{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T16:07:13Z","timestamp":1790006833108,"version":"4.0.1"},"reference-count":94,"publisher":"Oxford University Press (OUP)","license":[{"start":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T00:00:00Z","timestamp":1787702400000},"content-version":"am","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"publisher","award":["PJT 166167"],"award-info":[{"award-number":["PJT 166167"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"publisher","award":["MOP 142191"],"award-info":[{"award-number":["MOP 142191"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"publisher","award":["PJT 153115"],"award-info":[{"award-number":["PJT 153115"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000024","name":"Canadian Institutes of Health Research","doi-asserted-by":"publisher","award":["PJT 156125"],"award-info":[{"award-number":["PJT 156125"]}],"id":[{"id":"10.13039\/501100000024","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000196","name":"Canada Foundation for Innovation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000196","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000156","name":"Fonds de Recherche du Qu\u00e9bec \u2013 Sant\u00e9","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000156","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100019889","name":"Weston Family Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100019889","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100002600","name":"American Sleep Medicine Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100002600","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002914","name":"Concordia University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100002914","id-type":"DOI","asserted-by":"publisher"}]},{"name":"CIHR fellowship","award":["FRN: MFE -195237"],"award-info":[{"award-number":["FRN: MFE -195237"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Insufficient sleep impairs cognition, particularly hippocampal-dependent memory. However, effective strategies for addressing these effects in humans remain limited. Physical exercise enhances slow-wave sleep (SWS) and improves memory performance, suggesting it may provide a physiologically grounded buffer against sleep-loss\u2013related memory deficits. Here, we tested whether a single session of moderate-intensity exercise may protect long-term declarative memory following sleep restriction, and whether SWS mediates this effect. Eighty-eight healthy young adults (52% female; 24.0\u2009\u00b1\u20093.8\u00a0years) were randomly assigned to one of four groups based on exercise and sleep opportunity: average sleep (S8: 8.5\u00a0hours\/night), sleep restriction (S5: 4.5\u00a0hours\/night), exercise before average sleep (ExS8), and exercise before sleep restriction (ExS5). Participants completed either a 30-min evening moderate-intensity cycling (7: 00\u00a0p.m.; 55%\u201360% VO\u2082max) or seated rest prior to learning 80 face\u2013name pairs (10:00\u00a0p.m.), followed by immediate and delayed recall (120 face-name pairs: 80 studied, 40 novel) after polysomnography-recorded sleep. We found a significant exercise \u00d7 sleep interaction for delayed recall performance (discriminability index, d\u2032; p\u2009=\u2009.033, partial \u03b72\u2009=\u20090.056). The delayed recall d\u2032 of the ExS5 was significantly higher than the S5 (p\u2009=\u2009.028), but was comparable to S8 and ExS8 groups (ps\u2009&amp;gt;\u20090.05). The proportion of SWS mediated the association between exercise and delayed recall d\u2032 (B\u2009=\u20090.105, SE\u2009=\u20090.091, 95% bootstrap CI (0.073, 0.924)). These findings indicate that acute moderate-intensity exercise is associated with offsetting the negative impact of sleep restriction on long-term declarative memory in sleep restriction, an effect statistically mediated by SWS percentage.<\/jats:p>","DOI":"10.1093\/sleep\/zsag235","type":"journal-article","created":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T11:42:18Z","timestamp":1787658138000},"source":"Crossref","is-referenced-by-count":0,"title":["Slow-wave sleep mediates the compensatory effects of acute moderate-intensity exercise on declarative memory deficits in sleep restriction"],"prefix":"10.1093","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5238-9236","authenticated-orcid":false,"given":"Emmanuel","family":"Frimpong","sequence":"first","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Health, Kinesiology & Applied Physiology, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Centre de Recherche de l\u2019Institut Universitaire de G\u00e9riatrie de Montr\u00e9al , Sant\u00e9 Qu\u00e9bec Centre-Sud-de-l\u2019\u00eele-de-Montr\u00e9al Universitaire, Montr\u00e9al, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Melodee","family":"Mograss","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Health, Kinesiology & Applied Physiology, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Psychology, Concordia University , Montreal, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jocelyn","family":"Dautet-Plourde","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Health, Kinesiology & Applied Physiology, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nicole","family":"Milbrath","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Psychology, Concordia University , Montreal, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francois","family":"Pointeau","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Health, Kinesiology & Applied Physiology, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Centre de Recherche de l\u2019Institut Universitaire de G\u00e9riatrie de Montr\u00e9al , Sant\u00e9 Qu\u00e9bec Centre-Sud-de-l\u2019\u00eele-de-Montr\u00e9al Universitaire, Montr\u00e9al, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Manlu","family":"Xu","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Psychology, Concordia University , Montreal, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ngoc Long Khang","family":"Nguyen","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Psychology, Concordia University , Montreal, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tehila","family":"Zvionow","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Health, Kinesiology & Applied Physiology, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6796-6710","authenticated-orcid":false,"given":"Mylene","family":"Aubertin-Leheudre","sequence":"additional","affiliation":[{"name":"Centre de Recherche de l\u2019Institut Universitaire de G\u00e9riatrie de Montr\u00e9al , Sant\u00e9 Qu\u00e9bec Centre-Sud-de-l\u2019\u00eele-de-Montr\u00e9al Universitaire, Montr\u00e9al, QC ,","place":["Canada"]},{"name":"D\u00e9partement des Sciences de l\u2019activit\u00e9 physique, GRAPA , , Montr\u00e9al, QC ,","place":["Canada"]},{"name":"Universit\u00e9 du Qu\u00e9bec \u00e0 Montr\u00e9al , , Montr\u00e9al, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Louis","family":"Bherer","sequence":"additional","affiliation":[{"name":"Centre de Recherche de l\u2019Institut Universitaire de G\u00e9riatrie de Montr\u00e9al , Sant\u00e9 Qu\u00e9bec Centre-Sud-de-l\u2019\u00eele-de-Montr\u00e9al Universitaire, Montr\u00e9al, QC ,","place":["Canada"]},{"name":"Department of Medicine and Centre de recherche de l\u2019Institut de cardiologie de Montr\u00e9al , , Montr\u00e9al, QC ,","place":["Canada"]},{"name":"Universit\u00e9 de Montr\u00e9al , , Montr\u00e9al, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"V\u00e9ronique","family":"Pepin","sequence":"additional","affiliation":[{"name":"Department of Health, Kinesiology & Applied Physiology, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Centre de recherche , , Montr\u00e9al, QC ,","place":["Canada"]},{"name":"CIUSSS du Nord-de l\u2019\u00cele-de-Montr\u00e9al , , Montr\u00e9al, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Edwin","family":"Robertson","sequence":"additional","affiliation":[{"name":"School of Psychology and Neuroscience, University of Glasgow , Glasgow ,","place":["United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7984-514X","authenticated-orcid":false,"given":"Arsenio","family":"P\u00e1ez","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Health, Kinesiology & Applied Physiology, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Centre de Recherche de l\u2019Institut Universitaire de G\u00e9riatrie de Montr\u00e9al , Sant\u00e9 Qu\u00e9bec Centre-Sud-de-l\u2019\u00eele-de-Montr\u00e9al Universitaire, Montr\u00e9al, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thien Thanh","family":"Dang-Vu","sequence":"additional","affiliation":[{"name":"Sleep, Cognition & Neuroimaging Laboratory, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Department of Health, Kinesiology & Applied Physiology, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"School of Health, Concordia University , Montreal, QC ,","place":["Canada"]},{"name":"Centre de Recherche de l\u2019Institut Universitaire de G\u00e9riatrie de Montr\u00e9al , Sant\u00e9 Qu\u00e9bec Centre-Sud-de-l\u2019\u00eele-de-Montr\u00e9al Universitaire, Montr\u00e9al, QC ,","place":["Canada"]},{"name":"Department of Psychology, Concordia University , Montreal, QC ,","place":["Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,8,26]]},"reference":[{"issue":"2","key":"2026092111370635900_ref1","doi-asserted-by":"publisher","first-page":"681","DOI":"10.1152\/physrev.00032.2012","article-title":"About sleep\u2019s role in memory","volume":"93","author":"Rasch","year":"2013","journal-title":"Physiol Rev"},{"issue":"10","key":"2026092111370635900_ref2","doi-asserted-by":"publisher","first-page":"558","DOI":"10.1101\/lm.031674.113","article-title":"The impact of sleep loss on hippocampal function","volume":"20","author":"Prince","year":"2013","journal-title":"Learn Mem"},{"key":"2026092111370635900_ref3","doi-asserted-by":"publisher","first-page":"108661","DOI":"10.1016\/j.neuropsychologia.2023.108661","article-title":"Spindle-dependent memory consolidation in healthy adults: a meta-analysis","volume":"189","author":"Kumral","year":"2023","journal-title":"Neuropsychologia."