{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"institution":[{"name":"bioRxiv"}],"indexed":{"date-parts":[[2026,9,12]],"date-time":"2026-09-12T01:29:01Z","timestamp":1789176541480,"version":"build-2803163510"},"posted":{"date-parts":[[2025,2,26]]},"group-title":"Neuroscience","reference-count":68,"publisher":"openRxiv","license":[{"start":{"date-parts":[[2025,2,26]],"date-time":"2025-02-26T00:00:00Z","timestamp":1740528000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"}],"funder":[{"award":["IA\/S\/19\/2\/504649"],"award-info":[{"award-number":["IA\/S\/19\/2\/504649"]}],"id":[{"id":"https:\/\/ror.org\/04reqzt68","id-type":"ROR","asserted-by":"publisher"}]},{"name":"MoE","award":["MoE\/STARS-1\/454"],"award-info":[{"award-number":["MoE\/STARS-1\/454"]}]},{"name":"Department of Biotechnology","award":["BT\/HRD-NBA-NWB\/38\/2019-20"],"award-info":[{"award-number":["BT\/HRD-NBA-NWB\/38\/2019-20"]}]},{"name":"ANRF","award":["SPG\/2022\/000182"],"award-info":[{"award-number":["SPG\/2022\/000182"]}]},{"name":"ANRF","award":["CRG\/2023\/001950"],"award-info":[{"award-number":["CRG\/2023\/001950"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"accepted":{"date-parts":[[2025,6,13]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                <jats:p>\n                  Memory formation is necessary for the survival of animals across phyla. Here, we elucidate the mechanism underlying the formation of long-term associative memory (LTAM) formed by treating\n                  <jats:italic>Caenorhabditis elegans<\/jats:italic>\n                  with a volatile chemoattractant and heat. Previous work has shown that training animals with a paradigm involving heat and isoamyl alcohol (IAA) simultaneously, causes\n                  <jats:italic>C. elegans<\/jats:italic>\n                  to lose their attraction to IAA. In this study, we elaborate on the mechanism behind this LTAM formation and suggest that during training with heat and IAA,\n                  <jats:italic>C. elegans<\/jats:italic>\n                  release extracellular vesicles (EVs) that upon being taken up by the same trained animals or their untrained counterparts causes the organism to lose attraction to IAA. Our data suggests that the vesicles are highly specific to the training paradigms used and differ with differing cues. Finally, we show that this mechanism of transfer of LTAM appears to be conserved between\n                  <jats:italic>C. elegans<\/jats:italic>\n                  and\n                  <jats:italic>C. briggsae<\/jats:italic>\n                  allowing for both intra and interspecies transfer of memory.\n                <\/jats:p>","DOI":"10.1101\/2025.02.26.640282","type":"posted-content","created":{"date-parts":[[2025,2,26]],"date-time":"2025-02-26T14:50:23Z","timestamp":1740581423000},"source":"Crossref","is-referenced-by-count":2,"title":["Extracellular vesicles aid in the transfer of long-term associative memory between\n                  <i>Caenorhabditis elegans<\/i>"],"prefix":"10.64898","author":[{"given":"Monmita","family":"Bhar","sequence":"first","affiliation":[{"name":"Centre for Neuroscience, Indian Institute of Science (IISc), Bangalore, Karnataka, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tanumoy","family":"Nandi","sequence":"additional","affiliation":[{"name":"Centre for Neuroscience, Indian Institute of Science (IISc), Bangalore, Karnataka, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hari","family":"Narayanan","sequence":"additional","affiliation":[{"name":"Centre for Neuroscience, Indian Institute of Science (IISc), Bangalore, Karnataka, India"},{"name":"Present address: University of