{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T14:45:35Z","timestamp":1792075535559,"version":"4.4.0"},"reference-count":29,"publisher":"Wiley","issue":"8","license":[{"start":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T00:00:00Z","timestamp":1792022400000},"content-version":"vor","delay-in-days":14,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,10,1]],"date-time":"2026-10-01T00:00:00Z","timestamp":1790812800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100003502","name":"Ella ja Georg Ehrnroothin S\u00e4\u00e4ti\u00f6","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100003502","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100002341","name":"Academy of Finland","doi-asserted-by":"publisher","award":["355392"],"award-info":[{"award-number":["355392"]}],"id":[{"id":"10.13039\/501100002341","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Eur J of Neuroscience"],"published-print":{"date-parts":[[2026,10]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Previous studies suggest that when a tone\u2010conditioned stimulus (CS) predicting an airpuff towards the eye is presented at diastole during expiration, both sensory responses to the CS and trace eyeblink conditioning (TEBC) are enhanced. However, not all participants learn and individual variation in the body\u2013brain relationship remains overlooked. We studied whether timing the CS based on an individually optimized bodily state enhances sensory responses and TEBC. We assumed so, as sensory response magnitude reflects attention and stimulus salience: A salient CS should facilitate TEBC. Forty\u2010one young adults and 19 older adults completed a passive auditory task while electrocardiogram, respiration and electroencephalogram were recorded. The cardiorespiratory state associated with the largest N1\u2013P2 response was identified individually. Optimal states varied across participants and did not differ between age groups. Participants underwent TEBC at their optimal state (Personalized) or at diastole during expiration (Control). As expected, the CS evoked a larger N1\u2013P2 response in the Personalized than the Control group. However, conditioned responding, peak performance, and learning rate, or the proportion of good learners did not differ between groups. Older adults learned more slowly, made fewer conditioned responses, and were less aware of the tone\u2010airpuff association than young adults. These findings tentatively suggest that enhancing early sensory processing of the CS may not improve TEBC, although the low reliability of the individually determined optimal state limits this conclusion. Future studies should examine whether timing external stimuli to specific bodily states effectively influences behaviour in tasks that rely on rapid sensory processing.<\/jats:p>","DOI":"10.1111\/ejn.70722","type":"journal-article","created":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T14:00:38Z","timestamp":1792072838000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Optimized Cardiorespiratory State at Tone Onset Enhances Sensory Responses but Not Associative Learning in Healthy Adults"],"prefix":"10.1111","volume":"64","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-4244-485X","authenticated-orcid":false,"given":"Praghajieeth Raajhen","family":"Santhana Gopalan","sequence":"first","affiliation":[{"name":"Department of Psychology and Centre for Interdisciplinary Brain Research University of Jyv\u00e4skyl\u00e4  Jyv\u00e4skyl\u00e4 Finland"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2222-6118","authenticated-orcid":false,"given":"Miriam S.","family":"Nokia","sequence":"additional","affiliation":[{"name":"Department of Psychology and Centre for Interdisciplinary Brain Research University of Jyv\u00e4skyl\u00e4  Jyv\u00e4skyl\u00e4 Finland"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,10,15]]},"reference":[{"key":"e_1_2_11_2_1","doi-asserted-by":"crossref","DOI":"10.1093\/acprof:oso\/9780195301069.001.0001","volume-title":"Rhythms of the Brain","author":"Buzsaki G.","year":"2006"},{"issue":"11","key":"e_1_2_11_3_1","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1038\/s41583-018-0068-2","article-title":"Maintenance, Reserve and Compensation: The Cognitive Neuroscience of Healthy Ageing","volume":"19","author":"Cabeza R.","year":"2018","journal-title":"Nature Reviews Neuroscience"},{"issue":"23","key":"e_1_2_11_4_1","doi-asserted-by":"crossref","first-page":"8108","DOI":"10.1073\/pnas.0800374105","article-title":"Neural Substrates Underlying Human Delay and Trace Eyeblink Conditioning","volume":"105","author":"Cheng