{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T22:29:09Z","timestamp":1784240949480,"version":"3.55.0"},"reference-count":49,"publisher":"The Company of Biologists","issue":"15","license":[{"start":{"date-parts":[[2021,2,9]],"date-time":"2021-02-09T00:00:00Z","timestamp":1612828800000},"content-version":"vor","delay-in-days":558,"URL":"http:\/\/www.biologists.com\/user-licence-1-1\/"}],"funder":[{"DOI":"10.13039\/100007247","name":"Adolph C. and Mary Sprague Miller Institute for Basic Research in Science, University of California Berkeley","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100007247","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100000861","name":"Burroughs Wellcome Fund","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000861","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000183","name":"Army Research Office","doi-asserted-by":"publisher","award":["W911NF-17-1-0346"],"award-info":[{"award-number":["W911NF-17-1-0346"]}],"id":[{"id":"10.13039\/100000183","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100012957","name":"Friedrich-Schiller-Universit\u00e4t Jena","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100012957","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100006754","name":"Army Research Laboratory","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100006754","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2019,8,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title><jats:p>Terrestrial animals often must self-right from an upside-down orientation on the ground to survive. Here, we compared self-righting strategies of the Madagascar hissing, American and discoid cockroaches on a challenging flat, rigid, low-friction surface to quantify the mechanical principles. All three species almost always self-righted (97% probability) when given time (30\u2005s), frequently self-righted (63%) on the first attempt, and on that attempt did so in 1\u00a0s or less. When successful, two of the three species gained and used pitch and\/or roll rotational kinetic energy to overcome potential energy barriers (American 63% of all attempts and discoid 78%). By contrast, the largest, heaviest, wingless cockroach (Madagascar hissing) relied far less on the energy of motion and was the slowest to self-right. Two of the three species used rolling strategies to overcome low potential energy barriers. Successful righting attempts had greater rolling rotation than failed attempts as the center of mass rose to the highest position. Madagascar hissing cockroaches rolled using body deformation (98% of all trials) and the American cockroach rolled using leg forces (93%). By contrast, the discoid cockroach overcame higher and a wider range of potential energy barriers with simultaneous pitching and rolling using the wings (46% of all trials) and legs (49%) equally to self-right. Our quantification revealed the performance advantages of using rotational kinetic energy to overcome the potential energy barrier and rolling more to lower it, while maintaining diverse strategies for ground-based self-righting.<\/jats:p>","DOI":"10.1242\/jeb.186080","type":"journal-article","created":{"date-parts":[[2019,8,9]],"date-time":"2019-08-09T13:44:00Z","timestamp":1565358240000},"source":"Crossref","is-referenced-by-count":31,"title":["Cockroaches use diverse strategies to self-right on the ground"],"prefix":"10.1242","volume":"222","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7516-3646","authenticated-orcid":false,"given":"Chen","family":"Li","sequence":"first","affiliation":[{"name":"Department\u2008of Mechanical Engineering, Johns Hopkins University, Baltimore, MD 21218, USA"},{"name":"Department of Integrative Biology, University of California, Berkeley, Berkeley, CA 94720, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1373-8488","authenticated-orcid":false,"given":"Toni","family":"W\u00f6hrl","sequence":"additional","affiliation":[{"name":"Institute of Sports Science, Friedrich-Schiller-Universit\u00e4t Jena, 07743 Jena, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1302-3880","authenticated-orcid":false,"given":"Han K.","family":"Lam","sequence":"additional","affiliation":[{"name":"Department of Integrative Biology, University of California, Berkeley, Berkeley, CA 94720, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Robert J.","family":"Full","sequence":"additional","affiliation":[{"name":"Department of Integrative Biology, University of California, Berkeley, Berkeley, CA 94720, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2019,8,9]]},"reference":[{"key":"2021042623071461500_JEB186080C1","doi-asserted-by":"publisher","first-page":"150","DOI":"10.3758\/BF03335647","article-title":"The righting reflex in turtles: a description and comparison","volume":"20","author":"Ashe","year":"1970","journal-title":"Psychon. Sci."