{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,15]],"date-time":"2026-07-15T15:17:34Z","timestamp":1784128654782,"version":"3.55.0"},"reference-count":61,"publisher":"The Company of Biologists","issue":"4","license":[{"start":{"date-parts":[[2023,2,15]],"date-time":"2023-02-15T00:00:00Z","timestamp":1676419200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-1\/"}],"funder":[{"name":"National Science Foundation","award":["GR10003305"],"award-info":[{"award-number":["GR10003305"]}]},{"name":"National Science Foundation","award":["DMS1764406"],"award-info":[{"award-number":["DMS1764406"]}]},{"name":"National Science Foundation"},{"DOI":"10.13039\/100000893","name":"Simons Foundation","doi-asserted-by":"crossref","award":["SFARI 594594"],"award-info":[{"award-number":["SFARI 594594"]}],"id":[{"id":"10.13039\/100000893","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2023,2,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n               <jats:p>Centipedes coordinate body and limb flexion to generate propulsion. On flat, solid surfaces, the limb-stepping patterns can be characterized according to the direction in which limb-aggregates propagate, opposite to (retrograde) or with the direction of motion (direct). It is unknown how limb and body dynamics are modified in terrain with terradynamic complexity more representative of these animal's natural heterogeneous environments. Here, we investigated how centipedes that use retrograde and direct limb-stepping patterns, Scolopendra polymorpha and Scolopocryptops sexspinosus, respectively, coordinate their body and limbs to navigate laboratory environments which present footstep challenges and terrain rugosity. We recorded the kinematics and measured the locomotive performance of these animals traversing two rugose terrains with randomly distributed step heights and compared the kinematics with those on a flat frictional surface. Scolopendra polymorpha exhibited similar body and limb dynamics across all terrains and a decrease in speed with increased terrain rugosity. Unexpectedly, when placed in a rugose terrain, S. sexspinosus changed the direction of the limb-stepping pattern from direct to retrograde. Further, for both species, traversal of these rugose terrains was facilitated by hypothesized passive mechanics: upon horizontal collision of a limb with a block, the limb bent and later continued the stepping pattern. Although centipedes have many degrees of freedom, our results suggest these animals negotiate limb\u2013substrate interactions and navigate complex terrains leveraging the innate flexibility of their limbs to simplify control.<\/jats:p>","DOI":"10.1242\/jeb.244688","type":"journal-article","created":{"date-parts":[[2023,1,19]],"date-time":"2023-01-19T12:16:55Z","timestamp":1674130615000},"source":"Crossref","is-referenced-by-count":9,"title":["Active and passive mechanics for rugose terrain traversal in centipedes"],"prefix":"10.1242","volume":"226","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1988-9431","authenticated-orcid":false,"given":"Kelimar","family":"Diaz","sequence":"first","affiliation":[{"name":"Georgia Institute of Technology 1 Interdisciplinary Graduate Program in Quantitative Biosciences , , Atlanta, GA 30332 , USA"},{"name":"School of Physics, Georgia Institute of Technology 2 , Atlanta, GA 30332 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eva","family":"Erickson","sequence":"additional","affiliation":[{"name":"School of Physics, Georgia Institute of Technology 2 , Atlanta, GA 30332 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Baxi","family":"Chong","sequence":"additional","affiliation":[{"name":"Georgia Institute of Technology 1 Interdisciplinary Graduate Program in Quantitative Biosciences , , Atlanta, GA 30332 , USA"},{"name":"School of Physics, Georgia Institute of Technology 2 , Atlanta, GA 30332 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Daniel","family":"Soto","sequence":"additional","affiliation":[{"name":"School of Mechanical Engineering, Georgia Institute of Technology 3 , Atlanta, GA 30332 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6954-9857","authenticated-orcid":false,"given":"Daniel I.","family":"Goldman","sequence":"additional","affiliation":[{"name":"School of Physics, Georgia Institute of Technology 2 , Atlanta, GA 30332 , USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2023,2,23]]},"reference":[{"key":"2024021308280623000_JEB244688C1","volume-title":"Principles of Animal Locomotion","author":"Alexander","year":"2003"},{"key":"2024021308280623000_JEB244688C2","doi-asserted-by":"publisher","first-page":"1185","DOI":"10.1242\/jeb.198.5.1185","article-title":"Axial kinematics and muscle activity during terrestrial locomotion of the centipede Scolopendra heros","volume":"198","author":"Anderson","year":"1995","journal-title":"J. Exp. Biol."