{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,27]],"date-time":"2026-10-27T08:31:23Z","timestamp":1793089883999,"version":"4.5.0"},"reference-count":16,"publisher":"National Academy of Sciences","issue":"33","license":[{"start":{"date-parts":[[2022,8,8]],"date-time":"2022-08-08T00:00:00Z","timestamp":1659916800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.pnas.org"],"crossmark-restriction":true},"short-container-title":["Proc. Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2022,8,16]]},"abstract":"<jats:p>Sleep and sleep-like states are present across the animal kingdom, with recent studies convincingly demonstrating sleep-like states in arthropods, nematodes, and even cnidarians. However, the existence of different sleep phases across taxa is as yet unclear. In particular, the study of rapid eye movement (REM) sleep is still largely centered on terrestrial vertebrates, particularly mammals and birds. The most salient indicator of REM sleep is the movement of eyes during this phase. Movable eyes, however, have evolved only in a limited number of lineages\u2014an adaptation notably absent in insects and most terrestrial arthropods\u2014restricting cross-species comparisons. Jumping spiders, however, possess movable retinal tubes to redirect gaze, and in newly emerged spiderlings, these movements can be directly observed through their temporarily translucent exoskeleton. Here, we report evidence for an REM sleep\u2013like state in a terrestrial invertebrate: periodic bouts of retinal movements coupled with limb twitching and stereotyped leg curling behaviors during nocturnal resting in a jumping spider. Observed retinal movement bouts were consistent, including regular durations and intervals, with both increasing over the course of the night. That these characteristic REM sleep\u2013like behaviors exist in a highly visual, long-diverged lineage further challenges our understanding of this sleep state. Comparisons across such long-diverged lineages likely hold important questions and answers about the visual brain as well as the origin, evolution, and function of REM sleep.<\/jats:p>","DOI":"10.1073\/pnas.2204754119","type":"journal-article","created":{"date-parts":[[2022,8,8]],"date-time":"2022-08-08T15:16:07Z","timestamp":1659971767000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":54,"title":["Regularly occurring bouts of retinal movements suggest an REM sleep\u2013like state in jumping spiders"],"prefix":"10.1073","volume":"119","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4678-2231","authenticated-orcid":false,"given":"Daniela C.","family":"R\u00f6\u00dfler","sequence":"first","affiliation":[{"name":"Zukunftskolleg, University of Konstanz, Konstanz, 78464 Germany"},{"name":"Department of Biology, University of Konstanz, Konstanz, 78464 Germany"},{"name":"Department of Collective Behavior, Max Planck Institute of Animal Behavior, Konstanz, 78464 Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1762-5982","authenticated-orcid":false,"given":"Kris","family":"Kim","sequence":"additional","affiliation":[{"name":"John Harvard Distinguished Science Fellows Program, Harvard University, Cambridge, MA 02138"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9284-5964","authenticated-orcid":false,"given":"Massimo","family":"De Agr\u00f2","sequence":"additional","affiliation":[{"name":"Department of Biology, University of Florence, 50121 Florence, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6131-9734","authenticated-orcid":false,"given":"Alex","family":"Jordan","sequence":"additional","affiliation":[{"name":"Department of Collective Behavior, Max Planck Institute of Animal Behavior, Konstanz, 78464 Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8292-6031","authenticated-orcid":false,"given":"C Giovanni","family":"Galizia","sequence":"additional","affiliation":[{"name":"Zukunftskolleg, University of Konstanz, Konstanz, 78464 Germany"},{"name":"Department of Biology, University of Konstanz, Konstanz, 78464 Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4639-9339","authenticated-orcid":false,"given":"Paul S.","family":"Shamble","sequence":"additional","affiliation":[{"name":"John Harvard Distinguished Science Fellows Program, Harvard University, Cambridge, MA 02138"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"341","published-online":{"date-parts":[[2022,8,8]]},"reference":[{"key":"e_1_3_4_1_2","doi-asserted-by":"publisher","DOI":"10.1126\/science.118.3062.273"},{"key":"e_1_3_4_2_2","doi-asserted-by":"publisher","DOI":"10.1016\/B0-72-160797-7\/50009-4"},{"key":"e_1_3_4_3_2","first-page":"jeb174862","article-title":"Cyclic nature of the REM sleep-like state in the cuttlefish Sepia officinalis","volume":"222","author":"Iglesias T. L.","year":"2019","unstructured":"T. L. Iglesias, J. G. Boal, M. G. Frank, J. Zeil, R. T. Hanlon, Cyclic nature of the REM sleep-like state in the cuttlefish Sepia officinalis. J. Exp. Biol. 222, jeb174862 (2019).","journal-title":"J. Exp. Biol."},{"key":"e_1_3_4_4_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2019.11.045"},{"key":"e_1_3_4_5_2","doi-asserted-by":"publisher","DOI":"10.3389\/fpsyg.2020.567618"},{"key":"e_1_3_4_6_2","doi-asserted-by":"publisher","DOI":"10.1186\/s12983-021-00410-3"},{"key":"e_1_3_4_7_2","doi-asserted-by":"publisher","DOI":"10.1242\/jeb.51.2.471"},{"key":"e_1_3_4_8_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2016.03.013"},{"key":"e_1_3_4_9_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2017.10.026"},{"key":"e_1_3_4_10_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.anbehav.2005.01.008"},{"key":"e_1_3_4_11_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2018.05.022"},{"key":"e_1_3_4_12_2","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms12468"},{"key":"e_1_3_4_13_2","first-page":"367","article-title":"The \u2018scanning hypothesis\u2019 of rapid eye movements during REM sleep: A review of the evidence","volume":"149","author":"Arnulf I.","year":"2011","unstructured":"I. Arnulf, The \u2018scanning hypothesis\u2019 of rapid eye movements during REM sleep: A review of the evidence. Arch. Ital. Biol. 149, 367\u2013382 (2011).","journal-title":"Arch. Ital. Biol."},{"key":"e_1_3_4_14_2","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2015.09.069"},{"key":"e_1_3_4_15_2","doi-asserted-by":"publisher","DOI":"10.1007\/s12264-013-1313-3"},{"key":"e_1_3_4_16_2","unstructured":"D. C. R\u00f6\u00dfler Data for: Regularly occurring bouts of retinal movements suggest an REM sleep\u2013like state in jumping spiders. Zenodo. https:\/\/doi.org\/10.5281\/zenodo.6616655. Deposited 6 June 2022."}],"container-title":["Proceedings of the National Academy of Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pnas.org\/doi\/pdf\/10.1073\/pnas.2204754119","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,8,8]],"date-time":"2022-08-08T15:22:23Z","timestamp":1659972143000},"score":1,"resource":{"primary":{"URL":"https:\/\/pnas.org\/doi\/full\/10.1073\/pnas.2204754119"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2022,8,8]]},"references-count":16,"journal-issue":{"issue":"33","published-print":{"date-parts":[[2022,8,16]]}},"alternative-id":["10.1073\/pnas.2204754119"],"URL":"https:\/\/doi.org\/10.1073\/pnas.2204754119","relation":{},"ISSN":["0027-8424","1091-6490"],"issn-type":[{"value":"0027-8424","type":"print"},{"value":"1091-6490","type":"electronic"}],"subject":[],"published":{"date-parts":[[2022,8,8]]},"assertion":[{"value":"2022-03-22","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2022-05-27","order":1,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2022-08-08","order":2,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e2204754119"}}