},{"issue":"2","key":"2026092111370635900_ref4","doi-asserted-by":"publisher","first-page":"430","DOI":"10.1007\/s11910-013-0430-8","article-title":"Differential effects of non-REM and REM sleep on memory consolidation?","volume":"14","author":"Ackermann","year":"2014","journal-title":"Curr Neurol Neurosci Rep"},{"issue":"10","key":"2026092111370635900_ref5","doi-asserted-by":"publisher","first-page":"1598","DOI":"10.1038\/s41593-019-0467-3","article-title":"Mechanisms of systems memory consolidation during sleep","volume":"22","author":"Klinzing","year":"2019","journal-title":"Nat Neurosci"},{"issue":"3","key":"2026092111370635900_ref6","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1038\/nrn2762-c2","article-title":"Slow-wave sleep takes the leading role in memory reorganization","volume":"11","author":"Diekelmann","year":"2010","journal-title":"Nat Rev Neurosci"},{"issue":"10","key":"2026092111370635900_ref7","doi-asserted-by":"publisher","first-page":"4494","DOI":"10.1523\/JNEUROSCI.3127-12.2013","article-title":"The critical role of sleep spindles in hippocampal-dependent memory: a pharmacology study","volume":"33","author":"Mednick","year":"2013","journal-title":"J Neurosci"},{"issue":"3","key":"2026092111370635900_ref8","doi-asserted-by":"publisher","first-page":"zsx003","DOI":"10.1093\/sleep\/zsx003","article-title":"Enhanced memory consolidation via automatic sound stimulation during non-REM sleep","volume":"40","author":"Leminen","year":"2017","journal-title":"Sleep."},{"issue":"33","key":"2026092111370635900_ref9","doi-asserted-by":"publisher","first-page":"7314","DOI":"10.1523\/JNEUROSCI.0273-18.2018","article-title":"Closed-loop slow-wave tACS improves sleep-dependent long-term memory generalization by modulating endogenous oscillations","volume":"38","author":"Ketz","year":"2018","journal-title":"J Neurosci"},{"issue":"5","key":"2026092111370635900_ref10","doi-asserted-by":"publisher","first-page":"300","DOI":"10.3390\/brainsci10050300","article-title":"Boosting slow oscillations during sleep to improve memory function in elderly people: a review of the literature","volume":"10","author":"Salfi","year":"2020","journal-title":"Brain Sci"},{"key":"2026092111370635900_ref11","doi-asserted-by":"publisher","first-page":"646678","DOI":"10.3389\/fnins.2021.646678","article-title":"Sleep and memory consolidation dysfunction in psychiatric disorders: evidence for the involvement of extracellular matrix molecules","volume":"15","author":"Gisabella","year":"2021","journal-title":"Front Neurosci"},{"issue":"4","key":"2026092111370635900_ref12","doi-asserted-by":"publisher","first-page":"440","DOI":"10.1037\/bul0000410","article-title":"Sleep loss and emotion: a systematic review and meta-analysis of over 50 years of experimental research","volume":"150","author":"Palmer","year":"2024","journal-title":"Psychol Bull"},{"key":"2026092111370635900_ref13","doi-asserted-by":"publisher","first-page":"586","DOI":"10.1016\/j.neubiorev.2017.07.010","article-title":"The neurocognitive consequences of sleep restriction: a meta-analytic review","volume":"80","author":"Lowe","year":"2017","journal-title":"Neurosci Biobehav Rev"},{"issue":"1","key":"2026092111370635900_ref14","doi-asserted-by":"publisher","first-page":"8774","DOI":"10.1038\/s41598-020-65086-x","article-title":"Two nights of recovery sleep restores hippocampal connectivity but not episodic memory after total sleep deprivation","volume":"10","author":"Chai","year":"2020","journal-title":"Sci Rep"},{"key":"2026092111370635900_ref15","doi-asserted-by":"publisher","first-page":"64","DOI":"10.1016\/j.smrv.2019.03.008","article-title":"Functional brain alterations in acute sleep deprivation: an activation likelihood estimation meta-analysis","volume":"46","author":"Javaheripour","year":"2019","journal-title":"Sleep Med Rev"},{"issue":"4","key":"2026092111370635900_ref16","doi-asserted-by":"publisher","first-page":"534","DOI":"10.1162\/jocn.1997.9.4.534","article-title":"Effects of early and late nocturnal sleep on declarative and procedural memory","volume":"9","author":"Plihal","year":"1997","journal-title":"J Cogn Neurosci"},{"key":"2026092111370635900_ref17","doi-asserted-by":"publisher","first-page":"105929","DOI":"10.1016\/j.neubiorev.2024.105929","article-title":"A systematic and meta-analytic review of the impact of sleep restriction on memory formation","volume":"167","author":"Crowley","year":"2024","journal-title":"Neurosci Biobehav Rev."