{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T15:28:18Z","timestamp":1787153298799,"version":"build-2736575974"},"posted":{"date-parts":[[2026,8,16]]},"group-title":"Animal Behavior and Cognition","reference-count":105,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2026,8,16]],"date-time":"2026-08-16T00:00:00Z","timestamp":1786838400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"name":"CNRS","award":["Momentum grant"],"award-info":[{"award-number":["Momentum grant"]}]},{"DOI":"10.13039\/501100003135","name":"Fyssen Foundation","doi-asserted-by":"crossref","award":["Research Grant"],"award-info":[{"award-number":["Research Grant"]}],"id":[{"id":"10.13039\/501100003135","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100003135","name":"Fyssen Foundation","doi-asserted-by":"crossref","award":["Postdoctoral Grant"],"award-info":[{"award-number":["Postdoctoral Grant"]}],"id":[{"id":"10.13039\/501100003135","id-type":"DOI","asserted-by":"crossref"}]},{"award":["ANR-22-CE02-0002 (ForAnInstant)"],"award-info":[{"award-number":["ANR-22-CE02-0002 (ForAnInstant)"]}],"id":[{"id":"https:\/\/ror.org\/00rbzpz17","id-type":"ROR","asserted-by":"publisher"}]},{"name":"SEVAB Doctoral School","award":["MESRI PhD grant"],"award-info":[{"award-number":["MESRI PhD grant"]}]},{"award":["BES-2017-079643"],"award-info":[{"award-number":["BES-2017-079643"]}],"id":[{"id":"https:\/\/ror.org\/05r0vyz12","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2026,8,16]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                <jats:p>\n                  The Marginal Value Theorem (MVT) is an important part of Optimal Foraging Theory, predicting the optimal time to leave a food patch. It has been mostly studied in birds, insects and mammals, even though simpler organisms also need to forage efficiently in patchy environments. Here we test whether the nematode\n                  <jats:italic>Caenorhabditis elegans<\/jats:italic>\n                  implements the MVT. We recorded individual nematodes exploring patchy environments, across four inter-patch distances and three different food qualities, and found that\n                  <jats:italic>C. elegans<\/jats:italic>\n                  behavior matches MVT predictions: When food patches are further away, each food patch is exploited for a longer time. In previous studies animals achieved this by modulating the duration of visits to food patches. Similarly, we found that\n                  <jats:italic>C. elegans<\/jats:italic>\n                  also increases visit duration with inter-patch distance, but this only accounts for half of the increase in total exploitation time. The other half of the increase comes from\n                  <jats:italic>C. elegans<\/jats:italic>\n                  revisiting food patches multiple times, and the number of these revisits increasing with inter-patch distance. This increase in the number of revisits is not due to behavioral changes in response to distance, but rather to a passive interaction between trajectories and environment geometry. These results show that\n                  <jats:italic>C. elegans<\/jats:italic>\n                  can learn the statistics of an environment and use this information in a way consistent with the MVT, but also that part of the fitness-relevant outcomes can emerge passively.\n                <\/jats:p>\n                <jats:sec>\n                  <jats:title>SIGNIFICANCE<\/jats:title>\n                  <jats:p>\n                    Despite being key in understanding foraging in patchy resources, the Marginal Value Theorem (MVT) has been tested almost exclusively in relatively complex animals. We extensively tested the MVT in a simple, non-visual organism, showing that\n                    <jats:italic>Caenorhabditis elegans<\/jats:italic>\n                    increases patch exploitation time when inter-patch distance increases. This effect is partially driven by the same behavioral adaptation found in complex animals, but also by an increase in the number of patch revisits. This second driver, which had not been reported before and is probably key for non-visual organisms, requires no behavioral adaptation and produces around half of the fitness-relevant outcome. Our results highlight the need for adapting Optimal Foraging Theory to a wide range of taxa spanning from microbes to small invertebrates.\n                  <\/jats:p>\n                <\/jats:sec>","DOI":"10.64898\/2026.08.10.743854","type":"posted-content","created":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T04:40:15Z","timestamp":1786941615000},"source":"Crossref","is-referenced-by-count":0,"title":["The Marginal Value Theorem in\n                  <i>Caenorhabditis elegans<\/i>"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7951-2110","authenticated-orcid":false,"given":"Alid","family":"Al-Asmar","sequence":"first","affiliation":[{"name":"Universit\u00e9 de Toulouse, Centre National de la Recherche Scientifique, Centre de Recherches sur la Cognition Animale, Centre de Biologie Int\u00e9grative, Toulouse 31062, France"},{"name":"Laboratory of Mathematical and Physical Ethology, Research Center of Mathematics for Social Creativity, Research Institute for Electronic Science, Hokkaido University, Sapporo, Hokkaido Prefecture, Japan"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4806-0435","authenticated-orcid":false,"given":"Roger","family":"Lloret-Cabot","sequence":"additional","affiliation":[{"name":"Universit\u00e9 de Toulouse, Centre National de la Recherche Scientifique, Centre de Recherches sur la Cognition Animale, Centre de Biologie Int\u00e9grative, Toulouse 31062, France"},{"name":"Centre d\u2019Estudis Avan\u00e7ats de Blanes (CEAB-CSIC), Cala Sant Francesc 14, 17300 Girona, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4782-6139","authenticated-orcid":false,"given":"Alfonso","family":"P\u00e9rez-Escudero","sequence":"additional","affiliation":[{"name":"Universit\u00e9 de Toulouse, Centre National de la Recherche Scientifique, Centre de Recherches sur la Cognition Animale, Centre de Biologie Int\u00e9grative, Toulouse 31062, France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026081908050732000_2026.08.10.743854v1.1","first-page":"411","article-title":"Departure time versus departure rate: How to forage optimally when you are stupid","volume":"1","year":"1999","journal-title":"Evolutionary Ecology Research"},{"issue":"2","key":"2026081908050732000_2026.08.10.743854v1.2","doi-asserted-by":"crossref","first-page":"275","DOI":"10.1038\/s41567-020-01036-8","article-title":"Capturing the continuous complexity of behaviour in Caenorhabditis elegans","volume":"17","year":"2021","journal-title":"Nature Physics"},{"key":"2026081908050732000_2026.08.10.743854v1.3","doi-asserted-by":"crossref","unstructured":"Al-Asmar, A. , Madirolas, G. , Bello, M. 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