},{"key":"2024021308280623000_JEB244688C3","doi-asserted-by":"publisher","first-page":"012717","DOI":"10.1103\/PhysRevE.87.012717","article-title":"Instability-based mechanism for body undulations in centipede locomotion","volume":"87","author":"Aoi","year":"2013","journal-title":"Phys. Rev. E Stat. Nonlin. Soft. Matter. Phys."},{"key":"2024021308280623000_JEB244688C4","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-016-0001-8","article-title":"Advantage of straight walk instability in turning maneuver of multilegged locomotion: a robotics approach","volume":"6","author":"Aoi","year":"2016","journal-title":"Sci. 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Biomim."},{"key":"2024021308280623000_JEB244688C35","first-page":"3293","article-title":"Multi-functional foot use during running in the zebra-tailed lizard (Callisaurus draconoides)","volume":"215","author":"Li","year":"2012","journal-title":"J. Exp. Biol."},{"key":"2024021308280623000_JEB244688C36","doi-asserted-by":"crossref","DOI":"10.1109\/IEMBS.1995.579743","article-title":"Control implications of musculoskeletal mechanics","author":"Loeb","year":"1995"},{"key":"2024021308280623000_JEB244688C37","doi-asserted-by":"publisher","first-page":"603","DOI":"10.1126\/science.1231806","article-title":"Controlled flight of a biologically inspired, insect-scale robot","volume":"340","author":"Ma","year":"2013","journal-title":"Science"},{"key":"2024021308280623000_JEB244688C38","doi-asserted-by":"publisher","first-page":"314","DOI":"10.1126\/science.1172490","article-title":"Undulatory swimming in sand: subsurface locomotion of the sandfish lizard","volume":"325","author":"Maladen","year":"2009","journal-title":"Science"},{"key":"2024021308280623000_JEB244688C39","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1111\/j.1096-3642.1952.tb01855.x","article-title":"The evolution of arthopodan locomotory mechanisms - Part 3. The locomotion of Chilopoda and Pauropoda","volume":"42","author":"Manton","year":"1952","journal-title":"Zool. J. Linn. Soc."},{"key":"2024021308280623000_JEB244688C40","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1111\/j.1096-3642.1965.tb00500.x","article-title":"The evolution of arthropodan locomotory mechanisms. Part 8. Functional requirements and body design in Chilopoda, together with a comparative account of their skeleto-muscular systems and an appendix on a comparison between burrowing forces of annelids and chilopods and its bearing upon the evolution of the arthropodan haemocoel","volume":"45","author":"Manton","year":"1965","journal-title":"Zool. J. Linn. Soc."},{"key":"2024021308280623000_JEB244688C41","author":"Manton","year":"1977"},{"key":"2024021308280623000_JEB244688C42","doi-asserted-by":"crossref","DOI":"10.1109\/SSRR.2014.7017667","article-title":"Semi-autonomous centipede-like robot with flexible legs","author":"Masuda","year":"2014"},{"key":"2024021308280623000_JEB244688C43","doi-asserted-by":"publisher","first-page":"1281","DOI":"10.1038\/s41593-018-0209-y","article-title":"Deeplabcut: markerless pose estimation of user-defined body parts with deep learning","volume":"21","author":"Mathis","year":"2018","journal-title":"Nat. Neurosci."},{"key":"2024021308280623000_JEB244688C44","doi-asserted-by":"publisher","first-page":"4530","DOI":"10.1242\/jeb.083477","article-title":"Locomotion-and mechanics-mediated tactile sensing: antenna reconfiguration simplifies control during high-speed navigation in cockroaches","volume":"216","author":"Mongeau","year":"2013","journal-title":"J. Exp. Biol."},{"key":"2024021308280623000_JEB244688C45","doi-asserted-by":"crossref","DOI":"10.1109\/RoboSoft48309.2020.9116022","article-title":"A systematic approach to creating terrain-capable hybrid soft\/hard myriapod robots","author":"Ozkan-Aydin","year":"2020"},{"key":"2024021308280623000_JEB244688C46","doi-asserted-by":"publisher","first-page":"1088","DOI":"10.1126\/science.1145803","article-title":"Self-organization, embodiment, and biologically inspired robotics","volume":"318","author":"Pfeifer","year":"2007","journal-title":"Science"},{"key":"2024021308280623000_JEB244688C47","doi-asserted-by":"publisher","first-page":"629","DOI":"10.1016\/j.conb.2008.01.001","article-title":"Adaptive motor behavior in insects","volume":"17","author":"Ritzmann","year":"2007","journal-title":"Curr. Opin. Neurobiol."},{"key":"2024021308280623000_JEB244688C48","doi-asserted-by":"publisher","first-page":"483","DOI":"10.1002\/jez.1401190307","article-title":"Tarsal structure and climbing ability of cockroaches","volume":"119","author":"Roth","year":"1952","journal-title":"J. Exp. Zool."