},{"key":"2021042623071461500_JEB186080C2","doi-asserted-by":"crossref","DOI":"10.1353\/book.3295","volume-title":"Cockroaches: Ecology, Behavior, and Natural History","author":"Bell","year":"2007"},{"key":"2021042623071461500_JEB186080C4","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1111\/j.1469-7998.1990.tb04033.x","article-title":"A novel method of self-righting in the springtail Sminthurus viridis (Insecta: Collembola)","volume":"222","author":"Brackenbury","year":"1990","journal-title":"J. Zool."},{"key":"2021042623071461500_JEB186080C5","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1007\/BF00620403","article-title":"Behavioral switching in cockroaches: transformations of tactile reflexes during righting behavior","volume":"113","author":"Camhi","year":"1977","journal-title":"J. Comp. Physiol. A"},{"key":"2021042623071461500_JEB186080C6","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1038\/s41598-017-15787-7","article-title":"Self-righting potential and the evolution of shell shape in Gal\u00e1pagos tortoises","volume":"7","author":"Chiari","year":"2017","journal-title":"Sci. Rep."},{"key":"2021042623071461500_JEB186080C7","doi-asserted-by":"publisher","first-page":"1369","DOI":"10.1242\/jeb.051136","article-title":"Slipping, sliding and stability: locomotor strategies for overcoming low-friction surfaces","volume":"214","author":"Clark","year":"2011","journal-title":"J. Exp. Biol."},{"key":"2021042623071461500_JEB186080C8","doi-asserted-by":"publisher","first-page":"15681","DOI":"10.1073\/pnas.0601473103","article-title":"Running over rough terrain reveals limb control for intrinsic stability","volume":"103","author":"Daley","year":"2006","journal-title":"Proc. Natl. Acad. Sci."},{"key":"2021042623071461500_JEB186080C9","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1242\/jeb.133.1.111","article-title":"Motor activity during searching and walking movements of cockroach legs","volume":"133","author":"Delcomyn","year":"1987","journal-title":"J. Exp. Biol."},{"key":"2021042623071461500_JEB186080C10","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1098\/rspb.2007.1188","article-title":"Geometry and self-righting of turtles","volume":"275","author":"Domokos","year":"2008","journal-title":"Proc. R. Soc. London B Biol. Sci."},{"key":"2021042623071461500_JEB186080C11","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1111\/j.1469-7998.1973.tb04553.x","article-title":"The jump of the click beetle (Coleoptera: E1ateridae)-energetics and mechanics","volume":"169","author":"Evans","year":"1973","journal-title":"J. Zool."},{"key":"2021042623071461500_JEB186080C12","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1242\/jeb.204.4.637","article-title":"Coordinated righting behaviour in locusts","volume":"204","author":"Faisal","year":"2001","journal-title":"J. Exp. Biol."},{"key":"2021042623071461500_JEB186080C13","doi-asserted-by":"publisher","first-page":"481","DOI":"10.1111\/j.1469-185X.2007.00020.x","article-title":"Leave it all behind: A taxonomic perspective of autotomy in invertebrates","volume":"82","author":"Fleming","year":"2007","journal-title":"Biol. Rev."},{"key":"2021042623071461500_JEB186080C14","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1016\/j.asd.2004.05.007","article-title":"Righting kinematics in beetles (Insecta: Coleoptera)","volume":"33","author":"Frantsevich","year":"2004","journal-title":"Arthropod Struct. Dev."},{"key":"2021042623071461500_JEB186080C15","doi-asserted-by":"publisher","first-page":"279","DOI":"10.1007\/BF00657348","article-title":"Turning and righting in Geotrupes (Coleoptera, Scarabaeidae)","volume":"289","author":"Frantsevich","year":"1980","journal-title":"J. Comp. Physiol."},{"key":"2021042623071461500_JEB186080C16","doi-asserted-by":"crossref","first-page":"2441","DOI":"10.1242\/jeb.198.12.2441","article-title":"Maximum single leg force production: cockroaches righting on photoelastic gelatin","volume":"198","author":"Full","year":"1995","journal-title":"J. Exp. Biol."},{"key":"2021042623071461500_JEB186080C17","doi-asserted-by":"publisher","first-page":"69","DOI":"10.1111\/jzo.12047","article-title":"Variations in righting behaviour across Hermann's tortoise populations","volume":"291","author":"Golubovic","year":"2013","journal-title":"J. Zool."},{"key":"2021042623071461500_JEB186080C18","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1007\/BF00337390","article-title":"Effects of circum-oesophageal lesion on the behaviour of the stick insect Carausius morosus","volume":"145","author":"Graham","year":"1979","journal-title":"Biol. Cybern."