{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T19:07:13Z","timestamp":1790104033644,"version":"4.0.1"},"reference-count":58,"publisher":"The Company of Biologists","issue":"18","license":[{"start":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T00:00:00Z","timestamp":1789430400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-2\/"}],"funder":[{"name":"Natural Sciences and Engineering Research Council of Canada","award":["RGPIN-2020-05894"],"award-info":[{"award-number":["RGPIN-2020-05894"]}],"id":[{"id":"https:\/\/ror.org\/01h531d29","id-type":"ROR","asserted-by":"crossref"}]},{"name":"Natural Sciences and Engineering Research Council of Canada","award":["585453-2023"],"award-info":[{"award-number":["585453-2023"]}],"id":[{"id":"https:\/\/ror.org\/01h531d29","id-type":"ROR","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,9,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Navigation behaviour of aquatic animals has often been studied in unidirectional flow. Yet many aquatic habitats have highly variable flow directions, particularly those affected by waves. We used field measurements of natural flows and underwater video to survey how the nudibranch Hermissenda crassicornis navigates in wave-affected flow conditions. We compared navigation tracks relative to feeding events in low and high prey densities and low and high flow direction variability. Regardless of flow direction variability or prey density, we found evidence of chemical and flow cue-based navigation towards prey. Evidence of flow-based navigation was only significant when flow direction was less variable and calculated over time scales of wave-affected flow rather than tidal flow. Slugs oriented parallel to flow (either upstream or downstream) before feeding, possibly orienting to flow oscillations from wave action when prey is temporarily upstream (and thus using a form of chemical-gated rheotaxis). In higher flow direction variability, slug track metrics still suggested a navigational strategy was used to find prey, but the apparent absence of any orientation to flow suggested the slugs use a different strategy (possibly chemotaxis) in conditions that could produce highly mixed chemical cues. Finally, when slugs were distant from feeding, they moved cross-stream, which may increase the chances of intercepting prey chemical cues. Collectively, these findings possibly provide further evidence of context-dependent switching of gastropod navigational strategies. In addition, they also suggest that chemical-gated rheotaxis is a viable navigation strategy in habitats with either unidirectional or wave-affected flow.<\/jats:p>","DOI":"10.1242\/jeb.252174","type":"journal-article","created":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T08:56:40Z","timestamp":1788253000000},"source":"Crossref","is-referenced-by-count":1,"title":["Navigational behaviour of the sea slug\n                    <i>Hermissenda crassicornis<\/i>\n                    in natural wave-affected flow conditions"],"prefix":"10.1242","volume":"229","author":[{"ORCID":"https:\/\/orcid.org\/0009-0006-5600-8732","authenticated-orcid":false,"given":"Alexandra M. D.","family":"Reinhardt","sequence":"first","affiliation":[{"name":"St. Francis Xavier University Department of Biology , , Antigonish, NS , , B2G 2W5","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0971-7588","authenticated-orcid":false,"given":"Russell C.","family":"Wyeth","sequence":"additional","affiliation":[{"name":"St. Francis Xavier University Department of Biology , , Antigonish, NS , , B2G 2W5","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2026,9,22]]},"reference":[{"key":"2026092204325395000_JEB252174C1","doi-asserted-by":"publisher","first-page":"jeb250746","DOI":"10.1242\/jeb.250746","article-title":"Navigational behaviour of Lymnaea stagnalis in response to chemical and flow cues","volume":"228","author":"Alansari","year":"2025","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C2","doi-asserted-by":"publisher","first-page":"1","DOI":"10.18637\/jss.v031.i10","article-title":"CircStat: A MATLAB toolbox for circular statistics","volume":"31","author":"Berens","year":"2009","journal-title":"J. Stat. Softw."},{"key":"2026092204325395000_JEB252174C3","doi-asserted-by":"publisher","first-page":"211","DOI":"10.2307\/1942312","article-title":"Interactions between a sea pen and seven of its predators","volume":"44","author":"Birkeland","year":"1974","journal-title":"Ecol. Monogr."