},{"issue":"1","key":"2026092111370635900_ref18","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3390\/healthcare7010001","article-title":"The global problem of insufficient sleep and its serious public health implications","volume":"7","author":"Chattu","year":"2018","journal-title":"Healthcare (Basel)"},{"key":"2026092111370635900_ref19","doi-asserted-by":"publisher","first-page":"13","DOI":"10.1007\/978-3-319-41400-3_2","volume-title":"Clinical Handbook of Insomnia. Current Clinical Neurology","author":"Grewal","year":"2017"},{"issue":"1","key":"2026092111370635900_ref20","doi-asserted-by":"publisher","first-page":"38","DOI":"10.1249\/JES.0000000000000273","article-title":"Exercising the sleepying brain: exercise, sleep, and sleep loss on memory","volume":"50","author":"Roig","year":"2022","journal-title":"Exerc Sport Sci Rev"},{"key":"2026092111370635900_ref21","doi-asserted-by":"publisher","DOI":"10.3389\/fneur.2012.00048","article-title":"Exercise effects on sleep physiology","volume":"3","author":"Uchida","year":"2012","journal-title":"Front Neur"},{"issue":"3","key":"2026092111370635900_ref22","doi-asserted-by":"publisher","first-page":"266","DOI":"10.1016\/j.sleep.2007.04.017","article-title":"Increased slow wave sleep and reduced stage 2 sleep in children depending on exercise intensity","volume":"9","author":"Dworak","year":"2008","journal-title":"Sleep Med"},{"issue":"3","key":"2026092111370635900_ref23","doi-asserted-by":"publisher","first-page":"427","DOI":"10.1007\/s10865-015-9617-6","article-title":"The effects of physical activity on sleep: a meta-analytic review","volume":"38","author":"Kredlow","year":"2015","journal-title":"J Behav Med"},{"issue":"2","key":"2026092111370635900_ref24","doi-asserted-by":"publisher","first-page":"269","DOI":"10.1007\/s40279-018-1015-0","article-title":"Effects of evening exercise on sleep in healthy participants: a systematic review and meta-analysis","volume":"49","author":"Stutz","year":"2019","journal-title":"Sports Med"},{"issue":"1","key":"2026092111370635900_ref25","doi-asserted-by":"publisher","first-page":"168","DOI":"10.1152\/japplphysiol.00765.2018","article-title":"Diurnal repeated exercise promotes slow-wave activity and fast-sigma power during sleep with increase in body temperature: a human crossover trial","volume":"127","author":"Aritake-Okada","year":"2019","journal-title":"J Appl Physiol"},{"issue":"4","key":"2026092111370635900_ref26","doi-asserted-by":"publisher","first-page":"87","DOI":"10.3390\/brainsci9040087","article-title":"The temporal effects of acute exercise on episodic memory function: systematic review with meta-analysis","volume":"9","author":"Loprinzi","year":"2019","journal-title":"Brain Sci"},{"issue":"2","key":"2026092111370635900_ref27","doi-asserted-by":"publisher","first-page":"81","DOI":"10.1249\/JES.0000000000000078","article-title":"Time-dependent effects of cardiovascular exercise on memory","volume":"44","author":"Roig","year":"2016","journal-title":"Exerc Sport Sci Rev"},{"issue":"8","key":"2026092111370635900_ref28","doi-asserted-by":"publisher","first-page":"1645","DOI":"10.1016\/j.neubiorev.2013.06.012","article-title":"The effects of cardiovascular exercise on human memory: a review with meta-analysis","volume":"37","author":"Roig","year":"2013","journal-title":"Neurosci Biobehav Rev."},{"issue":"12","key":"2026092111370635900_ref29","doi-asserted-by":"publisher","first-page":"973","DOI":"10.1136\/bjsports-2012-091441","article-title":"Physical exercise and executive functions in preadolescent children, adolescents and young adults: a meta-analysis","volume":"48","author":"Verburgh","year":"2014","journal-title":"Br J Sports Med"},{"issue":"1","key":"2026092111370635900_ref30","doi-asserted-by":"publisher","first-page":"3112","DOI":"10.1038\/s41467-021-23520-2","article-title":"Endogenous memory reactivation during sleep in humans is clocked by slow oscillation-spindle complexes","volume":"12","author":"Schreiner","year":"2021","journal-title":"Nat Commun"},{"issue":"2","key":"2026092111370635900_ref31","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1007\/s40675-024-00291-y","article-title":"Electrophysiological mechanisms of memory consolidation in human non-rapid eye movement sleep","volume":"10","author":"Denis","year":"2024","journal-title":"Curr Sleep Med Rep"},{"issue":"44","key":"2026092111370635900_ref32","doi-asserted-by":"publisher","first-page":"e2123428119","DOI":"10.1073\/pnas.2123428119","article-title":"Shaping overnight consolidation via slow-oscillation closed-loop targeted memory reactivation","volume":"119","author":"Ngo","year":"2022","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"8","key":"2026092111370635900_ref33","doi-asserted-by":"publisher","first-page":"1429","DOI":"10.1038\/s41593-023-01381-w","article-title":"How coupled slow oscillations, spindles and ripples coordinate