{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,10]],"date-time":"2026-10-10T13:58:02Z","timestamp":1791640682977,"version":"4.3.4"},"reference-count":48,"publisher":"Wiley","issue":"6","license":[{"start":{"date-parts":[[2025,2,21]],"date-time":"2025-02-21T00:00:00Z","timestamp":1740096000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2025,2,21]],"date-time":"2025-02-21T00:00:00Z","timestamp":1740096000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"content-domain":{"domain":["physoc.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Journal of Physiology"],"published-print":{"date-parts":[[2025,3]]},"abstract":"<jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      Respiration shapes brain activity and synchronizes sensory and exploratory motor actions, with some evidence suggesting that it also affects pupil size. However, evidence for a coupling between respiration and pupil size remains scarce and inconclusive, hindered by small sample sizes and limited controls. Given the importance of pupil size in visual perception and as a reflection of brain state, understanding its relationship with respiration is essential. In five experiments using a pre\u2010registered protocol, we systematically investigated how respiratory phase affects pupil size across different conditions. In Experiment 1 (\n                      <jats:italic>n\u00a0=\u00a0<\/jats:italic>\n                      50), we examined nasal and oral breathing at rest under dim lighting with nearby fixation points, then replicated these results under identical conditions in Experiment 2 (\n                      <jats:italic>n\u00a0=\u00a0<\/jats:italic>\n                      53). Experiment 3 (\n                      <jats:italic>n\u00a0=\u00a0<\/jats:italic>\n                      112) extended this to active visual tasks, while Experiment 4 (\n                      <jats:italic>n\u00a0=<\/jats:italic>\n                      \u00a057) extended this to controlled breathing at different paces under ambient lighting with distant fixation. Finally, in Experiment 5 (\n                      <jats:italic>n\u00a0=<\/jats:italic>\n                      \u00a034), individuals with isolated congenital anosmia (born without olfactory bulbs) were used as a lesion\u2010type model during visual\u2013auditory tasks to assess whether the respiratory\u2013pupil link depends on olfactory bulb\u2010driven oscillations.\n                    <\/jats:p>\n                    <jats:p>\n                      Across all conditions \u2013 free and controlled breathing; different tasks, lighting and fixation distances; and with and without olfactory bulbs \u2013 we consistently found that pupil size is smallest around inhalation onset and largest during exhalation. We term this effect the pupillary respiratory\u2010phase response, the fourth known mechanism influencing pupil size, alongside the pupillary light, near fixation and psychosensory responses.\n                      <jats:boxed-text content-type=\"graphic\" position=\"anchor\">\n                        <jats:graphic xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" mimetype=\"image\/png\" position=\"anchor\" specific-use=\"enlarged-web-image\" xlink:href=\"graphic\/tjp16571-gra-0001-m.png\">\n                          <jats:alt-text>image<\/jats:alt-text>\n                        <\/jats:graphic>\n                      <\/jats:boxed-text>\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Key points<\/jats:title>\n                    <jats:p>\n                      <jats:list list-type=\"bullet\">\n                        <jats:list-item>\n                          <jats:p>The influence of respiration on pupil size dynamics has long been debated.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>In this study, we systematically investigated how pupil size changes across the breathing cycle through a series of five experiments, while varying tasks, lighting, fixation distance and brain region involvement.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>We show that pupil size is smallest around inhalation onset and largest during exhalation, with pupil dilatation occurring through most of inhalation and the early phase of exhalation, and pupil constriction occurring primarily during the latter part of exhalation.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>This pattern was consistent across all experimental conditions, demonstrating that it is robust and likely controlled by brainstem circuits.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>We term this effect the pupillary respiratory\u2010phase response, the fourth known mechanism influencing pupil size.<\/jats:p>\n                        <\/jats:list-item>\n                      <\/jats:list>\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1113\/jp287205","type":"journal-article","created":{"date-parts":[[2025,2,21]],"date-time":"2025-02-21T05:06:35Z","timestamp":1740114395000},"page":"1607-1625","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":20,"title":["The pupillary respiratory\u2010phase response: pupil size is smallest around inhalation onset and largest during exhalation"],"prefix":"10.1113","volume":"603","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2942-9962","authenticated-orcid":false,"given":"Martin","family":"Schaefer","sequence":"first","affiliation":[{"name":"Department of Clinical Neuroscience Karolinska Institutet  Stockholm Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3379-8584","authenticated-orcid":false,"given":"Sebastiaan","family":"Math\u00f4t","sequence":"additional","affiliation":[{"name":"Department of Psychology University of Groningen  Groningen The