},{"key":"2026092204325395000_JEB252174C4","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1002\/ar.b.20090","article-title":"Subcellular, cellular, and circuit mechanisms underlying classical conditioning in Hermissenda crassicornis","volume":"289","author":"Blackwell","year":"2006","journal-title":"Anat. Rec. B New Anat."},{"key":"2026092204325395000_JEB252174C5","doi-asserted-by":"publisher","first-page":"151323","DOI":"10.1016\/j.jembe.2020.151323","article-title":"Smells like home: The use of chemically-mediated rheotaxes by Limulus polyphemus larvae","volume":"525","author":"Butler","year":"2020","journal-title":"J. Exp. Mar. Biol. Ecol."},{"key":"2026092204325395000_JEB252174C6","doi-asserted-by":"publisher","first-page":"e0286384","DOI":"10.1371\/journal.pone.0286384","article-title":"Two decades of change in sea star abundance at a subtidal site in Puget Sound, Washington","volume":"18","author":"Casendino","year":"2023","journal-title":"PLoS ONE"},{"key":"2026092204325395000_JEB252174C7","doi-asserted-by":"publisher","first-page":"113","DOI":"10.1080\/10236240600688748","article-title":"Mechanosensory based orienting behaviors in fluvial and lacustrine populations of mottled sculpin (Cottus bairdi)","volume":"39","author":"Coombs","year":"2006","journal-title":"Mar. Freshw. Behav. Physiol."},{"key":"2026092204325395000_JEB252174C8","doi-asserted-by":"publisher","first-page":"jeb223008","DOI":"10.1242\/jeb.223008","article-title":"Rheotaxis revisited: a multi-behavioral and multisensory perspective on how fish orient to flow","volume":"223","author":"Coombs","year":"2020","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C9","doi-asserted-by":"publisher","first-page":"209","DOI":"10.1523\/JNEUROSCI.05-01-00209.1985","article-title":"Conditioned modification of phototactic behavior in Hermissenda. I. Analysis of light intensity","volume":"5","author":"Crow","year":"1985","journal-title":"J. Neurosci."},{"key":"2026092204325395000_JEB252174C10","doi-asserted-by":"publisher","first-page":"412","DOI":"10.1126\/science.209.4454.412","article-title":"Associative behavioral modification in Hermissenda: cellular correlates","volume":"209","author":"Crow","year":"1980","journal-title":"Science"},{"key":"2026092204325395000_JEB252174C11","doi-asserted-by":"publisher","first-page":"57","DOI":"10.1080\/10236249909379037","article-title":"Coordination of chemosensory orientation in the starfish Asterias forbesi","volume":"32","author":"Dale","year":"1999","journal-title":"Mar. Freshw. Behav. Physiol."},{"key":"2026092204325395000_JEB252174C12","doi-asserted-by":"publisher","first-page":"4083","DOI":"10.1242\/jeb.008664","article-title":"The flow generated by an active olfactory system of the red swamp crayfish","volume":"210","author":"Denissenko","year":"2007","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C13","doi-asserted-by":"publisher","first-page":"2511","DOI":"10.1007\/BF01014727","article-title":"Optimal search direction for an animal flying or swimming in a wind or current","volume":"15","author":"Dusenbery","year":"1989","journal-title":"J. Chem. Ecol."},{"key":"2026092204325395000_JEB252174C14","doi-asserted-by":"publisher","first-page":"1971","DOI":"10.1007\/BF01020509","article-title":"Upwind searching for an odor plume is sometimes optimal","volume":"16","author":"Dusenbery","year":"1990","journal-title":"J. Chem. Ecol."},{"key":"2026092204325395000_JEB252174C15","doi-asserted-by":"publisher","first-page":"809","DOI":"10.1242\/jeb.01438","article-title":"Slow-moving predatory gastropods track prey odors in fast and turbulent flow","volume":"208","author":"Ferner","year":"2005","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C16","doi-asserted-by":"publisher","first-page":"1056","DOI":"10.4319\/lo.1999.44.4.1056","article-title":"Odor transport in turbulent flows: Constraints on animal navigation","volume":"44","author":"Finelli","year":"1999","journal-title":"Limnol. Oceanogr."},{"key":"2026092204325395000_JEB252174C17","doi-asserted-by":"publisher","first-page":"3878","DOI":"10.1242\/jeb.167056","article-title":"Bringing the analysis of animal orientation data full circle: Model-based approaches with maximum likelihood","volume":"220","author":"Fitak","year":"2017","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C18","doi-asserted-by":"publisher","first-page":"1325","DOI":"10.1111\/2041-210X.12584","article-title":"BORIS: A free, versatile open-source event-logging software for video\/audio coding and live observations","volume":"7","author":"Friard","year":"2016","journal-title":"Method. Ecol. Evol."