Konstanz, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kamal","family":"Kishore","sequence":"additional","affiliation":[{"name":"Centre for Neuroscience, Indian Institute of Science (IISc), Bangalore, Karnataka, India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7527-3666","authenticated-orcid":false,"given":"Kavita","family":"Babu","sequence":"additional","affiliation":[{"name":"Centre for Neuroscience, Indian Institute of Science (IISc), Bangalore, Karnataka, India"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026041113064515000_2025.02.26.640282v3.1","doi-asserted-by":"publisher","DOI":"10.1101\/lm.2224411"},{"key":"2026041113064515000_2025.02.26.640282v3.2","doi-asserted-by":"publisher","DOI":"10.1101\/lm.960510"},{"key":"2026041113064515000_2025.02.26.640282v3.3","doi-asserted-by":"publisher","DOI":"10.1038\/srep29284"},{"key":"2026041113064515000_2025.02.26.640282v3.4","doi-asserted-by":"crossref","unstructured":"Bargmann CI (2006) Chemosensation in C. elegans. WormBook: 1\u201329","DOI":"10.1895\/wormbook.1.123.1"},{"key":"2026041113064515000_2025.02.26.640282v3.5","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(00)81752-3"},{"key":"2026041113064515000_2025.02.26.640282v3.6","doi-asserted-by":"publisher","DOI":"10.1002\/dev.20277"},{"key":"2026041113064515000_2025.02.26.640282v3.7","doi-asserted-by":"crossref","unstructured":"Bedecarrats A , Chen S , Pearce K , Cai D , Glanzman DL (2018) RNA from Trained Aplysia Can Induce an Epigenetic Engram for Long-Term Sensitization in Untrained Aplysia. eNeuro 5","DOI":"10.1523\/ENEURO.0038-18.2018"},{"key":"2026041113064515000_2025.02.26.640282v3.8","doi-asserted-by":"publisher","DOI":"10.1051\/medsci\/20031967725"},{"key":"2026041113064515000_2025.02.26.640282v3.9","doi-asserted-by":"publisher","DOI":"10.1007\/BF01733207"},{"key":"2026041113064515000_2025.02.26.640282v3.10","doi-asserted-by":"publisher","DOI":"10.1101\/SQB.1996.061.01.018"},{"key":"2026041113064515000_2025.02.26.640282v3.11","doi-asserted-by":"publisher","DOI":"10.1016\/0896-6273(95)90224-4"},{"key":"2026041113064515000_2025.02.26.640282v3.12","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.0006-19.2019"},{"key":"2026041113064515000_2025.02.26.640282v3.13","doi-asserted-by":"publisher","DOI":"10.1038\/345718a0"},{"key":"2026041113064515000_2025.02.26.640282v3.14","doi-asserted-by":"publisher","DOI":"10.1104\/pp.15.01217"},{"key":"2026041113064515000_2025.02.26.640282v3.15","doi-asserted-by":"publisher","DOI":"10.1186\/s12870-017-1022-6"},{"key":"2026041113064515000_2025.02.26.640282v3.16","doi-asserted-by":"publisher","DOI":"10.1038\/s41592-022-01507-1"},{"key":"2026041113064515000_2025.02.26.640282v3.17","doi-asserted-by":"crossref","unstructured":"Ferkey DM , Sengupta P , L\u2019Etoile ND (2021) Chemosensory signal transduction in Caenorhabditis elegans. Genetics 217","DOI":"10.1093\/genetics\/iyab004"},{"key":"2026041113064515000_2025.02.26.640282v3.18","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2015.05.034"},{"key":"2026041113064515000_2025.02.26.640282v3.19","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.neuro.31.060407.125631"},{"key":"2026041113064515000_2025.02.26.640282v3.20","doi-asserted-by":"publisher","DOI":"10.1007\/BF00332953"},{"key":"2026041113064515000_2025.02.26.640282v3.21","doi-asserted-by":"crossref","unstructured":"Hart AC (2006) Behavior. WormBook","DOI":"10.1895\/wormbook.1.87.1"},{"key":"2026041113064515000_2025.02.26.640282v3.22","doi-asserted-by":"publisher","DOI":"10.1186\/s40793-018-0333-z"},{"key":"2026041113064515000_2025.02.26.640282v3.23","doi-asserted-by":"publisher","DOI":"10.1016\/j.cellsig.2004.05.001"},{"key":"2026041113064515000_2025.02.26.640282v3.24","doi-asserted-by":"publisher","DOI":"10.1534\/genetics.107.082651"},{"key":"2026041113064515000_2025.02.26.640282v3.25","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.physiol.68.040204.100715"},{"key":"2026041113064515000_2025.02.26.640282v3.26","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-020-2699-5"},{"key":"2026041113064515000_2025.02.26.640282v3.27","doi-asserted-by":"crossref","unstructured":"Kauffman