{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,12]],"date-time":"2026-10-12T16:28:38Z","timestamp":1791822518287,"version":"4.3.4"},"reference-count":47,"publisher":"Public Library of Science (PLoS)","issue":"4","license":[{"start":{"date-parts":[[2021,4,8]],"date-time":"2021-04-08T00:00:00Z","timestamp":1617840000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Saxon State government"}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"abstract":"<jats:p>The original windlass mechanism describes a one-to-one coupling between metatarsal joint dorsiflexion and medial longitudinal arch rise. The description assumes a sufficiently stiff plantar aponeurosis and absence of foot muscle activity. However, recent research calls for a broader interpretation of the windlass mechanism that accounts for an extensible plantar aponeurosis and active foot muscles. In this study, we investigate the rise of the arch in response to toe dorsiflexion when sitting, standing, and walking to discuss the windlass mechanism\u2019s contributions in static and dynamic load scenarios. 3D motion analysis allowed a kinematic investigation of the rise and drop of the arch relative to the extent of toe dorsiflexion. The results suggest that static windlass effects poorly predict the relationship between arch dynamics and metatarsophalangeal joint motion during dynamic load scenarios, such as walking. We were able to show that toe dorsiflexion resulted in an immediate rise of the longitudinal arch during sitting and standing. In contrast, a decrease in arch height was observed during walking, despite toe dorsiflexion at the beginning of the push-off phase. Further, the longitudinal arch rose almost linearly with toe dorsiflexion in the static loading scenarios, while the dynamic load scenario revealed an exponential rise of the arch. In addition to that, the rate of change in arch height relative to toe motion was significantly lower when sitting and standing compared to walking. Finally, and most surprisingly, arch rise was found to correlate with toe dorsiflexion only in the dynamic loading scenario. These results challenge the traditional perspective of the windlass mechanism as the dominating source of foot rigidity for push-off against the ground during bipedal walking. It seems plausible that other mechanisms besides the windlass act to raise the foot arch.<\/jats:p>","DOI":"10.1371\/journal.pone.0249965","type":"journal-article","created":{"date-parts":[[2021,4,8]],"date-time":"2021-04-08T18:05:39Z","timestamp":1617905139000},"page":"e0249965","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":35,"title":["The rise of the longitudinal arch when sitting, standing, and walking: Contributions of the windlass mechanism"],"prefix":"10.1371","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7032-5025","authenticated-orcid":true,"given":"Freddy","family":"Sichting","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9162-5676","authenticated-orcid":true,"given":"Florian","family":"Ebrecht","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2021,4,8]]},"reference":[{"issue":"5","key":"pone.0249965.ref001","doi-asserted-by":"crossref","first-page":"1513","DOI":"10.1152\/jn.00256.2011","article-title":"Foot anatomy specialization for postural sensation and control","volume":"107","author":"WG Wright","year":"2012","journal-title":"J Neurophysiol"},{"issue":"Pt 5","key":"pone.0249965.ref002","doi-asserted-by":"crossref","first-page":"713","DOI":"10.1242\/jeb.019885","article-title":"Walking, running and the evolution of short toes in humans","volume":"212","author":"C Rolian","year":"2009","journal-title":"J Exp Biol"},{"issue":"Pt 17","key":"pone.0249965.ref003","article-title":"Rethinking the evolution of the human foot: insights from experimental research","volume":"221","author":"NB Holowka","year":"2018","journal-title":"J Exp Biol"},{"key":"pone.0249965.ref004","article-title":"Evolutionary anatomy of the plantar aponeurosis in primates, including humans","author":"F Sichting","year":"2020","journal-title":"J Anat"},{"issue":"1","key":"pone.0249965.ref005","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1002\/ajpa.1330200109","article-title":"Chimpanzee and human feet in bipedal walking","volume":"20","author":"H Elftman","year":"1935","journal-title":"Am. J. Phys. Anthropol"},{"issue":"1","key":"pone.0249965.ref006","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1002\/ajpa.1330830103","article-title":"Metatarsophalangeal