{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T18:58:52Z","timestamp":1785437932902,"version":"3.56.0"},"posted":{"date-parts":[[2025,7,3]]},"group-title":"Animal Behavior and Cognition","reference-count":62,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2025,7,3]],"date-time":"2025-07-03T00:00:00Z","timestamp":1751500800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"award":["R15 AI183095"],"award-info":[{"award-number":["R15 AI183095"]}],"id":[{"id":"https:\/\/ror.org\/043z4tv69","id-type":"ROR","asserted-by":"publisher"}]},{"award":["HHSN272201700014I"],"award-info":[{"award-number":["HHSN272201700014I"]}],"id":[{"id":"https:\/\/ror.org\/043z4tv69","id-type":"ROR","asserted-by":"publisher"}]},{"name":"National Science Foundation","award":["CNS-1920220"],"award-info":[{"award-number":["CNS-1920220"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2025,7,3]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>Over 700 million people are at risk of contracting schistosomiasis due to regular exposure to freshwater sources where infected snails, the obligate intermediate hosts of schistosomes, are endemic. Although mass drug administration of praziquantel effectively controls the disease in most regions, achieving elimination will require reducing populations of infected snails that shed the human-infective larval stage. Considerable effort has focused on parasite development and immunological responses after snail penetration, but comparatively little is known about the molecular and behavioral host seeking events that precede it, primarily due to technical and physical constraints. To address this gap, we developed a custom imaging and computational system for tracking and screening schistosome miracidia, the snail-infective larval form that hatches from eggs. Our system employs an array of cameras without magnification and acrylic devices that maintain miracidia within the focal plane, create a field of view over 200,000 times the area of a single miracidium, and support the formation of stable chemical gradients. Using this platform, we perform quantitative ethology of miracidia at an unprecedented scale and extract features that drive the emergent chemoklinokinetic behavior in response to snail cues. We demonstrate that miracidia accumulate at the edge of a gradient of snail cues by increasing key chemoklinokinetic features upon leaving the region of a cue, corroborating previous reports. In contrast, miracidia do not exhibit these behaviors when the cue is uniform, demonstrating that they represent a specific sensory response rather than generic neuromuscular activity. We further find that a previously identified stimulatory snail peptide only partially recapitulates the full chemoklinokinetic profile, and homologues from closely related species elicit divergent behavioral outcomes. Notably, some of these snail peptides can mask a natural gradient and inhibit miracidia penetration of snails. This work establishes a scalable behavioral platform for probing parasite- snail interactions and identifies a peptide scaffold that potently blocks snail penetration.<\/jats:p>","DOI":"10.1101\/2025.07.02.662618","type":"posted-content","created":{"date-parts":[[2025,7,3]],"date-time":"2025-07-03T20:15:29Z","timestamp":1751573729000},"source":"Crossref","is-referenced-by-count":1,"title":["Quantitative ethology of schistosome miracidia characterizes a conserved snail peptide that inhibits penetration"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0009-0003-2547-7112","authenticated-orcid":false,"given":"Rachel V.","family":"Horejsi","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-0282-554X","authenticated-orcid":false,"given":"Chase N.","family":"Nelson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-9938-8559","authenticated-orcid":false,"given":"Avery","family":"De Ruyter","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0005-4630-600X","authenticated-orcid":false,"given":"Helen","family":"Gensch","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-2720-5702","authenticated-orcid":false,"given":"Saige","family":"Maasz-Seawright","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-8935-3350","authenticated-orcid":false,"given":"Carly","family":"Weber","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-6979-1334","authenticated-orcid":false,"given":"Sophie","family":"Willett","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-3666-1348","authenticated-orcid":false,"given":"Sonja