},{"key":"2026092204325395000_JEB252174C19","volume-title":"Oregon Estuarine Invertebrates: Illustrated Guide to Common Species","author":"Hiebert","year":"2017"},{"key":"2026092204325395000_JEB252174C20","doi-asserted-by":"publisher","first-page":"199","DOI":"10.1007\/s10750-012-1044-x","article-title":"Nudibranch predation and dietary preference for the polyps of Aurelia labiata (Cnidaria: Scyphozoa)","volume":"690","author":"Hoover","year":"2012","journal-title":"Hydrobiologia"},{"key":"2026092204325395000_JEB252174C21","doi-asserted-by":"publisher","first-page":"1557","DOI":"10.1007\/s00265-012-1409-1","article-title":"Sea lamprey orient toward a source of a synthesized pheromone using odor-conditioned rheotaxis","volume":"66","author":"Johnson","year":"2012","journal-title":"Behav. Ecol. Sociobiol."},{"key":"2026092204325395000_JEB252174C22","doi-asserted-by":"publisher","first-page":"514","DOI":"10.1039\/C6NP00121A","article-title":"Finding food: How marine invertebrates use chemical cues to track and select food","volume":"34","author":"Kamio","year":"2017","journal-title":"Nat. Prod. Rep."},{"key":"2026092204325395000_JEB252174C23","doi-asserted-by":"publisher","first-page":"274","DOI":"10.2307\/1543362","article-title":"An initial study on the effects of signal intermittency on the odor plume tracking behavior of the American lobster, Homarus americanus","volume":"201","author":"Kozlowski","year":"2001","journal-title":"Biol. Bull."},{"key":"2026092204325395000_JEB252174C24","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1080\/1023624031000136279","article-title":"Changes in odour intermittency influence the success and search behaviour during orientation in the crayfish (Orconectes rusticus)","volume":"36","author":"Kozlowski","year":"2003","journal-title":"Mar. Freshw. Behav. Physiol."},{"key":"2026092204325395000_JEB252174C25","doi-asserted-by":"publisher","first-page":"753","DOI":"10.1163\/1937240X-00002378","article-title":"Behaviorally-selective chemoreceptor lesions reveal two different chemically mediated orientation strategies in the rusty crayfish, Orconectes rusticus","volume":"35","author":"Kraus-Epley","year":"2015","journal-title":"J. Crustac. Biol."},{"key":"2026092204325395000_JEB252174C26","doi-asserted-by":"publisher","first-page":"30","DOI":"10.1186\/s12898-019-0246-8","article-title":"The Hermans\u2013Rasson test as a powerful alternative to the Rayleigh test for circular statistics in biology","volume":"19","author":"Landler","year":"2019","journal-title":"BMC Ecol."},{"key":"2026092204325395000_JEB252174C27","doi-asserted-by":"publisher","first-page":"bio049866","DOI":"10.1242\/bio.049866","article-title":"Model selection versus traditional hypothesis testing in circular statistics: a simulation study","volume":"9","author":"Landler","year":"2020","journal-title":"Biol. Open"},{"key":"2026092204325395000_JEB252174C28","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1016\/S0163-1047(80)91599-X","article-title":"Complex responses to light of the nudibranch Hermissenda crassicornis (Gastropoda: Opisthobranchia)","volume":"28","author":"Lederhendler","year":"1980","journal-title":"Behav. Neural Biol."},{"key":"2026092204325395000_JEB252174C29","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1186\/s10152-017-0489-8","article-title":"Orientation and food search behaviour of a deep sea lobster in turbulent versus laminar odour plumes","volume":"71","author":"Major","year":"2017","journal-title":"Helgol. Mar. Res."