joints of Australopithecus afarensis","volume":"83","author":"B Latimer","year":"1990","journal-title":"Am. J. Phys. Anthropol"},{"issue":"1","key":"pone.0249965.ref007","first-page":"25","article-title":"The mechanics of the foot. II. The plantar aponeurosis and the arch","volume":"88","author":"JH Hicks","year":"1954","journal-title":"J Anat"},{"issue":"1","key":"pone.0249965.ref008","doi-asserted-by":"crossref","first-page":"18","DOI":"10.1177\/107110070002100104","article-title":"The biomechanical relationship between the tendoachilles, plantar fascia and metatarsophalangeal joint dorsiflexion angle","volume":"21","author":"RE Carlson","year":"2000","journal-title":"Foot Ankle Int"},{"issue":"8","key":"pone.0249965.ref009","doi-asserted-by":"crossref","first-page":"845","DOI":"10.3113\/FAI.2008.0845","article-title":"Nonlinear finite element analysis of the plantar fascia due to the windlass mechanism","volume":"29","author":"H-YK Cheng","year":"2008","journal-title":"Foot Ankle Int"},{"issue":"9","key":"pone.0249965.ref010","doi-asserted-by":"crossref","first-page":"1937","DOI":"10.1016\/j.jbiomech.2008.03.028","article-title":"Finite element analysis of plantar fascia under stretch-the relative contribution of windlass mechanism and Achilles tendon force","volume":"41","author":"H-YK Cheng","year":"2008","journal-title":"J Biomech"},{"issue":"4","key":"pone.0249965.ref011","doi-asserted-by":"crossref","first-page":"313","DOI":"10.7547\/0960313","article-title":"The relationship between navicular drop and first metatarsophalangeal joint motion","volume":"96","author":"JS Paton","year":"2006","journal-title":"J Am Podiatr Med Assoc"},{"issue":"5","key":"pone.0249965.ref012","doi-asserted-by":"crossref","first-page":"245","DOI":"10.7547\/87507315-91-5-245","article-title":"Function of the windlass mechanism in excessively pronated feet","volume":"91","author":"A Aquino","year":"2001","journal-title":"J Am Podiatr Med Assoc"},{"issue":"6","key":"pone.0249965.ref013","doi-asserted-by":"crossref","first-page":"598","DOI":"10.1111\/joa.12050","article-title":"An investigation of the dynamic relationship between navicular drop and first metatarsophalangeal joint dorsal excursion","volume":"222","author":"NL Griffin","year":"2013","journal-title":"J Anat"},{"issue":"1943","key":"pone.0249965.ref014","first-page":"20202095","article-title":"The extensibility of the plantar fascia influences the windlass mechanism during human running","volume":"288","author":"L Welte","year":"2021","journal-title":"Proc Biol Sci"},{"issue":"3","key":"pone.0249965.ref015","doi-asserted-by":"crossref","first-page":"254","DOI":"10.1111\/j.1469-7580.2010.01261.x","article-title":"Dynamics of longitudinal arch support in relation to walking speed: contribution of the plantar aponeurosis","volume":"217","author":"P Caravaggi","year":"2010","journal-title":"J Anat"},{"issue":"168","key":"pone.0249965.ref016","doi-asserted-by":"crossref","first-page":"20200208","DOI":"10.1098\/rsif.2020.0208","article-title":"Foot stiffening during the push-off phase of human walking is linked to active muscle contraction, and not the windlass mechanism","volume":"17","author":"DJ Farris","year":"2020","journal-title":"J R Soc Interface"},{"issue":"5","key":"pone.0249965.ref017","doi-asserted-by":"crossref","first-page":"1645","DOI":"10.1073\/pnas.1812820116","article-title":"The functional importance of human foot muscles for bipedal locomotion","volume":"116","author":"DJ Farris","year":"2019","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"5","key":"pone.0249965.ref018","doi-asserted-by":"crossref","first-page":"290","DOI":"10.1136\/bjsports-2013-092690","article-title":"The foot core system: a new paradigm for understanding intrinsic foot muscle function","volume":"49","author":"PO McKeon","year":"2015","journal-title":"Br J Sports Med"},{"key":"pone.0249965.ref019","first-page":"1551","volume-title":"Gray\u2019s anatomy: The anatomical basis of clinical practice","author":"H Gray","year":"2008","edition":"4"},{"issue":"4","key":"pone.0249965.ref020","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1007\/s00421-014-3056-x","article-title":"Coordination of intrinsic and extrinsic foot muscles during walking","volume":"115","author":"KE Zelik","year":"2015","journal-title":"Eur