D. T.","year":"2008","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"3","key":"e_1_2_11_5_1","doi-asserted-by":"crossref","first-page":"684","DOI":"10.1037\/a0017978","article-title":"The Effects of Aging in Delay and Trace Human Eyeblink Conditioning","volume":"25","author":"Cheng D. T.","year":"2010","journal-title":"Psychology and Aging"},{"key":"e_1_2_11_6_1","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1016\/j.jneumeth.2003.10.009","article-title":"EEGLAB: An Open\u2010Source Toolbox for Analysis of Single\u2010Trial EEG Dynamics","volume":"134","author":"Delorme A.","year":"2004","journal-title":"Journal of Neuroscience Methods"},{"issue":"10","key":"e_1_2_11_7_1","doi-asserted-by":"crossref","first-page":"1670","DOI":"10.1038\/s41593-023-01425-1","article-title":"Interoceptive Rhythms in the Brain","volume":"26","author":"Engelen T.","year":"2023","journal-title":"Nature Neuroscience"},{"issue":"3","key":"e_1_2_11_8_1","doi-asserted-by":"crossref","first-page":"208","DOI":"10.1016\/j.tics.2019.12.006","article-title":"Theta Oscillations in Human Memory","volume":"24","author":"Herweg N. A.","year":"2020","journal-title":"Trends in Cognitive Sciences"},{"issue":"4108","key":"e_1_2_11_9_1","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1126\/science.182.4108.177","article-title":"Electrical Signs of Selective Attention in the Human Brain","volume":"182","author":"Hillyard S. A.","year":"1973","journal-title":"Science"},{"issue":"1","key":"e_1_2_11_10_1","doi-asserted-by":"crossref","DOI":"10.1038\/s41467-023-40250-9","article-title":"Modulatory Dynamics of Periodic and Aperiodic Activity in Respiration\u2010Brain Coupling","volume":"14","author":"Kluger D. S.","year":"2023","journal-title":"Nature Communications"},{"issue":"4","key":"e_1_2_11_11_1","doi-asserted-by":"crossref","first-page":"747","DOI":"10.1037\/0735-7044.115.4.747","article-title":"Awareness in Classical Differential Eyeblink Conditioning in Young and Aging Humans","volume":"115","author":"Knuttinen M. G.","year":"2001","journal-title":"Behavioral Neuroscience"},{"issue":"37","key":"e_1_2_11_12_1","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2311953121","article-title":"Interoceptive Signals Shape the Earliest Markers and Neural Pathway to Awareness at the Visual Threshold","volume":"121","author":"Leupin V.","year":"2024","journal-title":"Proceedings of the National Academy of Sciences of the United States of America"},{"issue":"5","key":"e_1_2_11_13_1","doi-asserted-by":"crossref","DOI":"10.1111\/psyp.14506","article-title":"Effects of the Cardiac Cycle on Auditory Processing: A Preregistered Study on Mismatch Negativity","volume":"61","author":"Li L.","year":"2024","journal-title":"Psychophysiology"},{"key":"e_1_2_11_14_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.ijpsycho.2025.112521","article-title":"Effects of Cardiac and Respiratory Phases on Auditory Evoked Potentials","volume":"209","author":"Mizuhara K.","year":"2025","journal-title":"International Journal of Psychophysiology: Official Journal of the International Organization of Psychophysiology"},{"issue":"6","key":"e_1_2_11_15_1","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1101\/lm.038166.115","article-title":"Phase Matters: Responding to and Learning About Peripheral Stimuli Depends on Hippocampal \u03b8 Phase at Stimulus Onset","volume":"22","author":"Nokia M. S.","year":"2015","journal-title":"Learning & Memory (Cold Spring Harbor, N.Y.)"},{"issue":"11","key":"e_1_2_11_16_1","doi-asserted-by":"crossref","first-page":"1228","DOI":"10.1002\/hipo.23551","article-title":"CA3\u2010CA1 Long\u2010Term Potentiation Occurs Regardless of Respiration and Cardiac Cycle Phases in Urethane\u2010Anesthetized Rats","volume":"33","author":"Nokia M. S.","year":"2023","journal-title":"Hippocampus"},{"key":"e_1_2_11_17_1","doi-asserted-by":"crossref","DOI":"10.1155\/2011\/156869","article-title":"FieldTrip: Open Source Software for Advanced Analysis of MEG, EEG, and Invasive Electrophysiological Data","volume":"2011","author":"Oostenveld R.","year":"2011","journal-title":"Computational Intelligence and Neuroscience"},{"key":"e_1_2_11_18_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.neubiorev.2022.104908","article-title":"Cardiorespiratory Rhythms, Brain Oscillatory Activity and Cognition: Review of Evidence and Proposal for Significance","volume":"142","author":"Parviainen T.","year":"2022","journal-title":"Neuroscience and Biobehavioral Reviews"},{"issue":"6","key":"e_1_2_11_19_1","doi-asserted-by":"crossref","first-page":"532","DOI":"10.1037\/0033-295X.87.6.532","article-title":"A Model for Pavlovian Learning: Variations in the Effectiveness of Conditioned but Not of Unconditioned Stimuli","volume":"87","author":"Pearce J. M.","year":"1980","journal-title":"Psychological