},{"key":"2024021308280623000_JEB244688C49","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1007\/978-3-642-70094-1_9","article-title":"Behavior theory and invertebrate learning","volume-title":"The Biology of Learning","author":"Sahley","year":"1984"},{"key":"2024021308280623000_JEB244688C50","doi-asserted-by":"publisher","first-page":"4191","DOI":"10.1242\/jeb.00663","article-title":"The aerodynamics of insect flight","volume":"206","author":"Sane","year":"2003","journal-title":"J. Exp. Biol."},{"key":"2024021308280623000_JEB244688C51","doi-asserted-by":"publisher","first-page":"4798","DOI":"10.1073\/pnas.1808675116","article-title":"Mechanical diffraction reveals the role of passive dynamics in a slithering snake","volume":"116","author":"Schiebel","year":"2019","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"2024021308280623000_JEB244688C52","doi-asserted-by":"crossref","first-page":"300","DOI":"10.1007\/978-3-030-64313-3_29","article-title":"Robophysical modeling of bilaterally activated and soft limbless locomotors","volume-title":"Conference on Biomimetic and Biohybrid Systems","author":"Schiebel","year":"2020"},{"key":"2024021308280623000_JEB244688C53","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1109\/48.757275","article-title":"Review of fish swimming modes for aquatic locomotion","volume":"24","author":"Sfakiotakis","year":"1999","journal-title":"IEEE J. Ocean. Eng."},{"key":"2024021308280623000_JEB244688C54","doi-asserted-by":"publisher","first-page":"440","DOI":"10.1242\/jeb.121939","article-title":"Locomotor benefits of being a slender and slick sand swimmer","volume":"218","author":"Sharpe","year":"2015","journal-title":"J. Exp. Biol."},{"key":"2024021308280623000_JEB244688C55","unstructured":"Soto, D.\n           (2022). Simplifying robotic locomotion by escaping traps via an active tail. PhD thesis, Georgia Institute of Technology."},{"key":"2024021308280623000_JEB244688C56","article-title":"Enhancing legged robot navigation of rough terrain via tail tapping","author":"Soto","year":"2021"},{"key":"2024021308280623000_JEB244688C57","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1088\/1748-3182\/2\/1\/002","article-title":"Distributed mechanical feedback in arthropods and robots simplifies control of rapid running on challenging terrain","volume":"2","author":"Spagna","year":"2007","journal-title":"Bioinspir. Biomim."},{"key":"2024021308280623000_JEB244688C58","doi-asserted-by":"publisher","first-page":"433","DOI":"10.1242\/jeb.012385","article-title":"Neuromechanical response of musculo-skeletal structures in cockroaches during rapid running on rough terrain","volume":"211","author":"Sponberg","year":"2008","journal-title":"J. Exp. Biol."},{"key":"2024021308280623000_JEB244688C59","volume-title":"Crawling and Burrowing Mechanics and Energetics of Animal Locomotion","author":"Trueman","year":"1977"},{"key":"2024021308280623000_JEB244688C60","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-019-53258-3","article-title":"Decoding the essential interplay between central and peripheral control in adaptive locomotion of amphibious centipedes","volume":"9","author":"Yasui","year":"2019","journal-title":"Sci. Rep."},{"key":"2024021308280623000_JEB244688C61","article-title":"Decentralized control mechanism underlying interlimb coordination of centipedes","author":"Yasui","year":"2017"},{"key":"2024021308280623000_JEB244688C62","doi-asserted-by":"publisher","first-page":"e0171421","DOI":"10.1371\/journal.pone.0171421","article-title":"Decentralized control scheme for myriapod robot inspired by adaptive and resilient centipede locomotion","volume":"12","author":"Yasui","year":"2017","journal-title":"PloS One"}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.244688\/2449619\/jeb244688.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.244688\/3367263\/jeb244688.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.244688\/3367263\/jeb244688.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,2,13]],"date-time":"2024-02-13T08:28:20Z","timestamp":1707812900000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/226\/4\/jeb244688\/287689\/Active-and-passive-mechanics-for-rugose-terrain"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2023,2,15]]},"references-count":61,"journal-issue":{"issue":"4","published-print":{"date-parts":[[2023,2,15]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.244688","relation":{},"ISSN":["0022-0949","1477-9145"],"issn-type":[{"value":"0022-0949","type":"print"},{"value":"1477-9145","type":"electronic"}],"subject":[],"published":{"date-parts":[[2023,2,15]]},"article-number":"jeb244688"}}