{"key":"e_1_2_5_59_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4939-3267-2_2"},{"key":"e_1_2_5_60_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00865.2011"},{"key":"e_1_2_5_61_1","doi-asserted-by":"publisher","DOI":"10.1016\/B978-0-12-819410-2.00017-5"},{"key":"e_1_2_5_62_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.brainresrev.2007.08.006"},{"key":"e_1_2_5_63_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00511.2005"},{"key":"e_1_2_5_64_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-023-39034-y"},{"key":"e_1_2_5_65_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00525.2009"},{"key":"e_1_2_5_66_1","doi-asserted-by":"publisher","DOI":"10.1038\/nm.2160"},{"key":"e_1_2_5_67_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pcbi.1009677"},{"key":"e_1_2_5_68_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.3000410"},{"key":"e_1_2_5_69_1","doi-asserted-by":"publisher","DOI":"10.1016\/0006-8993(80)90207-3"},{"key":"e_1_2_5_70_1","doi-asserted-by":"publisher","DOI":"10.1123\/mcj.4.1.1"},{"key":"e_1_2_5_71_1","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.204.13.2277"},{"key":"e_1_2_5_72_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469-7793.1998.293bu.x"},{"key":"e_1_2_5_73_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.xnsj.2023.100235"},{"key":"e_1_2_5_74_1","unstructured":"Rahmati S. M. A. Klishko A. N. Markin S. N. Shevtsova N. A. Frigon A. Rybak I. A. &Prilutsky B. I.(2023).Hindlimb musculoskeletal structure and mechanical demands of locomotion determine basic organization of spinal locomotor networks in the cat. Program No. 416.03. 2023 Neuroscience Meeting Planner.Washington DC:Society for Neuroscience 2023. Online."},{"key":"e_1_2_5_75_1","first-page":"163861","article-title":"A whole\u2010body musculoskeletal model of the mouse","volume":"9","author":"Tata Ramalingasetty S.","year":"2021","journal-title":"Institute of Electrical and Electronics Engineers Access"},{"key":"e_1_2_5_76_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-neuro-061010-113746"},{"key":"e_1_2_5_77_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jbiomech.2017.09.002"},{"key":"e_1_2_5_78_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.2006.118703"},{"key":"e_1_2_5_79_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.98841.3"},{"key":"e_1_2_5_80_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.2006.118711"},{"key":"e_1_2_5_81_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gaitpost.2017.10.007"},{"key":"e_1_2_5_82_1","doi-asserted-by":"publisher","DOI":"10.1113\/JP277515"},{"key":"e_1_2_5_83_1","first-page":"253","volume-title":"Progress in motor control effects of age, disorder, and rehabilitation","author":"Shapkova E. Y.","year":"2004"},{"key":"e_1_2_5_84_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-1-4939-3267-2_5"},{"key":"e_1_2_5_85_1","first-page":"659","article-title":"Control of walking and running by means of electric stimulation of the midbrain","volume":"11","author":"Shik M. L.","year":"1966","journal-title":"Biophysics"},{"key":"e_1_2_5_86_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-024-07504-y"},{"key":"e_1_2_5_87_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-021-90058-0"},{"key":"e_1_2_5_88_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2022.1041015"},{"key":"e_1_2_5_89_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00236.2014"},{"key":"e_1_2_5_90_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2009.09.002"},{"key":"e_1_2_5_91_1","first-page":"7","article-title":"Minimal spanning tree based fuzzy clustering","volume":"8","author":"Vathy\u2010Fogarassy A.","year":"2005","journal-title":"World Academy of Science, Engineering and Technology"},{"key":"e_1_2_5_92_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1908-15.2015"},{"key":"e_1_2_5_93_1","doi-asserted-by":"publisher","DOI":"10.1038\/srep36275"},{"key":"e_1_2_5_94_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00230447"}],"container-title":["The Journal of Physiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/pdf\/10.1113\/JP288089","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,6,1]],"date-time":"2025-06-01T01:10:19Z","timestamp":1748740219000},"score":1,"resource":{"primary":{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/10.1113\/JP288089"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,5]]},"references-count":93,"journal-issue":{"issue":"10","published-print":{"date-parts":[[2025,5]]}},"alternative-id":["10.1113\/JP288089"],"URL":"https:\/\/doi.org\/10.1113\/jp288089","archive":["Portico"],"relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2024.09.19.613891","asserted-by":"object"}]},"ISSN":["0022-3751","1469-7793"],"issn-type":[{"value":"0022-3751","type":"print"},{"value":"1469-7793","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,5]]},"assertion":[{"value":"2024-11-09","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2025-03-25","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2025-05-05","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}