{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,25]],"date-time":"2026-05-25T09:04:14Z","timestamp":1779699854673,"version":"3.53.1"},"reference-count":90,"publisher":"Oxford University Press (OUP)","issue":"1","license":[{"start":{"date-parts":[[2026,5,6]],"date-time":"2026-05-06T00:00:00Z","timestamp":1778025600000},"content-version":"vor","delay-in-days":125,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,1,14]]},"abstract":"<jats:title>Synopsis<\/jats:title>\n                  <jats:p>The transition from fins to limbs represents a pivotal evolutionary shift that enabled vertebrates to engage terrestrial environments through coordinated transformations in appendicular structure and neuromuscular control. Here, we propose that this transition can be understood as a biomechanical reorganization of force transmission across multi-joint musculoskeletal linkages. Rather than a simple rotational modification transforming fins directly into forelimbs, this process involved progressive reorganization of sarcopterygian appendages, including axial rotation, directional force production, and coordinated stance-swing dynamics that likely emerged prior to fully terrestrial locomotion. This review applies the two-joint link model, originally developed for human limb biomechanics, to extant sarcopterygians such as the coelacanth (Latimeria) and to early-diverging tetrapods, including archosaurs such as the American alligator (Alligator mississippiensis). The model demonstrates how sequential activation of biarticular and monoarticular muscles generates predictable, axis-aligned endpoint force outputs, providing a mechanistic explanation for how ancestral fins and early limbs could support weight bearing, propulsion, and postural stabilization during the water-to-land transition. The framework further clarifies the evolution and persistence of diagonal-couplet lateral sequence gait, a locomotor pattern widespread among tetrapods and consistent with Paleozoic trackway evidence. Integration of electromyographic data with conserved spinal circuitry including central pattern generators, interneurons, and the topographic organization of lateral motor column motor pools reveals how intrinsic spinal architecture governs muscle activation sequences and limb-level force redirection. Although supraspinal pathways modulate locomotion, core coordination of diagonal-couplet gait emerges from spinal mechanisms. A central property of the model is musculoskeletal redundancy: multiple muscle combinations can generate equivalent endpoint forces within linked multi-joint systems, thereby preserving functional capacity across postural variation. This robustness arises from the four-bar linkage organization of biarticular muscles, a design principle conserved across vertebrate musculoskeletal systems. Together, these findings position the two-joint link model as a unifying neuromechanical framework for understanding fin-to-limb evolution, the emergence of tetrapod gait, and the conserved spinal organization underlying vertebrate locomotion.<\/jats:p>","DOI":"10.1093\/iob\/obag020","type":"journal-article","created":{"date-parts":[[2026,5,2]],"date-time":"2026-05-02T12:27:18Z","timestamp":1777724838000},"source":"Crossref","is-referenced-by-count":0,"title":["From Fin to Limb: Orientational Shift and Evolution of Diagonal-Couplet Gait in Tetrapods"],"prefix":"10.1093","volume":"8","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9066-3219","authenticated-orcid":false,"given":"T","family":"Miyake","sequence":"first","affiliation":[{"name":"Ehime University Graduate School of Medicine Department of Medical Education, , To-on, Ehime 791-0295 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"K","family":"Nishikawa","sequence":"additional","affiliation":[{"name":"Graduate School of Human and Environmental Studies, Kyoto University , Yoshida Nihonmatsu-cho, Sakyo 606-8501, Kyoto ,","place":["Japan"]},{"name":"Graduate School of Global Environmental Studies, Kyoto University , Yoshida Hon-machi, Sakyo, Kyoto 606-8501 ,","place":["Japan"]},{"name":"Institute for Environment and Development (LESTARI), Universiti Kebangsaan , Malaysia, UKM Bangi, Selangor 43600 ,","place":["Malaysia"]},{"name":"Faculty of Science, Chulalongkorn University Department of Biology, , Bangkok 10330 ,","place":["Thailand"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"M","family":"Iwata","sequence":"additional","affiliation":[{"name":"Aquamarine Inawashiro, Kingfishers Aquarium , Osada, Inawashiro-machi, Fukushima 969-3283 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"H","family":"Kamiyama","sequence":"additional","affiliation":[{"name":"Atagawa Tropical & Alligator Garden, Naramoto Higashiizu-cho , Shizuoka 413-0302 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"K","family":"Ozaki","sequence":"additional","affiliation":[{"name":"Graduate School of Human and Environmental Studies, Kyoto University , Yoshida Nihonmatsu-cho, Sakyo 606-8501, Kyoto ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"H","family":"Koie","sequence":"additional","affiliation":[{"name":"College of Bioresource Sciences, Nihon University Laboratory of Veterinary Physiology, Department of Veterinary Medicine, , Fujisawa 252-0880 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"A","family":"Yozu","sequence":"additional","affiliation":[{"name":"Nippon Medical School Department of Rehabilitation Medicine, , Sendagi, Tokyo 113-8602 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"T","family":"Hirasawa","sequence":"additional","affiliation":[{"name":"School of Science, The University of Tokyo Department of Earth and Planetary Environmental Science, , Tokyo 113-0033 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"N","family":"Kobayashi","sequence":"additional","affiliation":[{"name":"Ehime University Graduate School of Medicine Department of Medical Education, , To-on, Ehime 791-0295 ,","place":["Japan"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,5,6]]},"reference":[{"key":"2026052504164080500_bib1","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1111\/j.0014-3820.2004.tb01673.x","article-title":"Evolutionary dynamics of complex biomechanical systems: an example using the four-bar mechanism","volume":"58","author":"Alfaro","year":"2004","journal-title":"Evolution"},{"key":"2026052504164080500_bib2","doi-asserted-by":"crossref","first-page":"046015","DOI":"10.1088\/1748-3190\/ad5129","article-title":"Utilising redundancy in musculoskeletal systems for adaptive stiffness and muscle failure compensation: a model-free inverse statics approach","volume":"19","author":"Almanzor","year":"2024","journal-title":"Bioinspir. 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