A , Parsons L , Stein G , Wills A , Kaletsky R , Murphy C (2011) C. elegans positive butanone learning, short-term, and long-term associative memory assays. J Vis Exp","DOI":"10.3791\/2490-v"},{"key":"2026041113064515000_2025.02.26.640282v3.28","doi-asserted-by":"crossref","first-page":"11","DOI":"10.1016\/j.toxlet.2020.05.017","article-title":"Glycochenodeoxycholic acid impairs transcription factor E3-dependent autophagy-lysosome machinery by disrupting reactive oxygen species homeostasis in L02 cells","volume":"331","year":"2020","journal-title":"Toxicol Lett"},{"key":"2026041113064515000_2025.02.26.640282v3.29","doi-asserted-by":"publisher","DOI":"10.1016\/j.isci.2022.104627"},{"key":"2026041113064515000_2025.02.26.640282v3.30","doi-asserted-by":"crossref","unstructured":"Lobo T , Haasnoot GH , Nawrocka A , Bruggeman CW , Razzauti A , Peterman EJG , Laurent P (2025) Sensory stimuli and cilium trafficking defects trigger the release of ciliary extracellular vesicles from multiple ciliary locations. bioRxiv: 2025.2006.2006.658075","DOI":"10.1101\/2025.06.06.658075"},{"key":"2026041113064515000_2025.02.26.640282v3.31","doi-asserted-by":"publisher","DOI":"10.1091\/mbc.E15-01-0009"},{"key":"2026041113064515000_2025.02.26.640282v3.32","doi-asserted-by":"publisher","DOI":"10.1016\/bs.ctdb.2021.03.001"},{"key":"2026041113064515000_2025.02.26.640282v3.33","doi-asserted-by":"publisher","DOI":"10.1039\/D4NA00393D"},{"key":"2026041113064515000_2025.02.26.640282v3.34","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2021.07.022"},{"key":"2026041113064515000_2025.02.26.640282v3.35","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2011.06.027"},{"key":"2026041113064515000_2025.02.26.640282v3.36","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2022.03.005"},{"key":"2026041113064515000_2025.02.26.640282v3.37","doi-asserted-by":"publisher","DOI":"10.3389\/fnbeh.2017.00080"},{"key":"2026041113064515000_2025.02.26.640282v3.38","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2017.09.006 29129530"},{"key":"2026041113064515000_2025.02.26.640282v3.39","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2004.12.073"},{"key":"2026041113064515000_2025.02.26.640282v3.40","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4221-11.2012"},{"key":"2026041113064515000_2025.02.26.640282v3.41","doi-asserted-by":"crossref","unstructured":"Porta-de-la-Riva M , Fontrodona L , Villanueva A , Ceron J (2012) Basic Caenorhabditis elegans methods: synchronization and observation. J Vis Exp: e4019","DOI":"10.3791\/4019-v"},{"key":"2026041113064515000_2025.02.26.640282v3.42","doi-asserted-by":"crossref","unstructured":"Qi B , Kniazeva M , Han M (2017) A vitamin-B2-sensing mechanism that regulates gut protease activity to impact animal\u2019s food behavior and growth. Elife 6","DOI":"10.7554\/eLife.26243"},{"key":"2026041113064515000_2025.02.26.640282v3.43","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2012.02.057"},{"key":"2026041113064515000_2025.02.26.640282v3.44","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2009.05.029"},{"key":"2026041113064515000_2025.02.26.640282v3.45","doi-asserted-by":"crossref","unstructured":"Razzauti A , Laurent P (2021) Ectocytosis prevents accumulation of ciliary cargo in C. elegans sensory neurons. Elife 