},{"key":"2021042623071461500_JEB186080C19","doi-asserted-by":"publisher","first-page":"034001","DOI":"10.1088\/1748-3182\/3\/3\/034001","article-title":"Software techniques for two- and three-dimensional kinematic measurements of biological and biomimetic systems","volume-title":"Bioinspir. Biomim.","author":"Hedrick","year":"2008"},{"key":"2021042623071461500_JEB186080C20","first-page":"E64","article-title":"Running without feet: the role of tarsi during high-speed horizontal locomotion in cockroaches","volume":"51","author":"Jayaram","year":"2011","journal-title":"Integr. Comp. Biol."},{"key":"2021042623071461500_JEB186080C21","doi-asserted-by":"publisher","first-page":"4215","DOI":"10.1073\/pnas.0711944105","article-title":"Active tails enhance arboreal acrobatics in geckos","volume":"105","author":"Jusufi","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"2021042623071461500_JEB186080C22","doi-asserted-by":"publisher","first-page":"530","DOI":"10.1046\/j.1365-2435.1999.00343.x","article-title":"The size-grain hypothesis and interspecific scaling in ants","volume":"13","author":"Kaspari","year":"1999","journal-title":"Funct. Ecol."},{"key":"2021042623071461500_JEB186080C23","doi-asserted-by":"publisher","first-page":"251","DOI":"10.1016\/j.asd.2004.06.003","article-title":"Mechanical aspects of legged locomotion control","volume":"33","author":"Koditschek","year":"2004","journal-title":"Arthropod Struct. Dev."},{"key":"2021042623071461500_JEB186080C24","doi-asserted-by":"crossref","first-page":"1919","DOI":"10.1242\/jeb.200.13.1919","article-title":"Three-dimensional kinematics and limb kinetic energy of running cockroaches","volume":"200","author":"Kram","year":"1997","journal-title":"J. Exp. Biol."},{"key":"2021042623071461500_JEB186080C25","doi-asserted-by":"publisher","first-page":"3293","DOI":"10.1242\/jeb.061937","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":"2021042623071461500_JEB186080C26","doi-asserted-by":"publisher","first-page":"046003","DOI":"10.1088\/1748-3190\/10\/4\/046003","article-title":"Terradynamically streamlined shapes in animals and robots enhance traversability through densely cluttered terrain","volume":"10","author":"Li","year":"2015","journal-title":"Bioinspir. Biomim."},{"key":"2021042623071461500_JEB186080C27","first-page":"2128","article-title":"Cockroach-inspired winged robot reveals principles of ground-based dynamic self-righting","author":"Li","year":"2016"},{"key":"2021042623071461500_JEB186080C28","doi-asserted-by":"publisher","first-page":"881","DOI":"10.1080\/01691864.2017.1372213","article-title":"Mechanical principles of dynamic terrestrial self-righting using wings","volume":"31","author":"Li","year":"2017","journal-title":"Adv. Robot."},{"key":"2021042623071461500_JEB186080C29","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1038\/nature10710","article-title":"Tail-assisted pitch control in lizards, robots and dinosaurs","volume":"481","author":"Libby","year":"2012","journal-title":"Nature"},{"key":"2021042623071461500_JEB186080C30","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1016\/j.physbeh.2016.01.001","article-title":"Asymmetry of righting reflexes in sea turtles and its behavioral correlates","volume":"157","author":"Malashichev","year":"2016","journal-title":"Physiol. Behav."},{"key":"2021042623071461500_JEB186080C31","doi-asserted-by":"publisher","first-page":"373","DOI":"10.1111\/j.1469-7998.2006.00079.x","article-title":"Shaping up to fight: sexual selection influences body shape and size in the fighting tortoise (Chersina angulata)","volume":"269","author":"Mann","year":"2006","journal-title":"J. Zool."},{"key":"2021042623071461500_JEB186080C32","volume-title":"Handbook of Biological Statistics","author":"McDonald","year":"2009"},{"key":"2021042623071461500_JEB186080C33","doi-asserted-by":"publisher","first-page":"1039","DOI":"10.1152\/japplphysiol.01177.2001","article-title":"Energy cost of walking and running at extreme uphill and downhill slopes","volume":"93","author":"Minetti","year":"2002","journal-title":"J. Appl. Physiol."},{"key":"2021042623071461500_JEB186080C34","first-page":"E370","article-title":"Leg vibrations help cockroaches self-right using wings","volume":"57","author":"Othayoth","year":"2017","journal-title":"Integr. Comp. Biol."},{"key":"2021042623071461500_JEB186080C35","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1017\/S0263574715001009","article-title":"Motion planning and implementation for the self-recovery of an overturned multi-legged robot","volume":"5","author":"Peng","year":"2015","journal-title":"Robotica"},{"key":"2021042623071461500_JEB186080C36","doi-asserted-by":"publisher","first-page":"1407","DOI":"10.1016\/0022-1910(77)90166-4","article-title":"A quantitative analysis of rhythmic leg movements during three different behaviors in the cockroach, Periplaneta americana","volume":"23","author":"Reingold","year":"1977","journal-title":"J. Insect Physiol."