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Here we assess this by measuring single neuron activity (1,420 neurons) and local field potentials (921 channels) in the medial temporal lobe of 16 patients with epilepsy performing a working-memory task for temporal order. During memory maintenance, we observe theta oscillations, preferential firing of single neurons to theta phase and a close relationship between phase of firing and item position. However, the firing order did not match item order. Training recurrent neural networks to perform an analogous task, we also show the generation of theta oscillations, theta phase-dependent firing related to item position and, again, no match between firing and item order. Rather, our results suggest a mechanistic link between phase order, stimulus timing and oscillation frequency. In both biological and artificial neural networks, we provide evidence supporting the role of phase of firing in working-memory processing.<\/jats:p>","DOI":"10.1038\/s41593-025-01893-7","type":"journal-article","created":{"date-parts":[[2025,3,24]],"date-time":"2025-03-24T06:19:05Z","timestamp":1742797145000},"page":"873-882","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":26,"title":["Phase of firing does not reflect temporal order in sequence memory of humans and recurrent neural networks"],"prefix":"10.1038","volume":"28","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2873-2943","authenticated-orcid":false,"given":"Stefanie","family":"Liebe","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3323-2986","authenticated-orcid":false,"given":"Johannes","family":"Niediek","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3051-1325","authenticated-orcid":false,"given":"Matthijs","family":"Pals","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3969-9782","authenticated-orcid":false,"given":"Thomas P.","family":"Reber","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jennifer","family":"Faber","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jan","family":"Bostr\u00f6m","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2531-6701","authenticated-orcid":false,"given":"Christian E.","family":"Elger","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jakob H.","family":"Macke","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1305-8028","authenticated-orcid":false,"given":"Florian","family":"Mormann","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2025,3,24]]},"reference":[{"key":"1893_CR1","doi-asserted-by":"publisher","first-page":"1026","DOI":"10.1016\/j.cub.2017.02.013","volume":"27","author":"S Kornblith","year":"2017","unstructured":"Kornblith, S., Quian Quiroga, R., Koch, C., Fried, I. & Mormann, F. 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Subjects in our study were selected based on medical criteria and voluntary participation was ensured for each experiment. Selection of participants could not result in stigmatization, incrimination or discrimination. Participants\u2019 safety was guaranteed by presence of medical personnel throughout continuous monitoring for the entire duration of the hospital stay. We have carefully considered research contributions and authorship criteria when involved in multiparty collaborations with the goal of promoting greater equity in research collaborations.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics statement"}}]}}