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However when the training demands exceed the ability to recover, it may lead to a state of persistent maladaptation. A prolonged imbalance between training or competition and recovery can cause extended periods of performance impairment, known as functional or non\u2010functional overreaching, which may progress to overtraining syndrome (OTS). This condition can significantly impair athletic development and, in severe cases, cause athletes to retire earlier than expected. Skeletal muscle is key for physical performance, including the ability to move and utilise energy. Impaired physical performance is a hallmark of OTS, reflecting the central role of skeletal muscle dysfunction in its development and progression. In this review we discuss the potential mechanisms contributing to skeletal muscle impairments in OTS. These include central and peripheral neural fatigue, endocrine dysregulation and altered hormonal responses, impaired mitochondrial bioenergetics and oxidative stress and systemic inflammation. We also draw parallels between maladaptive response in overreaching and OTS and those typically observed in chronic inflammatory and catabolic conditions, as these may offer insights into underlying mechanistic understanding of OTS, as well as future treatment strategies. Because replicating the full clinical picture of OTS in humans in experimental settings is constrained by practical challenges, we further discuss animal models that may allow for controlled investigations of external variables and thus enable enhanced understanding of its complex pathophysiology.\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\/tjp70416-gra-0001-m.png\">\n                        <jats:alt-text>image<\/jats:alt-text>\n                      <\/jats:graphic>\n                    <\/jats:boxed-text>\n                  <\/jats:p>","DOI":"10.1113\/jp287708","type":"journal-article","created":{"date-parts":[[2026,3,8]],"date-time":"2026-03-08T21:27:07Z","timestamp":1773005227000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":6,"title":["Intramuscular pathways of maladaptation in overtraining syndrome"],"prefix":"10.1113","author":[{"given":"Emily","family":"Shorter","sequence":"first","affiliation":[{"name":"Molecular Muscle Physiology &amp; Pathophysiology Group, Department of Physiology and Pharmacology Karolinska Institutet  Stockholm Sweden"},{"name":"Department of Physiology, Nutrition and Biomechanics the Swedish School of Sport and Health Sciences  Stockholm Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Oscar","family":"Horwath","sequence":"additional","affiliation":[{"name":"Department of Physiology, Nutrition and Biomechanics the Swedish School of Sport and Health Sciences  Stockholm Sweden"},{"name":"Division of Pediatric Neurology, Department of Women's and Children's Health Karolinska Institutet  Stockholm Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sabrina Tzivia","family":"Barsky","sequence":"additional","affiliation":[{"name":"Faculty of Kinesiology and Physical Education University of Toronto  Toronto Ontario Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ferdinand","family":"von Walden","sequence":"additional","affiliation":[{"name":"Division of Pediatric Neurology, Department of Women's and Children's Health Karolinska Institutet  Stockholm Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Johanna T.","family":"Lanner","sequence":"additional","affiliation":[{"name":"Molecular Muscle Physiology &amp; Pathophysiology Group, Department of Physiology and Pharmacology Karolinska Institutet  Stockholm Sweden"},{"name":"Department of Physiology, Nutrition and Biomechanics the Swedish School of Sport and Health Sciences  Stockholm Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,2,26]]},"reference":[{"key":"e_1_2_2_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-024-49460-1"},{"key":"e_1_2_2_3_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jmb.2008.08.087"},{"key":"e_1_2_2_4_1","doi-asserted-by":"publisher","DOI":"10.15190\/d.2023.18"},{"key":"e_1_2_2_5_1","doi-asserted-by":"publisher","DOI":"10.1519\/JSC.0000000000000421"},{"key":"e_1_2_2_6_1","doi-asserted-by":"publisher","DOI":"10.1084\/jem.20070075"},{"key":"e_1_2_2_7_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.2011.212845"},{"key":"e_1_2_2_8_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jnutbio.2009.01.005"},{"key":"e_1_2_2_9_1","doi-asserted-by":"publisher","DOI":"10.1210\/jcem-60-4-803"},{"key":"e_1_2_2_10_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00421-002-0737-7"},{"key":"e_1_2_2_11_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.physbeh.2021.113413"},{"key":"e_1_2_2_12_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.physbeh.2023.114344"},{"key":"e_1_2_2_13_1","first-page":"609","article-title":"Central and peripheral fatigue in sustained maximum voluntary contractions of human quadriceps muscle","volume":"54","author":"Bigland Ritchie B.","year":"1978","journal-title":"Clinical Science and Molecular Medicine"},{"key":"e_1_2_2_14_1","doi-asserted-by":"publisher","DOI":"10.1002\/mus.880070902"},{"key":"e_1_2_2_15_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1990.sp018008"},{"key":"e_1_2_2_16_1","first-page":"789","article-title":"Mitochondrial dysfunction results from oxidative stress in the skeletal muscle of diet\u2010induced insulin\u2010resistant mice","volume":"118","author":"Bonnard C.","year":"2008","journal-title":"Journal of Clinical Investigation"},{"key":"e_1_2_2_17_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0185993"},{"issue":"12","key":"e_1_2_2_18_1","article-title":"p38 MAPKs \u2010 roles in skeletal muscle physiology, disease mechanisms, and as potential therapeutic targets","volume":"6","author":"Brennan C. 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