{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T12:04:30Z","timestamp":1790337870232,"version":"4.1.0"},"reference-count":39,"publisher":"Wiley","issue":"5","license":[{"start":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T00:00:00Z","timestamp":1789430400000},"content-version":"vor","delay-in-days":14,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T00:00:00Z","timestamp":1788220800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["J Eukaryotic Microbiology"],"published-print":{"date-parts":[[2026,9]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>\n                    For decades, the capacity for associative learning was considered a hallmark of multicellular organisms possessing complex nervous systems. However, a growing body of evidence in the field of basal cognition is challenging this neuro\u2010centric paradigm, demonstrating that unicellular eukaryotes, or protozoans, exhibit sophisticated behavioral plasticity. This review synthesizes historical and contemporary research into associative learning\u2014specifically classical and operant conditioning\u2014in prominent protozoan models such as Paramecium, Stentor, and the acellular slime mold\n                    <jats:styled-content style=\"fixed-case\">\n                      <jats:italic>Physarum polycephalum<\/jats:italic>\n                    <\/jats:styled-content>\n                    . We examine the experimental frameworks that have successfully demonstrated anticipatory behavior and stimulus\u2013response pairing, while critically addressing the methodological skepticism that has historically clouded the field. Furthermore, we explore the hypothesized subcellular mechanisms driving these processes, including cytoskeletal restructuring, epigenetic modifications, and biochemical signaling cascades that serve as a non\u2010neural \u201cmemory.\u201d By reevaluating the cognitive boundaries of the single cell, this paper argues that learning is a fundamental property of biological systems rather than a specialized function of neural tissue. The findings discussed herein have significant implications for our understanding of the evolution of intelligence, the development of synthetic biological systems, and the broadening definition of biological memory.\n                  <\/jats:p>","DOI":"10.1111\/jeu.70120","type":"journal-article","created":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T08:37:21Z","timestamp":1789461441000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Learning Without Neurons: A Review of Associative Memory in Protists"],"prefix":"10.1111","volume":"73","author":[{"ORCID":"https:\/\/orcid.org\/0009-0006-9438-3082","authenticated-orcid":false,"given":"Yashika","family":"Jorwal","sequence":"first","affiliation":[{"name":"Loreto Convent School  New Delhi India"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,15]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1305049110"},{"key":"e_1_2_10_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/451385a"},{"key":"e_1_2_10_4_1","doi-asserted-by":"publisher","DOI":"10.3389\/fpsyg.2016.00902"},{"key":"e_1_2_10_5_1","doi-asserted-by":"publisher","DOI":"10.1080\/19420889.2018.1489184"},{"key":"e_1_2_10_6_1","doi-asserted-by":"publisher","DOI":"10.7551\/mitpress\/14642.003.0006"},{"key":"e_1_2_10_7_1","doi-asserted-by":"publisher","DOI":"10.1002\/bies.201800229"},{"key":"e_1_2_10_8_1","doi-asserted-by":"publisher","DOI":"10.21468\/SciPostPhysLectNotes.101"},{"key":"e_1_2_10_9_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.isci.2021.102131"},{"key":"e_1_2_10_10_1","doi-asserted-by":"publisher","DOI":"10.1101\/SQB.1992.057.01.018"},{"key":"e_1_2_10_11_1","doi-asserted-by":"publisher","DOI":"10.1186\/s13408-018-0067-7"},{"key":"e_1_2_10_12_1","doi-asserted-by":"publisher","DOI":"10.1002\/0471142727.mb2129s109"},{"key":"e_1_2_10_13_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-019-11677-w"},{"key":"e_1_2_10_14_1","unstructured":"Dehghani N. andM.Levin.2024.\u201cBio\u2010Inspired AI: Integrating Biological Complexity Into Artificial Intelligence.\u201darXiv 2411.15243."},{"key":"e_1_2_10_15_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2020.05.036"},{"key":"e_1_2_10_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2024.10.041"},{"key":"e_1_2_10_17_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.biosystems.2005.06.011"},{"key":"e_1_2_10_18_1","doi-asserted-by":"publisher","DOI":"10.1037\/h0063093"},{"key":"e_1_2_10_19_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.61907"},{"issue":"1","key":"e_1_2_10_20_1","first-page":"3","article-title":"Learning, Memory and the Brain","volume":"35","author":"Horn G.","year":"1991","journal-title":"Indian Journal of Physiology and Pharmacology"},{"key":"e_1_2_10_21_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41580-020-0214-3"},{"key":"e_1_2_10_22_1","doi-asserted-by":"publisher","DOI":"10.1126\/scisignal.aay3585"},{"key":"e_1_2_10_23_1","doi-asserted-by":"publisher","DOI":"10.3389\/fpsyg.2019.02688"},{"key":"e_1_2_10_24_1","doi-asserted-by":"publisher","DOI":"10.1037\/xan0000306"},{"key":"e_1_2_10_25_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10339-005-0016-8"},{"key":"e_1_2_10_26_1","doi-asserted-by":"publisher","DOI":"10.3389\/fmicb.2015.00264"},{"issue":"41","key":"e_1_2_10_27_1","article-title":"Form Follows Function: Network Topology and Transport in \n                     Physarum polycephalum","volume":"50","author":"Oettmeier C.","year":"2017","journal-title":"Journal of Physics D: Applied Physics"},{"key":"e_1_2_10_28_1","doi-asserted-by":"publisher","DOI":"10.1038\/s44319-023-00004-6"},{"key":"e_1_2_10_29_1","doi-asserted-by":"publisher","DOI":"10.1007\/s10071-023-01782-1"},{"key":"e_1_2_10_30_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1215037109"},{"key":"e_1_2_10_31_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.ge.21.120187.000403"},{"key":"e_1_2_10_32_1","doi-asserted-by":"publisher","DOI":"10.1016\/bs.mie.2017.01.019"},{"key":"e_1_2_10_33_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajpcell.00447.2007"},{"key":"e_1_2_10_34_1","doi-asserted-by":"publisher","DOI":"10.1002\/bies.201900115"},{"key":"e_1_2_10_35_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-020-58820-y"},{"key":"e_1_2_10_36_1","unstructured":"Watson R. C. L.Buckley R.Mills andA.Davies.2010.\u201cAssociative Memory in Gene Regulation Networks.\u201dIn Proceedings of the 12th International Conference on the Synthesis and Simulation of Living Systems (ALIFE 12) 194\u2013201. 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