},{"key":"2021042623071461500_JEB186080C37","doi-asserted-by":"publisher","first-page":"jeb.182642","DOI":"10.1242\/jeb.182642","article-title":"Biomechanical factors influencing successful self-righting in the pleurodire turtle, Emydura subglobosa","author":"Rubin","year":"2018","journal-title":"J. Exp. Biol."},{"key":"2021042623071461500_JEB186080C38","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1080\/10407413.2016.1163983","article-title":"The reciprocity of environment and action in self-righting beetles: the textures of the ground and an object, and the claws","volume":"28","author":"Sasaki","year":"2016","journal-title":"Ecol. Psychol."},{"key":"2021042623071461500_JEB186080C39","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1007\/BF00620404","article-title":"A modified walking rhythm employed during righting behavior in the cockroach Gromphadorhina portentosa","volume":"316","author":"Sherman","year":"1977","journal-title":"J. Comp. Physiol."},{"key":"2021042623071461500_JEB186080C40","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":"2021042623071461500_JEB186080C41","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":"2021042623071461500_JEB186080C42","doi-asserted-by":"publisher","first-page":"315","DOI":"10.1016\/j.bbr.2006.06.023","article-title":"Lateralized righting behavior in the tortoise (Testudo hermanni)","volume":"173","author":"Stancher","year":"2006","journal-title":"Behav. Brain Res."},{"key":"2021042623071461500_JEB186080C43","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1242\/jeb.197.1.251","article-title":"Dynamic and static stability in hexapedal runners","volume":"197","author":"Ting","year":"1994","journal-title":"J. Exp. Biol."},{"key":"2021042623071461500_JEB186080C44","first-page":"276","article-title":"Observations on the swimming, righting, and burrowing movements of young horseshoe crabs, limulus polyphemus","volume":"70","author":"Vosatka","year":"1970","journal-title":"Ohio J. Sci."},{"key":"2021042623071461500_JEB186080C45","doi-asserted-by":"crossref","first-page":"2135","DOI":"10.1242\/jeb.205.14.2135","article-title":"Scaling of rotational inertia in murine rodents and two species of lizard","volume":"205","author":"Walter","year":"2002","journal-title":"J. Exp. Biol."},{"key":"2021042623071461500_JEB186080C46","doi-asserted-by":"publisher","first-page":"353","DOI":"10.1017\/S0952836903003820","article-title":"Sexual dimorphism of body size and shell shape in European tortoises","volume":"260","author":"Willemsen","year":"2003","journal-title":"J. Zool."},{"key":"2021042623071461500_JEB186080C47","first-page":"E255","article-title":"In silico experiments reveal the importance of randomness of motions in cockroach's winged self-righting","volume":"59","author":"Xuan","year":"2019","journal-title":"Integr. Comp. Biol."},{"key":"2021042623071461500_JEB186080C48","doi-asserted-by":"publisher","first-page":"978","DOI":"10.1007\/s00300-006-0140-7","article-title":"The effects of temperature on walking and righting in temperate and Antarctic crustaceans","volume":"29","author":"Young","year":"2006","journal-title":"Polar Biol."},{"key":"2021042623071461500_JEB186080C49","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1002\/neu.480170406","article-title":"A model of pattern generation of cockroach walking reconsidered","volume":"17","author":"Zill","year":"1986","journal-title":"J. Neurobiol."},{"key":"2021042623071461500_JEB186080C50","doi-asserted-by":"publisher","first-page":"20133072","DOI":"10.1098\/rspb.2013.3072","article-title":"Static antennae act as locomotory guides that compensate for visual motion blur in a diurnal, keen-eyed predator","volume":"281","author":"Zurek","year":"2014","journal-title":"Proc. R. Soc. B"}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1242\/jeb.186080","content-type":"unspecified","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.186080\/1975863\/jeb186080.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.186080\/1975863\/jeb186080.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,9,18]],"date-time":"2023-09-18T23:48:34Z","timestamp":1695080914000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/222\/15\/jeb186080\/223383\/Cockroaches-use-diverse-strategies-to-self-right"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,8,1]]},"references-count":49,"journal-issue":{"issue":"15","published-print":{"date-parts":[[2019,8,1]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.186080","relation":{},"ISSN":["1477-9145","0022-0949"],"issn-type":[{"value":"1477-9145","type":"electronic"},{"value":"0022-0949","type":"print"}],"subject":[],"published":{"date-parts":[[2019,8,1]]},"article-number":"jeb186080"}}