neuronal processing and communication during human sleep","volume":"26","author":"Staresina","year":"2023","journal-title":"Nat Neurosci"},{"issue":"7","key":"2026092111370635900_ref34","doi-asserted-by":"publisher","first-page":"470","DOI":"10.1016\/j.tins.2018.03.003","article-title":"Slow-wave activity enhancement to improve cognition","volume":"41","author":"Wilckens","year":"2018","journal-title":"Trends Neurosci"},{"issue":"5","key":"2026092111370635900_ref35","doi-asserted-by":"publisher","first-page":"1154","DOI":"10.1016\/j.neubiorev.2010.12.003","article-title":"The function of the sleep spindle: a physiological index of intelligence and a mechanism for sleep-dependent memory consolidation","volume":"35","author":"Fogel","year":"2011","journal-title":"Neurosci Biobehav Rev"},{"issue":"4","key":"2026092111370635900_ref36","doi-asserted-by":"publisher","first-page":"zsad017","DOI":"10.1093\/sleep\/zsad017","article-title":"Sleep spindles in the healthy brain from birth through 18 years","volume":"46","author":"Kwon","year":"2023","journal-title":"Sleep."},{"issue":"2","key":"2026092111370635900_ref37","doi-asserted-by":"publisher","first-page":"805","DOI":"10.1152\/physrev.00042.2018","article-title":"Sleep spindles: mechanisms and functions","volume":"100","author":"Fernandez","year":"2020","journal-title":"Physiol Rev"},{"key":"2026092111370635900_ref38","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2025.121105","article-title":"Multi-task exercise increases delta power in non-rapid eye movement sleep among older female adults: a randomized crossover trial","volume":"310","author":"Lee","year":"2025","journal-title":"NeuroImage."},{"key":"2026092111370635900_ref39","doi-asserted-by":"publisher","first-page":"4410","DOI":"10.1038\/s41598-021-83817-6","article-title":"Exercise improves the quality of slow-wave sleep by increasing slow-wave stability","volume":"11","author":"Park","year":"2021","journal-title":"Sci Rep"},{"key":"2026092111370635900_ref40","doi-asserted-by":"publisher","first-page":"14371","DOI":"10.1038\/s41598-021-93813-5","article-title":"A single session of moderate intensity exercise influences memory, endocannabinoids and brain derived neurotrophic factor levels in men","volume":"11","author":"Marin Bosch","year":"2021","journal-title":"Sci Rep"},{"issue":"6","key":"2026092111370635900_ref41","doi-asserted-by":"publisher","first-page":"336","DOI":"10.1123\/jsep.2018-0072","article-title":"The effect of acute exercise on encoding and consolidation of long-term memory","volume":"40","author":"Labban","year":"2018","journal-title":"J Sport Exerc Psychol"},{"issue":"1","key":"2026092111370635900_ref42","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.jshs.2019.07.012","article-title":"Updated overview on interplay between physical exercise, neurotrophins, and cognitive function in humans","volume":"9","author":"Lippi","year":"2020","journal-title":"J Sport Health Sci"},{"issue":"5","key":"2026092111370635900_ref43","doi-asserted-by":"publisher","first-page":"536","DOI":"10.1016\/j.semcdb.2011.07.002","article-title":"Effects of environmental enrichment and voluntary exercise on neurogenesis, learning and memory, and pattern separation: BDNF as a critical variable?","volume":"22","author":"Bekinschtein","year":"2011","journal-title":"Semin Cell Dev Biol"},{"issue":"6","key":"2026092111370635900_ref44","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1016\/S0166-2236(02)02143-4","article-title":"Exercise: a behavioral intervention to enhance brain health and plasticity","volume":"25","author":"Cotman","year":"2002","journal-title":"Trends Neurosci"},{"issue":"1","key":"2026092111370635900_ref45","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-025-07880-z","article-title":"High intensity exercise before sleep boosts memory encoding the next morning","volume":"15","author":"Ramirez","year":"2025","journal-title":"Sci Rep"},{"issue":"4","key":"2026092111370635900_ref46","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1556\/2060.105.2018.4.28","article-title":"Hypothesized mechanisms through which acute exercise influences episodic memory","volume":"105","author":"Loprinzi","year":"2018","journal-title":"Physiol Int"},{"issue":"6","key":"2026092111370635900_ref47","doi-asserted-by":"publisher","first-page":"375","DOI":"10.1177\/1559827614544437","article-title":"The bidirectional relationship between exercise and sleep: implications for exercise adherence and sleep improvement","volume":"8","author":"Kline","year":"2014","journal-title":"Am J Lifestyle Med"},{"issue":"7","key":"2026092111370635900_ref48","doi-asserted-by":"publisher","first-page":"zsad119","DOI":"10.1093\/sleep\/zsad119","article-title":"Acute evening high-intensity interval training may attenuate the detrimental effects of sleep restriction on long-term declarative memory","volume":"46","author":"Frimpong","year":"2023","journal-title":"Sleep."