A.","family":"Olson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5909-4190","authenticated-orcid":false,"given":"Nicolas J.","family":"Wheeler","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2025070713354548000_2025.07.02.662618v1.1","doi-asserted-by":"crossref","unstructured":"Grimes JE , Croll D , Harrison WE , Utzinger J , Freeman MC , Templeton MR . The roles of water, sanitation and hygiene in reducing schistosomiasis: a review. Parasit Vectors. 2015 Mar 13;8:156.","DOI":"10.1186\/s13071-015-0766-9"},{"key":"2025070713354548000_2025.07.02.662618v1.2","doi-asserted-by":"publisher","DOI":"10.1186\/s40249-021-00919-z"},{"key":"2025070713354548000_2025.07.02.662618v1.3","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0008388"},{"key":"2025070713354548000_2025.07.02.662618v1.4","doi-asserted-by":"publisher","DOI":"10.4269\/ajtmh.19-0193"},{"key":"2025070713354548000_2025.07.02.662618v1.5","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0004794"},{"key":"2025070713354548000_2025.07.02.662618v1.6","doi-asserted-by":"publisher","DOI":"10.1139\/z72-115"},{"key":"2025070713354548000_2025.07.02.662618v1.7","doi-asserted-by":"publisher","DOI":"10.2307\/3276148"},{"key":"2025070713354548000_2025.07.02.662618v1.8","doi-asserted-by":"publisher","DOI":"10.2307\/3280200"},{"key":"2025070713354548000_2025.07.02.662618v1.9","doi-asserted-by":"publisher","DOI":"10.1007\/BF00389946"},{"key":"2025070713354548000_2025.07.02.662618v1.10","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0006948"},{"key":"2025070713354548000_2025.07.02.662618v1.11","doi-asserted-by":"publisher","DOI":"10.1186\/s13071-022-05439-9"},{"key":"2025070713354548000_2025.07.02.662618v1.12","doi-asserted-by":"crossref","unstructured":"Du X , McManus DP , Fogarty CE , Jones MK , You H. Schistosoma mansoni Fibroblast Growth Factor Receptor A Orchestrates Multiple Functions in Schistosome Biology and in the Host- Parasite Interplay. Front Immunol [Internet]. 2022 Jun 22 [cited 2025 Jun 12];13. Available from: https:\/\/www.frontiersin.org\/journals\/immunology\/articles\/10.3389\/fimmu.2022.868077\/full","DOI":"10.3389\/fimmu.2022.868077"},{"key":"2025070713354548000_2025.07.02.662618v1.13","doi-asserted-by":"crossref","unstructured":"Du X , McManus DP , French JD , Collinson N , Sivakumaran H , MacGregor SR , et al. CRISPR interference for sequence-specific regulation of fibroblast growth factor receptor A in Schistosoma mansoni. Front Immunol [Internet]. 2023 Jan 13 [cited 2025 Jun 12];13. Available from: https:\/\/www.frontiersin.org\/journals\/immunology\/articles\/10.3389\/fimmu.2022.1105719\/full","DOI":"10.3389\/fimmu.2022.1105719"},{"key":"2025070713354548000_2025.07.02.662618v1.14","doi-asserted-by":"publisher","DOI":"10.1007\/BF00929167"},{"key":"2025070713354548000_2025.07.02.662618v1.15","doi-asserted-by":"publisher","DOI":"10.2307\/3278950"},{"key":"2025070713354548000_2025.07.02.662618v1.16","doi-asserted-by":"publisher","DOI":"10.2307\/3280200"},{"key":"2025070713354548000_2025.07.02.662618v1.17","doi-asserted-by":"crossref","unstructured":"Miura M , Mitsui Y , Aoki Y , Kato K . A new accumulation assay of Schistosoma mansoni miracidia using square capillary glass tubes. Exp Parasitol. 2022 Aug 1;239:108313.","DOI":"10.1016\/j.exppara.2022.108313"},{"key":"2025070713354548000_2025.07.02.662618v1.18","doi-asserted-by":"publisher","DOI":"10.1038\/260702a0"},{"issue":"4","key":"2025070713354548000_2025.07.02.662618v1.19","first-page":"e4004","article-title":"Host preference of field-derived Schistosoma mansoni is influenced by snail host compatibility and infection status","volume":"13","year":"2022","journal-title":"Ecosphere Wash DC"},{"key":"2025070713354548000_2025.07.02.662618v1.20","doi-asserted-by":"publisher","DOI":"10.1016\/0020-7519(94)00158-K"},{"key":"2025070713354548000_2025.07.02.662618v1.21","doi-asserted-by":"crossref","unstructured":"Fogarty CE , Phan P , Duke MG , McManus DP , Wyeth RC , Cummins SF , et al. Identification of Schistosoma mansoni miracidia attractant candidates in infected Biomphalaria glabrata using behaviour-guided comparative proteomics. Front Immunol. 