},{"key":"2026092204325395000_JEB252174C30","doi-asserted-by":"publisher","first-page":"603","DOI":"10.2307\/1935526","article-title":"Feeding behavior of asteroids and escape responses of their prey in the Puget Sound region","volume":"49","author":"Mauzey","year":"1968","journal-title":"Ecology"},{"key":"2026092204325395000_JEB252174C31","doi-asserted-by":"publisher","first-page":"4149","DOI":"10.1242\/jeb.111153","article-title":"One rhinophore probably provides sufficient sensory input for odour-based navigation by the nudibranch mollusc Tritonia diomedea","volume":"217","author":"McCullagh","year":"2014","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C32","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1242\/jeb.00063","article-title":"Fine-scale patterns of odor encounter by the antennules of mantis shrimp tracking turbulent plumes in wave-affected and unidirectional flow","volume":"206","author":"Mead","year":"2003","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C33","doi-asserted-by":"publisher","first-page":"183","DOI":"10.1016\/B978-0-12-391857-4.00009-4","volume-title":"Imaging and Spectroscopic Analysis of Living Cells","author":"Meijering","year":"2012"},{"key":"2026092204325395000_JEB252174C34","volume-title":"Intertidal Invertebrates of California","author":"Morris","year":"1980"},{"key":"2026092204325395000_JEB252174C35","doi-asserted-by":"publisher","first-page":"487","DOI":"10.1093\/plankt\/fbh040","article-title":"Host location by larvae of a parasitic barnacle: Larval chemotaxis and plume tracking in flow","volume":"26","author":"Pasternak","year":"2004","journal-title":"J. Plankton Res."},{"key":"2026092204325395000_JEB252174C36","doi-asserted-by":"publisher","first-page":"jeb245213","DOI":"10.1242\/jeb.245213","article-title":"State-dependent, visually guided behaviors in the nudibranch Berghia stephanieae","volume":"226","author":"Quinlan","year":"2023","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C37","unstructured":"Reinhardt, A. M.\n           (2026). Navigational behaviour of Hermissenda crassicornis in different natural cue environments. MSc thesis, St. Francis Xavier University. https:\/\/hdl.handle.net\/20.500.14648\/65761"},{"key":"2026092204325395000_JEB252174C38","doi-asserted-by":"publisher","first-page":"3","DOI":"10.2307\/1541369","article-title":"Navigational strategies and models","volume":"23","author":"Rodrigo","year":"2002","journal-title":"Psicol\u00f3gica"},{"key":"2026092204325395000_JEB252174C39","doi-asserted-by":"publisher","first-page":"676","DOI":"10.1038\/nmeth.2019","article-title":"Fiji: An open-source platform for biological-image analysis","volume":"9","author":"Schindelin","year":"2012","journal-title":"Nat. Methods"},{"key":"2026092204325395000_JEB252174C40","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1016\/S0003-3472(05)80068-5","article-title":"Statistical analysis of animal orientation data","volume":"43","author":"Schnute","year":"1992","journal-title":"Anim. Behav."},{"key":"2026092204325395000_JEB252174C41","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1080\/10236244.2015.1123870","article-title":"Detection of substrate-associated odour cues versus prey-associated cues by the oral veil in Tritonia diomedea","volume":"49","author":"Schwarz","year":"2016","journal-title":"Mar. Freshw. Behav. Physiol."},{"key":"2026092204325395000_JEB252174C42","doi-asserted-by":"publisher","first-page":"225","DOI":"10.1007\/BF00379881","article-title":"Variable wind directions and anemotactic strategies of searching for an odour plume","volume":"63","author":"Sabelis","year":"1984","journal-title":"Oecologia"},{"key":"2026092204325395000_JEB252174C43","doi-asserted-by":"publisher","first-page":"447","DOI":"10.1093\/icb\/icv073","article-title":"Additional navigational strategies can augment odor-gated rheotaxis for navigation under conditions of variable flow","volume":"55","author":"Vasey","year":"2015","journal-title":"Integr. Comp. Biol."},{"key":"2026092204325395000_JEB252174C44","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1111\/fme.70026","article-title":"Fisheries management and ecology assessment of a baited remote underwater video method to evaluate American lobster (Homarus americanus) response to baits in nature","volume":"2025","author":"Walls","year":"2025","journal-title":"Fish. Manag. Ecol."},{"key":"2026092204325395000_JEB252174C45","doi-asserted-by":"publisher","first-page":"1034","DOI":"10.4319\/lo.2001.46.5.1034","article-title":"Chemosensory guidance cues in a turbulent chemical odor plume","volume":"46","author":"Webster","year":"2001","journal-title":"Limnol. Oceanogr."},{"key":"2026092204325395000_JEB252174C46","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1146\/annurev.fluid.010908.165240","article-title":"The hydrodynamics of chemical cues among aquatic Organisms","volume":"41","author":"Webster","year":"2009","journal-title":"Annu. Rev. Fluid Mech."