},{"key":"2026092111370635900_ref49","doi-asserted-by":"publisher","first-page":"793875","DOI":"10.3389\/fpsyg.2022.793875","article-title":"Cardiorespiratory fitness may protect memory for poorer sleepers","volume":"13","author":"Kuhn","year":"2022","journal-title":"Front Psychol"},{"issue":"9","key":"2026092111370635900_ref50","doi-asserted-by":"publisher","DOI":"10.1093\/sleep\/zsaa062","article-title":"Exercising before a nap benefits memory better than napping or exercising alone","volume":"43","author":"Mograss","year":"2020","journal-title":"Sleep."},{"issue":"6","key":"2026092111370635900_ref51","doi-asserted-by":"publisher","first-page":"569","DOI":"10.1080\/15402002.2016.1253013","article-title":"Physical activity and cognition: a mediating role of efficient sleep","volume":"16","author":"Wilckens","year":"2018","journal-title":"Behav Sleep Med"},{"issue":"9","key":"2026092111370635900_ref52","doi-asserted-by":"publisher","first-page":"1170","DOI":"10.1093\/gerona\/glu038","article-title":"Sleep and executive function in older women: the moderating effect of physical activity","volume":"69","author":"Lambiase","year":"2014","journal-title":"J Gerontol A Biol Sci Med Sci"},{"key":"2026092111370635900_ref53","doi-asserted-by":"publisher","first-page":"115753","DOI":"10.1016\/j.bbr.2025.115753","article-title":"Physical exercise prevents behavioral and neurobiological deficits induced by sleep deprivation in rodents through the regulation of BDNF and 5-HT levels: a systematic review and meta-analysis","volume":"495","author":"De Matos","year":"2025","journal-title":"Behav Brain Res"},{"issue":"4","key":"2026092111370635900_ref54","doi-asserted-by":"publisher","first-page":"1702","DOI":"10.1016\/j.neuroimage.2009.12.023","article-title":"Acute moderate exercise elicits increased dorsolateral prefrontal activation and improves cognitive performance with Stroop test","volume":"50","author":"Yanagisawa","year":"2010","journal-title":"NeuroImage."},{"issue":"8","key":"2026092111370635900_ref55","doi-asserted-by":"publisher","first-page":"1381","DOI":"10.1249\/01.MSS.0000078924.61453.FB","article-title":"International physical activity questionnaire: 12-country reliability and validity","volume":"35","author":"Craig","year":"2003","journal-title":"Med Sci Sports Exerc"},{"issue":"2","key":"2026092111370635900_ref56","doi-asserted-by":"publisher","first-page":"355","DOI":"10.1038\/s41366-018-0138-6","article-title":"The impact of obesity in cognitive and memory dysfunction in obstructive sleep apnea syndrome","volume":"43","author":"Shen","year":"2019","journal-title":"Int J Obes (Lond)"},{"issue":"2","key":"2026092111370635900_ref57","doi-asserted-by":"publisher","first-page":"193","DOI":"10.1016\/0165-1781(89)90047-4","article-title":"The Pittsburgh Sleep Quality Index: a new instrument for psychiatric practice and research","volume":"28","author":"Buysse","year":"1989","journal-title":"Psychiatry Res"},{"issue":"6","key":"2026092111370635900_ref58","doi-asserted-by":"publisher","first-page":"540","DOI":"10.1093\/sleep\/14.6.540","article-title":"A new method for measuring daytime sleepiness: the Epworth sleepiness scale","volume":"14","author":"Johns","year":"1991","journal-title":"Sleep."},{"issue":"5","key":"2026092111370635900_ref59","doi-asserted-by":"publisher","first-page":"601","DOI":"10.1093\/sleep\/34.5.601TeX","article-title":"The Insomnia Severity Index: psychometric indicators to detect insomnia cases and evaluate treatment response","volume":"34","author":"Morin","year":"2011","journal-title":"Sleep."},{"issue":"1","key":"2026092111370635900_ref60","doi-asserted-by":"publisher","first-page":"52","DOI":"10.1111\/j.1365-2869.1994.tb00104.x","article-title":"The Ullanlinna Narcolepsy Scale: validation of a measure of symptoms in the narcoleptic syndrome","volume":"3","author":"Hublin","year":"1994","journal-title":"J Sleep Res"},{"key":"2026092111370635900_ref61","doi-asserted-by":"publisher","volume-title":"J Pers Assess","DOI":"10.1207\/s15327752jpa6703_13"},{"issue":"6","key":"2026092111370635900_ref62","doi-asserted-by":"publisher","first-page":"893","DOI":"10.1037\/\/0022-006x.56.6.893","article-title":"An inventory for measuring clinical anxiety: psychometric properties","volume":"56","author":"Beck","year":"1988","journal-title":"J Consult Clin Psychol"},{"issue":"2","key":"2026092111370635900_ref63","first-page":"97","article-title":"A self-assessment questionnaire to determine morningness-eveningness in human circadian rhythms","volume":"4","author":"Horne","year":"1976","journal-title":"Int J Chronobiol"},{"issue":"2","key":"2026092111370635900_ref64","doi-asserted-by":"publisher","first-page":"175","DOI":"10.3758\/BF03193146","article-title":"G*Power 3: a flexible statistical power analysis program for the social, behavioral, and biomedical