2022 Oct 10;13:954282.","DOI":"10.3389\/fimmu.2022.954282"},{"key":"2025070713354548000_2025.07.02.662618v1.22","doi-asserted-by":"crossref","unstructured":"Dinguirard N , Heinemann C , Yoshino TP . Mass Isolation and In Vitro Cultivation of Intramolluscan Stages of the Human Blood Fluke Schistosoma Mansoni. J Vis Exp [Internet]. 2018 Jan 1 [cited 2025 Jun 12];(131). Available from: https:\/\/europepmc.org\/articles\/PMC5908647","DOI":"10.3791\/56345-v"},{"key":"2025070713354548000_2025.07.02.662618v1.23","doi-asserted-by":"crossref","unstructured":"Caterer Z , Horejsi RV , Weber C , Mathisen B , Nelson CN , Bagatta M , et al. A graphical user interface for wrmXpress 2.0 streamlines helminth phenotypic screening. Int J Parasitol Drugs Drug Resist. 2025 Apr 1;27:100588.","DOI":"10.1016\/j.ijpddr.2025.100588"},{"key":"2025070713354548000_2025.07.02.662618v1.24","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-021-02997-z"},{"key":"2025070713354548000_2025.07.02.662618v1.25","unstructured":"Allan DB , Caswell T , Keim NC , van der Wel CM , Verweij RW . soft-matter\/trackpy: v0.6.4 [Internet]. Zenodo; 2024 [cited 2025 Apr 2]. Available from: https:\/\/zenodo.org\/records\/12708864"},{"key":"2025070713354548000_2025.07.02.662618v1.26","doi-asserted-by":"crossref","unstructured":"M\u00f6lder F , Jablonski KP , Letcher B , Hall MB , Tomkins-Tinch CH , Sochat V , et al. Sustainable data analysis with Snakemake [Internet]. F1000Research; 2021 [cited 2025 Jun 12]. Available from: https:\/\/f1000research.com\/articles\/10-33","DOI":"10.12688\/f1000research.29032.2"},{"key":"2025070713354548000_2025.07.02.662618v1.27","doi-asserted-by":"crossref","unstructured":"Yoo AB , Jette MA , Grondona M . SLURM: Simple Linux Utility for Resource Management. In: Feitelson D , Rudolph L , Schwiegelshohn U , editors. Job Scheduling Strategies for Parallel Processing. Berlin, Heidelberg: Springer; 2003. p. 44\u201360.","DOI":"10.1007\/10968987_3"},{"key":"2025070713354548000_2025.07.02.662618v1.28","doi-asserted-by":"publisher","DOI":"10.18637\/jss.v067.i0"},{"key":"2025070713354548000_2025.07.02.662618v1.29","unstructured":"Powell M. The BOBYQA algorithm for bound constrained optimization without derivatives. In 2009 [cited 2025 Jun 12]. Available from: https:\/\/www.semanticscholar.org\/paper\/The-BOBYQA-algorithm-for-bound-constrained-without-Powell\/0d2edc46f81f9a0b0b62937507ad977b46729f64"},{"key":"2025070713354548000_2025.07.02.662618v1.30","doi-asserted-by":"publisher","DOI":"10.18637\/jss.v082.i13"},{"key":"2025070713354548000_2025.07.02.662618v1.31","unstructured":"Cohen J . Statistical Power Analysis for the Behavioral Sciences. Rev. ed. Oxford: Lawrence Erlbaum Associates; 1988."},{"issue":"2","key":"2025070713354548000_2025.07.02.662618v1.32","first-page":"65","article-title":"A Simple Sequentially Rejective Multiple Test Procedure","volume":"6","year":"1979","journal-title":"Scand J Stat"},{"key":"2025070713354548000_2025.07.02.662618v1.33","doi-asserted-by":"publisher","DOI":"10.1111\/j.2517-6161.1995.tb02031.x)"},{"key":"2025070713354548000_2025.07.02.662618v1.34","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms15451"},{"key":"2025070713354548000_2025.07.02.662618v1.35","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gku1117"},{"key":"2025070713354548000_2025.07.02.662618v1.36","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gks596"},{"key":"2025070713354548000_2025.07.02.662618v1.37","doi-asserted-by":"publisher","DOI":"10.4269\/ajtmh.23-0235"},{"key":"2025070713354548000_2025.07.02.662618v1.38","unstructured":"Nong W , Yu Y , Aase-Remedios ME , Xie Y , So WL , Li Y , et al. Genome of the ramshorn snail Biomphalaria straminea-an obligate intermediate host of schistosomiasis. GigaScience. 2022 Feb 15;11:giac012."