Review"},{"issue":"5","key":"e_1_2_11_20_1","doi-asserted-by":"crossref","first-page":"797","DOI":"10.1152\/jn.00356.2023","article-title":"Cardiorespiratory Rhythm\u2010Contingent Trace Eyeblink Conditioning in Elderly Adults","volume":"131","author":"Santhana Gopalan P. R.","year":"2024","journal-title":"Journal of Neurophysiology"},{"issue":"3","key":"e_1_2_11_21_1","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1016\/j.bandc.2009.08.002","article-title":"Cardiac Modulation of Startle: Effects on Eye Blink and Higher Cognitive Processing","volume":"71","author":"Schulz A.","year":"2009","journal-title":"Brain and Cognition"},{"issue":"1708","key":"e_1_2_11_22_1","doi-asserted-by":"crossref","DOI":"10.1098\/rstb.2016.0019","article-title":"Respiratory Modulation of Startle Eye Blink: A New Approach to Assess Afferent Signals From the Respiratory System","volume":"371","author":"Schulz A.","year":"2016","journal-title":"Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences"},{"key":"e_1_2_11_23_1","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1016\/j.ijpsycho.2020.08.005","article-title":"Cardiac Cycle Phases Affect Auditory\u2010Evoked Potentials, Startle Eye Blink and Pre\u2010Motor Reaction Times in Response to Acoustic Startle Stimuli","volume":"157","author":"Schulz A.","year":"2020","journal-title":"International Journal of Psychophysiology: Official Journal of the International Organization of Psychophysiology"},{"key":"e_1_2_11_24_1","doi-asserted-by":"crossref","DOI":"10.3389\/fpubh.2017.00258","article-title":"An Overview of Heart Rate Variability Metrics and Norms","volume":"5","author":"Shaffer F.","year":"2017","journal-title":"Frontiers in Public Health"},{"issue":"3","key":"e_1_2_11_25_1","doi-asserted-by":"crossref","first-page":"732","DOI":"10.1016\/j.neuroscience.2009.01.041","article-title":"The Role of the Cerebellum in Classical Conditioning of Discrete Behavioral Responses","volume":"162","author":"Thompson R. F.","year":"2009","journal-title":"Neuroscience"},{"issue":"6","key":"e_1_2_11_26_1","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1038\/s41583-025-00920-7","article-title":"Global Coordination of Brain Activity by the Breathing Cycle","volume":"26","author":"Tort A. B. L.","year":"2025","journal-title":"Nature Reviews. Neuroscience"},{"issue":"2","key":"e_1_2_11_27_1","doi-asserted-by":"crossref","first-page":"830","DOI":"10.1152\/jn.00128.2018","article-title":"Learning by Heart: Cardiac Cycle Reveals an Effective Time Window for Learning","volume":"120","author":"Waselius T.","year":"2018","journal-title":"Journal of Neurophysiology"},{"issue":"9","key":"e_1_2_11_28_1","doi-asserted-by":"crossref","DOI":"10.1111\/psyp.13387","article-title":"Breathe Out and Learn: Expiration\u2010Contingent Stimulus Presentation Facilitates Associative Learning in Trace Eyeblink Conditioning","volume":"56","author":"Waselius T.","year":"2019","journal-title":"Psychophysiology"},{"issue":"3","key":"e_1_2_11_29_1","doi-asserted-by":"crossref","first-page":"767","DOI":"10.1152\/jn.00298.2021","article-title":"Cardiac Cycle and Respiration Phase Affect Responses to the Conditioned Stimulus in Young Adults Trained in Trace Eyeblink Conditioning","volume":"127","author":"Waselius T.","year":"2022","journal-title":"Journal of Neurophysiology"},{"issue":"3","key":"e_1_2_11_30_1","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1037\/a0023520","article-title":"Exploring Prefrontal Cortical Memory Mechanisms With Eyeblink Conditioning","volume":"125","author":"Weiss C.","year":"2011","journal-title":"Behavioral Neuroscience"}],"container-title":["European Journal of Neuroscience"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/ejn.70722","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1111\/ejn.70722","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1111\/ejn.70722","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T14:00:43Z","timestamp":1792072843000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1111\/ejn.70722"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,10]]},"references-count":29,"journal-issue":{"issue":"8","published-print":{"date-parts":[[2026,10]]}},"alternative-id":["10.1111\/ejn.70722"],"URL":"https:\/\/doi.org\/10.1111\/ejn.70722","archive":["Portico"],"relation":{},"ISSN":["0953-816X","1460-9568"],"issn-type":[{"value":"0953-816X","type":"print"},{"value":"1460-9568","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,10]]},"assertion":[{"value":"2026-06-11","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-07","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-10-15","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70722"}}