10","DOI":"10.7554\/eLife.67670"},{"key":"2026041113064515000_2025.02.26.640282v3.46","doi-asserted-by":"publisher","DOI":"10.1007\/s11357-020-00204-1"},{"key":"2026041113064515000_2025.02.26.640282v3.47","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth.2019"},{"key":"2026041113064515000_2025.02.26.640282v3.48","doi-asserted-by":"publisher","DOI":"10.1016\/S0092-8674(00)81068-5"},{"key":"2026041113064515000_2025.02.26.640282v3.49","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(94)90028-0"},{"key":"2026041113064515000_2025.02.26.640282v3.50","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pgen.1011178"},{"key":"2026041113064515000_2025.02.26.640282v3.51","doi-asserted-by":"crossref","unstructured":"Seto R , Brown R , Kaletsky R , Parsons LR , Moore RS , Murphy CT (2024) Pseudomonas fluorescens 15 small RNA Pfs1 mediates transgenerational epigenetic inheritance of pathogen avoidance in C. elegans through the Ephrin receptor VAB-1. bioRxiv","DOI":"10.1101\/2024.05.23.595334"},{"key":"2026041113064515000_2025.02.26.640282v3.52","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.neuro.21.1.127"},{"key":"2026041113064515000_2025.02.26.640282v3.53","doi-asserted-by":"publisher","DOI":"10.3758\/BF03200077"},{"key":"2026041113064515000_2025.02.26.640282v3.54","doi-asserted-by":"publisher","DOI":"10.1016\/j.celrep.2018.10.053"},{"key":"2026041113064515000_2025.02.26.640282v3.55","doi-asserted-by":"publisher","DOI":"10.1037\/a0024370"},{"key":"2026041113064515000_2025.02.26.640282v3.56","doi-asserted-by":"crossref","unstructured":"Wang J , Barr MM , Wehman AM (2024a) Extracellular vesicles. Genetics 227","DOI":"10.1093\/genetics\/iyae088"},{"key":"2026041113064515000_2025.02.26.640282v3.57","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2021.06.040"},{"key":"2026041113064515000_2025.02.26.640282v3.58","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2024.05.015"},{"key":"2026041113064515000_2025.02.26.640282v3.59","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2014.01.002"},{"key":"2026041113064515000_2025.02.26.640282v3.60","doi-asserted-by":"publisher","DOI":"10.1186\/s12964-025-02074-6"},{"key":"2026041113064515000_2025.02.26.640282v3.61","doi-asserted-by":"publisher","DOI":"10.1007\/s00018-024-05522-7"},{"key":"2026041113064515000_2025.02.26.640282v3.62","doi-asserted-by":"publisher","DOI":"10.1016\/j.cell.2024.11.018"},{"key":"2026041113064515000_2025.02.26.640282v3.63","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-022-05561-9"},{"key":"2026041113064515000_2025.02.26.640282v3.64","doi-asserted-by":"crossref","unstructured":"Yang B , Wang J , Zheng X , Wang X (2023) Nematode Pheromones: Structures and Functions. Molecules 28","DOI":"10.3390\/molecules28052409"},{"key":"2026041113064515000_2025.02.26.640282v3.65","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(95)90375-5"},{"key":"2026041113064515000_2025.02.26.640282v3.66","doi-asserted-by":"publisher","DOI":"10.1016\/S0959-4388(96)80082-1"},{"key":"2026041113064515000_2025.02.26.640282v3.67","doi-asserted-by":"publisher","DOI":"10.1016\/0092-8674(94)90399-9"},{"key":"2026041113064515000_2025.02.26.640282v3.68","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M116.741132"}],"container-title":[],"original-title":[],"link":[{"URL":"https:\/\/syndication.highwire.org\/content\/doi\/10.1101\/2025.02.26.640282","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,4,11]],"date-time":"2026-04-11T20:06:57Z","timestamp":1775938017000},"score":1,"resource":{"primary":{"URL":"http:\/\/biorxiv.org\/lookup\/doi\/10.1101\/2025.02.26.640282"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,2,26]]},"references-count":68,"URL":"https:\/\/doi.org\/10.1101\/2025.02.26.640282","relation":{},"subject":[],"published":{"date-parts":[[2025,2,26]]},"subtype":"preprint"}}