},{"key":"2026092204325395000_JEB252174C47","doi-asserted-by":"publisher","first-page":"188","DOI":"10.2307\/1542523","article-title":"The fluid dynamical context of chemosensory behavior","volume":"198","author":"Weissburg","year":"2000","journal-title":"Biol. Bull."},{"key":"2026092204325395000_JEB252174C48","doi-asserted-by":"publisher","first-page":"1428","DOI":"10.2307\/1940072","article-title":"Life and death in moving fluids: hydrodynamic effects on chemosensory-mediated predation","volume":"74","author":"Weissburg","year":"1993","journal-title":"Ecology"},{"key":"2026092204325395000_JEB252174C49","doi-asserted-by":"publisher","first-page":"349","DOI":"10.1242\/jeb.197.1.349","article-title":"Odor plumes and how blue crabs use them in finding prey","volume":"197","author":"Weissburg","year":"1994","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C50","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-319-24277-4","volume-title":"ggplot2: Elegant Graphics for Data Analysis","author":"Wickham","year":"2016"},{"key":"2026092204325395000_JEB252174C51","doi-asserted-by":"publisher","first-page":"1323","DOI":"10.1016\/j.anbehav.2012.08.024","article-title":"Temporal and spatial sampling strategies maintain tracking success of whelks to prey patches of differing distributions","volume":"84","author":"Wilson","year":"2012","journal-title":"Anim. Behav."},{"key":"2026092204325395000_JEB252174C52","doi-asserted-by":"publisher","first-page":"143","DOI":"10.1145\/1978942.1978963","article-title":"The aligned rank transform for nonparametric factorial analyses using only anova procedures","author":"Wobbrock","year":"2011"},{"key":"2026092204325395000_JEB252174C53","doi-asserted-by":"publisher","first-page":"42","DOI":"10.3389\/fnbeh.2010.00042","article-title":"Should animals navigating over short distances switch to a magnetic compass sense?","volume":"4","author":"Wyeth","year":"2010","journal-title":"Front. Behav. Neurosci."},{"key":"2026092204325395000_JEB252174C54","doi-asserted-by":"publisher","first-page":"jeb185843","DOI":"10.1242\/jeb.185843","article-title":"Olfactory navigation in aquatic gastropods","volume":"222","author":"Wyeth","year":"2019","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C56","doi-asserted-by":"publisher","first-page":"1441","DOI":"10.1242\/jeb.02164","article-title":"Odours detected by rhinophores mediate orientation to flow in the nudibranch mollusc, Tritonia diomedea","volume":"209","author":"Wyeth","year":"2006","journal-title":"J. Exp. Biol."},{"key":"2026092204325395000_JEB252174C57","doi-asserted-by":"publisher","first-page":"97","DOI":"10.2307\/4134599","article-title":"Orientation and navigation relative to water flow, prey, conspecifics, and predators by the nudibranch mollusc Tritonia diomedea","volume":"210","author":"Wyeth","year":"2006","journal-title":"Biol. Bull."},{"key":"2026092204325395000_JEB252174C58","doi-asserted-by":"publisher","first-page":"105","DOI":"10.1086\/713663","article-title":"Rare-earth magnets influence movement patterns of the magnetically sensitive nudibranch Tritonia exsulans in its natural habitat","volume":"240","author":"Wyeth","year":"2021","journal-title":"Biol. Bull."},{"key":"2026092204325395000_JEB252174C59","doi-asserted-by":"publisher","first-page":"111","DOI":"10.2307\/1542075","article-title":"Odor plumes and animal navigation in turbulent water flow: a field study","volume":"188","author":"Zimmer-Faust","year":"1995","journal-title":"Biol. Bull."}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.252174\/3715071\/jeb252174.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.252174\/3718902\/jeb252174.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.252174\/3718902\/jeb252174.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,22]],"date-time":"2026-09-22T08:33:05Z","timestamp":1790065985000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/229\/18\/jeb252174\/372986\/Navigational-behaviour-of-the-sea-slug-Hermissenda"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,15]]},"references-count":58,"journal-issue":{"issue":"18","published-print":{"date-parts":[[2026,9,15]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.252174","relation":{},"ISSN":["0022-0949","1477-9145"],"issn-type":[{"value":"0022-0949","type":"print"},{"value":"1477-9145","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,15]]},"article-number":"jeb252174"}}