},{"key":"2025070713354548000_2025.07.02.662618v1.39","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0011208"},{"key":"2025070713354548000_2025.07.02.662618v1.40","doi-asserted-by":"crossref","unstructured":"AliView: a fast and lightweight alignment viewer and editor for large datasets | Bioinformatics | Oxford Academic [Internet]. [cited 2025 Mar 31]. Available from: https:\/\/academic.oup.com\/bioinformatics\/article\/30\/22\/3276\/2391211","DOI":"10.1093\/bioinformatics\/btu531"},{"key":"2025070713354548000_2025.07.02.662618v1.41","doi-asserted-by":"publisher","DOI":"10.1016\/j.isci.2022.105357"},{"key":"2025070713354548000_2025.07.02.662618v1.42","doi-asserted-by":"publisher","DOI":"10.1016\/0020-7519(94)00158-K"},{"key":"2025070713354548000_2025.07.02.662618v1.43","unstructured":"Haas W , Haberl B . Host Recognition by Trematode Miracidia and Cercariae. In: Advances in Trematode Biology. CRC Press; 1997."},{"key":"2025070713354548000_2025.07.02.662618v1.44","doi-asserted-by":"publisher","DOI":"10.2307\/3281558"},{"key":"2025070713354548000_2025.07.02.662618v1.45","doi-asserted-by":"publisher","DOI":"10.1078\/0944-2006-00125"},{"key":"2025070713354548000_2025.07.02.662618v1.46","doi-asserted-by":"publisher","DOI":"10.1017\/S0031182000051878"},{"key":"2025070713354548000_2025.07.02.662618v1.47","doi-asserted-by":"publisher","DOI":"10.1016\/0166-6851(86)90125-8"},{"key":"2025070713354548000_2025.07.02.662618v1.48","doi-asserted-by":"publisher","DOI":"10.1016\/0020-7519(94)00158-K"},{"key":"2025070713354548000_2025.07.02.662618v1.49","doi-asserted-by":"publisher","DOI":"10.1093\/oxfordjournals.molbev.a003769"},{"key":"2025070713354548000_2025.07.02.662618v1.50","doi-asserted-by":"publisher","DOI":"10.1186\/s12864-024-10103-w"},{"key":"2025070713354548000_2025.07.02.662618v1.51","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-28634-9"},{"key":"2025070713354548000_2025.07.02.662618v1.52","doi-asserted-by":"publisher","DOI":"10.1186\/s40249-024-01187-3"},{"key":"2025070713354548000_2025.07.02.662618v1.53","unstructured":"Executive summary. In: WHO guideline on control and elimination of human schistosomiasis [Internet] [Internet]. World Health Organization; 2022 [cited 2025 Jun 16]. Available from: https:\/\/www.ncbi.nlm.nih.gov\/books\/NBK578389\/"},{"key":"2025070713354548000_2025.07.02.662618v1.54","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pntd.0007833"},{"key":"2025070713354548000_2025.07.02.662618v1.55","doi-asserted-by":"publisher","DOI":"10.2307\/3279673"},{"key":"2025070713354548000_2025.07.02.662618v1.56","doi-asserted-by":"publisher","DOI":"10.1186\/1471-2121-12-6"},{"key":"2025070713354548000_2025.07.02.662618v1.57","doi-asserted-by":"crossref","unstructured":"Gomez-Marin A , Stephens GJ , Louis M . Active sampling and decision making in Drosophila chemotaxis. Nat Commun. 2011 Aug 23;2:441.","DOI":"10.1038\/ncomms1455"},{"key":"2025070713354548000_2025.07.02.662618v1.58","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.19-21-09557.1999"},{"key":"2025070713354548000_2025.07.02.662618v1.59","doi-asserted-by":"crossref","unstructured":"Ram\u00edrez-G\u00f3mez HV , Jimenez Sabinina V , Vel\u00e1zquez P\u00e9rez M , Beltran C , Carneiro J , Wood CD , et al. Sperm chemotaxis is driven by the slope of the chemoattractant concentration field. Barkai N, Goldstein RE, Dunkel J, Austin R, editors. eLife. 2020 Mar 9;9:e50532.","DOI":"10.7554\/eLife.50532"},{"key":"2025070713354548000_2025.07.02.662618v1.60","doi-asserted-by":"crossref","unstructured":"Attenborough T , Rawlinson KA , Diaz Soria CL , Ambridge K , Sankaranarayanan G , Graham J , et al. A single-cell atlas of the miracidium larva of Schistosoma mansoni reveals cell types, developmental pathways, and tissue architecture. Banerjee U, editor. eLife. 2024 Aug 27;13:RP95628.","DOI":"10.7554\/eLife.95628"},{"key":"2025070713354548000_2025.07.02.662618v1.61","doi-asserted-by":"publisher","DOI":"10.1371\/journal.ppat.1006718"},{"key":"2025070713354548000_2025.07.02.662618v1.62","doi-asserted-by":"publisher","DOI":"10.17912\/micropub.biology.000704"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1101\/2025.07.02.662618","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,1,15]],"date-time":"2026-01-15T09:03:25Z","timestamp":1768467805000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.1101\/2025.07.02.662618"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,7,3]]},"references-count":62,"URL":"https:\/\/doi.org\/10.1101\/2025.07.02.662618","relation":{},"subject":[],"published":{"date-parts":[[2025,7,3]]},"subtype":"preprint"}}