J Appl Physiol"},{"key":"pone.0249965.ref021","doi-asserted-by":"crossref","first-page":"469","DOI":"10.2106\/00004623-196446030-00001","article-title":"Phasic activity of intrinsic muscles of the foot","volume":"46","author":"R Mann","year":"1964","journal-title":"J Bone Joint Surg Am"},{"issue":"1","key":"pone.0249965.ref022","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1002\/ar.1091610101","article-title":"Electromyography and cinematography of leg and foot (\"normal\" and flat) during walking","volume":"161","author":"EG Gray","year":"1968","journal-title":"Anat Rec"},{"issue":"102","key":"pone.0249965.ref023","doi-asserted-by":"crossref","first-page":"20141076","DOI":"10.1098\/rsif.2014.1076","article-title":"Active regulation of longitudinal arch compression and recoil during walking and running","volume":"12","author":"LA Kelly","year":"2015","journal-title":"J R Soc Interface"},{"issue":"119","key":"pone.0249965.ref024","doi-asserted-by":"crossref","DOI":"10.1098\/rsif.2016.0174","article-title":"Shoes alter the spring-like function of the human foot during running","volume":"13","author":"LA Kelly","year":"2016","journal-title":"J R Soc Interface"},{"issue":"1","key":"pone.0249965.ref025","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1249\/MSS.0000000000001420","article-title":"The Influence of Foot-Strike Technique on the Neuromechanical Function of the Foot","volume":"50","author":"LA Kelly","year":"2018","journal-title":"Med Sci Sports Exerc"},{"issue":"6","key":"pone.0249965.ref026","doi-asserted-by":"crossref","first-page":"471","DOI":"10.1016\/j.aanat.2014.07.003","article-title":"Changes in length of the plantar aponeurosis during the stance phase of gait\u2014an in vivo dynamic fluoroscopic study","volume":"196","author":"G Fessel","year":"2014","journal-title":"Ann Anat"},{"issue":"6","key":"pone.0249965.ref027","first-page":"662","article-title":"An electromyographic study of certain muscles of the leg and foot in the standing position","volume":"98","author":"JV Basmajian","year":"1954","journal-title":"Surg Gynecol Obstet"},{"issue":"6","key":"pone.0249965.ref028","doi-asserted-by":"crossref","first-page":"1184","DOI":"10.2106\/00004623-196345060-00006","article-title":"The role of muscles in arch support of the foot: an electromyographic study","volume":"45","author":"JV Basmajian","year":"1963","journal-title":"J Bone Joint Surg Am"},{"issue":"1","key":"pone.0249965.ref029","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1016\/j.clinbiomech.2011.07.013","article-title":"Recruitment of the plantar intrinsic foot muscles with increasing postural demand.","volume":"27","author":"LA Kelly","year":"2012","journal-title":"Clin Biomech (Bristol, Avon)"},{"issue":"93","key":"pone.0249965.ref030","doi-asserted-by":"crossref","first-page":"20131188","DOI":"10.1098\/rsif.2013.1188","article-title":"Intrinsic foot muscles have the capacity to control deformation of the longitudinal arch","volume":"11","author":"LA Kelly","year":"2014","journal-title":"J R Soc Interface"},{"issue":"1","key":"pone.0249965.ref031","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1159\/000141055","article-title":"The foot as a support","volume":"25","author":"JH Hicks","year":"1955","journal-title":"Acta Anat (Basel)"},{"issue":"1","key":"pone.0249965.ref032","doi-asserted-by":"crossref","first-page":"135","DOI":"10.1111\/j.1469-7998.1983.tb04266.x","article-title":"A dynamic similarity hypothesis for the gaits of quadrupedal mammals","volume":"201","author":"RM Alexander","year":"1983","journal-title":"Journal of Zoology"},{"key":"pone.0249965.ref033","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1186\/s13047-018-0257-2","article-title":"A minimal markerset for three-dimensional foot function assessment: measuring navicular drop and drift under dynamic conditions","volume":"11","author":"P Eichelberger","year":"2018","journal-title":"J Foot Ankle Res"},{"issue":"1","key":"pone.0249965.ref034","doi-asserted-by":"crossref","first-page":"e84347","DOI":"10.1371\/journal.pone.0084347","article-title":"Anatomy and biomechanical properties of the plantar aponeurosis: a cadaveric study","volume":"9","author":"D Chen","year":"2014","journal-title":"PLoS ONE"},{"issue":"3","key":"pone.0249965.ref035","doi-asserted-by":"crossref","first-page":"190","DOI":"10.1016\/S0268-0033(97)00038-7","article-title":"The windlass mechanism during normal walking and passive first metatarsalphalangeal joint extension","volume":"13","author":"A Kappel-Bargas","year":"1998","journal-title":"Clinical Biomechanics"},{"key":"pone.0249965.ref036","unstructured":"Yawar A, Korpas L, Lugo-Bolanos M, Mandre S, Venkadesan M. Contribution of the transverse arch to foot stiffness in humans. arXiv preprint arXiv:1706.04610 2017."