sciences","volume":"39","author":"Faul","year":"2007","journal-title":"Behav Res Methods"},{"issue":"5","key":"2026092111370635900_ref65","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1249\/00005768-198205000-00012","article-title":"Psychophysical bases of perceived exertion","volume":"14","author":"Borg","year":"1982","journal-title":"Med Sci Sports Exerc"},{"key":"2026092111370635900_ref66","volume-title":"ACSM\u2019s Resource Manual for Guidelines for Exercise Testing and Prescription","author":"Guthrie","year":"2013","edition":"6"},{"key":"2026092111370635900_ref67","doi-asserted-by":"publisher","first-page":"101535","DOI":"10.1016\/j.smrv.2021.101535","article-title":"The effects of evening high-intensity exercise on sleep in healthy adults: a systematic review and meta-analysis","volume":"60","author":"Frimpong","year":"2021","journal-title":"Sleep Med Rev"},{"issue":"9","key":"2026092111370635900_ref68","doi-asserted-by":"publisher","first-page":"1789","DOI":"10.1007\/s00421-014-2873-2","article-title":"Sleep onset is disrupted following pre-sleep exercise that causes large physiological excitement at bedtime","volume":"114","author":"Oda","year":"2014","journal-title":"Eur J Appl Physiol"},{"issue":"12","key":"2026092111370635900_ref69","doi-asserted-by":"publisher","first-page":"3301","DOI":"10.1093\/cercor\/bht188","article-title":"Impaired prefrontal sleep spindle regulation of hippocampal-dependent learning in older adults","volume":"24","author":"Mander","year":"2014","journal-title":"Cereb Cortex"},{"issue":"1","key":"2026092111370635900_ref70","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1037\/0096-3445.117.1.34","article-title":"Pragmatics of measuring recognition memory: applications to dementia and amnesia","volume":"117","author":"Snodgrass","year":"1988","journal-title":"J Exp Psychol Gen"},{"issue":"1","key":"2026092111370635900_ref71","doi-asserted-by":"publisher","first-page":"11579","DOI":"10.1038\/s41598-018-30030-7","article-title":"The discrimination ratio derived from novel object recognition tasks as a measure of recognition memory sensitivity, not bias","volume":"8","author":"Sivakumaran","year":"2018","journal-title":"Sci Rep"},{"key":"2026092111370635900_ref72"},{"issue":"4","key":"2026092111370635900_ref73","doi-asserted-by":"publisher","first-page":"277","DOI":"10.2165\/00007256-199621040-00004","article-title":"The effects of acute and chronic exercise on sleep: a meta-analytic review","volume":"21","author":"Kubitz","year":"1996","journal-title":"Sports Med"},{"key":"2026092111370635900_ref74","volume-title":"Introduction to Mediation, Moderation, and Conditional Process Analysis: A Regression-Based Approach","author":"Hayes","year":"2013"},{"issue":"4","key":"2026092111370635900_ref75","doi-asserted-by":"publisher","first-page":"155","DOI":"10.2114\/jpa2.28.155","article-title":"Positive effects of acute and moderate physical exercise on cognitive function","volume":"28","author":"Kashihara","year":"2009","journal-title":"J Physiol Anthropol"},{"key":"2026092111370635900_ref76","doi-asserted-by":"publisher","first-page":"87","DOI":"10.1016\/j.brainres.2012.02.068","article-title":"The effects of acute exercise on cognitive performance: a meta-analysis","volume":"1453","author":"Chang","year":"2012","journal-title":"Brain Res"},{"issue":"9","key":"2026092111370635900_ref77","doi-asserted-by":"publisher","first-page":"192","DOI":"10.1101\/lm.053683.122","article-title":"Targeted memory reactivation during slow-wave sleep vs. sleep stage N2: no significant differences in a vocabulary task","volume":"30","author":"Wick","year":"2023","journal-title":"Learn Mem"},{"issue":"1","key":"2026092111370635900_ref78","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/j.neuron.2017.02.004","article-title":"Sleep and human aging","volume":"94","author":"Mander","year":"2017","journal-title":"Neuron."},{"issue":"1","key":"2026092111370635900_ref79","doi-asserted-by":"publisher","first-page":"12","DOI":"10.1016\/j.neuron.2013.12.025","article-title":"Sleep and the price of plasticity: from synaptic and cellular homeostasis to memory consolidation and integration","volume":"81","author":"Tononi","year":"2014","journal-title":"Neuron."},{"key":"2026092111370635900_ref80","doi-asserted-by":"publisher","first-page":"441","DOI":"10.1007\/164_2019_243","volume-title":"Sleep-Wake Neurobiology and Pharmacology","author":"Dijk","year":"2019"},{"issue":"11","key":"2026092111370635900_ref81","doi-asserted-by":"publisher","first-page":"1679","DOI":"10.1038\/nn.4119","article-title":"Hierarchical nesting of slow oscillations, spindles and ripples in the human hippocampus during sleep","volume":"18","author":"Staresina","year":"2015","journal-title":"Nat Neurosci"},{"issue":"1","key":"2026092111370635900_ref82","doi-asserted-by":"publisher","first-page":"49","DOI":"10.1016\/j.smrv.2005.05.002","article-title":"Sleep function and synaptic homeostasis","volume":"10","author":"Tononi","year":"2006","journal-title":"Sleep Med Rev"},{"issue":"1-2","key":"2026092111370635900_ref83","doi-asserted-by":"publisher","first-page":"157","DOI":"10.1016\/0166-4328(95)00012-I","article-title":"The sequential hypothesis of the function of sleep","volume":"69","author":"Giuditta","year":"1995","journal-title":"Behav Brain Res"},{"issue":"41","key":"2026092111370635900_ref84","doi-asserted-by":"publisher","first-page":"10487","DOI":"10.1073\/pnas.1805668115","article-title":"Rapid stimulation of human dentate gyrus function with acute mild exercise","volume":"115","author":"Suwabe","year":"2018","journal-title":"Proc Natl Acad Sci U S A"},{"issue":"8","key":"2026092111370635900_ref85","doi-asserted-by":"publisher","first-page":"800","DOI":"10.1111\/gbb.12394","article-title":"A single bout of exercise increases hippocampal Bdnf: influence of chronic exercise and noradrenaline","volume":"16","author":"Venezia","year":"2017","journal-title":"Genes Brain Behav"},{"key":"2026092111370635900_ref86","doi-asserted-by":"publisher","first-page":"137830","DOI":"10.1016\/j.neulet.2024.137830","article-title":"In vivo bioluminescence imaging revealed the change of the time window of BDNF expression in the brain elicited by a single bout of exercise following repeated exercise","volume":"834","author":"Ikegami","year":"2024","journal-title":"Neurosci Lett"},{"issue":"5","key":"2026092111370635900_ref87","doi-asserted-by":"publisher","first-page":"339","DOI":"10.1038\/nrn2822","article-title":"New neurons and new memories: how does adult hippocampal neurogenesis affect learning and memory?","volume":"11","author":"Deng","year":"2010","journal-title":"Nat Rev Neurosci"},{"issue":"1","key":"2026092111370635900_ref88","doi-asserted-by":"publisher","first-page":"a018903","DOI":"10.1101\/cshperspect.a018903","article-title":"Maturation and functional integration of new granule cells into the adult hippocampus","volume":"8","author":"Toni","year":"2016","journal-title":"Cold Spring Harb Perspect Biol"},{"issue":"3","key":"2026092111370635900_ref89","doi-asserted-by":"publisher","first-page":"260","DOI":"10.1038\/6365","article-title":"Learning enhances adult neurogenesis in the hippocampal formation","volume":"2","author":"Gould","year":"1999","journal-title":"Nat Neurosci"},{"issue":"3","key":"2026092111370635900_ref90","doi-asserted-by":"publisher","first-page":"302","DOI":"10.1093\/sleep\/32.3.302","article-title":"Slow wave sleep and REM sleep awakenings do not affect sleep dependent memory consolidation","volume":"32","author":"Genzel","year":"2009","journal-title":"Sleep."},{"issue":"1","key":"2026092111370635900_ref91","doi-asserted-by":"publisher","first-page":"9057","DOI":"10.1038\/s41598-024-59696-y","article-title":"Comparing targeted memory reactivation during slow wave sleep and sleep stage 2","volume":"14","author":"Carbone","year":"2024","journal-title":"Sci Rep"},{"issue":"10","key":"2026092111370635900_ref92","doi-asserted-by":"publisher","first-page":"2389","DOI":"10.1016\/j.neuropsychologia.2012.06.008","article-title":"Sleep stage II contributes to the consolidation of declarative memories","volume":"50","author":"Ruch","year":"2012","journal-title":"Neuropsychologia."},{"issue":"4","key":"2026092111370635900_ref93","doi-asserted-by":"publisher","first-page":"e14126","DOI":"10.1111\/jsr.14126","article-title":"An acute bout of high-intensity exercise affects nocturnal sleep and sleep-dependent memory consolidation","volume":"33","author":"Frisch","journal-title":"J Sleep Res"},{"key":"2026092111370635900_ref94","doi-asserted-by":"publisher","first-page":"108054","DOI":"10.1016\/j.nlm.2025.108054","article-title":"Memory impairments observed after a half night sleep restriction are not mediated by working memory, attention, or inhibitory control mechanisms","volume":"219","author":"Heckman","year":"2025","journal-title":"Neurobiol Learn Mem"}],"container-title":["SLEEP"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/academic.oup.com\/sleep\/advance-article-pdf\/doi\/10.1093\/sleep\/zsag235\/70810924\/zsag235.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/academic.oup.com\/sleep\/advance-article-pdf\/doi\/10.1093\/sleep\/zsag235\/70810924\/zsag235.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,21]],"date-time":"2026-09-21T15:37:15Z","timestamp":1790005035000},"score":1,"resource":{"primary":{"URL":"https:\/\/academic.oup.com\/sleep\/advance-article\/doi\/10.1093\/sleep\/zsag235\/8771500"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,26]]},"references-count":94,"URL":"https:\/\/doi.org\/10.1093\/sleep\/zsag235","relation":{},"ISSN":["0161-8105","1550-9109"],"issn-type":[{"value":"0161-8105","type":"print"},{"value":"1550-9109","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,26]]},"article-number":"zsag235"}}