},{"issue":"3","key":"pone.0249965.ref037","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1177\/107110070302400307","article-title":"The in vivo elastic properties of the plantar fascia during the contact phase of walking","volume":"24","author":"A Gefen","year":"2003","journal-title":"Foot Ankle Int"},{"key":"pone.0249965.ref038","doi-asserted-by":"crossref","first-page":"e7487","DOI":"10.7717\/peerj.7487","article-title":"Ankle and midtarsal joint quasi-stiffness during walking with added mass","volume":"7","author":"AM Kern","year":"2019","journal-title":"PeerJ"},{"issue":"1","key":"pone.0249965.ref039","doi-asserted-by":"crossref","first-page":"15404","DOI":"10.1038\/s41598-017-15218-7","article-title":"Energy neutral: the human foot and ankle subsections combine to produce near zero net mechanical work during walking","volume":"7","author":"KZ Takahashi","year":"2017","journal-title":"Sci Rep"},{"issue":"7000","key":"pone.0249965.ref040","doi-asserted-by":"crossref","first-page":"147","DOI":"10.1038\/325147a0","article-title":"The spring in the arch of the human foot","volume":"325","author":"RF Ker","year":"1987","journal-title":"Nature"},{"issue":"Pt 15","key":"pone.0249965.ref041","doi-asserted-by":"crossref","first-page":"2491","DOI":"10.1242\/jeb.025767","article-title":"A dynamic model of the windlass mechanism of the foot: evidence for early stance phase preloading of the plantar aponeurosis","volume":"212","author":"P Caravaggi","year":"2009","journal-title":"J Exp Biol"},{"issue":"3","key":"pone.0249965.ref042","doi-asserted-by":"crossref","first-page":"546","DOI":"10.2106\/00004623-200403000-00013","article-title":"Dynamic loading of the plantar aponeurosis in walking","volume":"86","author":"A Erdemir","year":"2004","journal-title":"J Bone Joint Surg Am"},{"issue":"4","key":"pone.0249965.ref043","doi-asserted-by":"crossref","first-page":"e0152602","DOI":"10.1371\/journal.pone.0152602","article-title":"The Role of Arch Compression and Metatarsophalangeal Joint Dynamics in Modulating Plantar Fascia Strain in Running","volume":"11","author":"KA McDonald","year":"2016","journal-title":"PLoS ONE"},{"issue":"5","key":"pone.0249965.ref044","doi-asserted-by":"crossref","first-page":"704","DOI":"10.1016\/j.jbiomech.2016.02.023","article-title":"Elastic energy within the human plantar aponeurosis contributes to arch shortening during the push-off phase of running","volume":"49","author":"JC Wager","year":"2016","journal-title":"J Biomech"},{"issue":"Pt 3","key":"pone.0249965.ref045","first-page":"599","article-title":"Plantar aponeurosis and internal architecture of the ball of the foot","volume":"121","author":"F Bojsen-Moller","year":"1976","journal-title":"J Anat"},{"issue":"5","key":"pone.0249965.ref046","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1177\/107110070102200513","article-title":"The effect of heel elevation on strain within the plantar aponeurosis: in vitro study","volume":"22","author":"GF Kogler","year":"2001","journal-title":"Foot Ankle Int"},{"issue":"10","key":"pone.0249965.ref047","doi-asserted-by":"crossref","first-page":"905","DOI":"10.1007\/s00276-011-0873-z","article-title":"Investigation of the mechanical properties of the plantar aponeurosis","volume":"33","author":"PG Pavan","year":"2011","journal-title":"Surg Radiol Anat"}],"container-title":["PLOS ONE"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0249965","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,4,8]],"date-time":"2021-04-08T18:06:13Z","timestamp":1617905173000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0249965"}},"subtitle":[],"editor":[{"given":"Yury","family":"Ivanenko","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"editor"}]}],"short-title":[],"issued":{"date-parts":[[2021,4,8]]},"references-count":47,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2021,4,8]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0249965","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,4,8]]}}}