Netherlands"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1011-5234","authenticated-orcid":false,"given":"Mikael","family":"Lundqvist","sequence":"additional","affiliation":[{"name":"Department of Clinical Neuroscience Karolinska Institutet  Stockholm Sweden"},{"name":"Department of Brain and Cognitive Sciences Massachusetts Institute of Technology  Cambridge MA USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3529-8981","authenticated-orcid":false,"given":"Johan N.","family":"Lundstr\u00f6m","sequence":"additional","affiliation":[{"name":"Department of Clinical Neuroscience Karolinska Institutet  Stockholm Sweden"},{"name":"Monell Chemical Senses Center  Philadelphia PA USA"},{"name":"Department of Otorhinolaryngology Karolinska University Hospital  Stockholm Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2282-5903","authenticated-orcid":false,"given":"Artin","family":"Arshamian","sequence":"additional","affiliation":[{"name":"Department of Clinical Neuroscience Karolinska Institutet  Stockholm Sweden"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2025,2,21]]},"reference":[{"key":"e_1_2_5_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/eye.2016.288"},{"key":"e_1_2_5_3_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3360-17.2018"},{"key":"e_1_2_5_4_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev-neuro-090121-014424"},{"key":"e_1_2_5_5_1","doi-asserted-by":"publisher","DOI":"10.18637\/jss.v031.i10"},{"key":"e_1_2_5_6_1","doi-asserted-by":"publisher","DOI":"10.1152\/ajplegacy.1975.228.4.1094"},{"key":"e_1_2_5_7_1","doi-asserted-by":"publisher","DOI":"10.1152\/physrev.00007.2004"},{"key":"e_1_2_5_8_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41583-018-0003-6"},{"key":"e_1_2_5_9_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tins.2006.06.013"},{"issue":"2","key":"e_1_2_5_10_1","first-page":"55","article-title":"[Respiratory effects on the dynamics of the pupillary light reflex in man]","volume":"4","author":"Golenhofen K.","year":"1967","journal-title":"Respiratorische Einflusse Auf Die Dynamik Des Pupillen\u2010Lichtreflexes Beim Menschen"},{"key":"e_1_2_5_11_1","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2016.00115"},{"key":"e_1_2_5_12_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00405-006-0173-0"},{"key":"e_1_2_5_13_1","unstructured":"Inkscape Project. (2018).Inkscape(Version 0.92) [Computer software].https:\/\/inkscape.org"},{"key":"e_1_2_5_14_1","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms4572"},{"key":"e_1_2_5_15_1","unstructured":"JASP Team. (2024).JASP(Version 0.18.3.0) [Computer software].https:\/\/jasp\u2010stats.org\/"},{"key":"e_1_2_5_16_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tics.2020.03.005"},{"key":"e_1_2_5_17_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-28090-5"},{"issue":"4","key":"e_1_2_5_18_1","first-page":"561","article-title":"[Time parameters of the retinal control of light intensity and influencing parameters]","volume":"38","author":"Kaulen S.","year":"1979","journal-title":"Zeitkennwerte Der Retinalen Leuchtstarkeregelung Und Beeinflussende Faktoren"},{"key":"e_1_2_5_19_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tins.2008.11.008"},{"key":"e_1_2_5_20_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2023.102784"},{"key":"e_1_2_5_21_1","doi-asserted-by":"publisher","DOI":"10.1101\/sqb.2014.79.024794"},{"key":"e_1_2_5_22_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1173-24.2024"},{"key":"e_1_2_5_23_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.70907"},{"key":"e_1_2_5_24_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2020.117272"},{"key":"e_1_2_5_25_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.3001457"},{"key":"e_1_2_5_26_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00429-017-1587-8"},{"key":"e_1_2_5_27_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.83654"},{"key":"e_1_2_5_28_1","doi-asserted-by":"publisher","DOI":"10.5334\/joc.18"},{"key":"e_1_2_5_29_1","doi-asserted-by":"publisher","DOI":"10.7717\/peerj.8220"},{"key":"e_1_2_5_30_1","doi-asserted-by":"publisher","DOI":"10.3758\/s13428-022-01957-7"},{"key":"e_1_2_5_31_1","doi-asserted-by":"publisher","DOI":"10.1111\/psyp.13091"},{"key":"e_1_2_5_32_1","doi-asserted-by":"publisher","DOI":"10.3390\/brainsci11101324"},{"key":"e_1_2_5_33_1","doi-asserted-by":"publisher","DOI":"10.1097\/CPM.0b013e31828fc9d6"},{"key":"e_1_2_5_34_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.resp.2018.07.005"},{"key":"e_1_2_5_35_1","doi-asserted-by":"publisher","DOI":"10.1093\/chemse\/bjy045"},{"key":"e_1_2_5_36_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41562-019-0556-z"},{"key":"e_1_2_5_37_1","doi-asserted-by":"publisher","DOI":"10.1002\/hbm.26522"},{"key":"e_1_2_5_38_1","doi-asserted-by":"publisher","DOI":"10.1080\/02713683.2020.1725060"},{"key":"e_1_2_5_39_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00424-022-02729-0"},{"key":"e_1_2_5_40_1","doi-asserted-by":"publisher","DOI":"10.1111\/psyp.14493"},{"key":"e_1_2_5_41_1","doi-asserted-by":"publisher","DOI":"10.1523\/ENEURO.0293\u201016.2017"},{"key":"e_1_2_5_42_1","doi-asserted-by":"publisher","DOI":"10.3389\/fneur.2018.01069"},{"key":"e_1_2_5_43_1","doi-asserted-by":"crossref","unstructured":"Thunell E. Peter M. Iravani B. Porada D. K. Prenner K. Darki F. &Lundstr\u00f6m J. N.(2024).Unisensory visual and auditory objects are processed in olfactory cortex independently of odor association(p. 2023.04.20.537709). bioRxiv.https:\/\/doi.org\/10.1101\/2023.04.20.537709","DOI":"10.1101\/2023.04.20.537709"},{"key":"e_1_2_5_44_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.tins.2018.01.007"},{"key":"e_1_2_5_45_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.resp.2024.104227"},{"key":"e_1_2_5_46_1","doi-asserted-by":"publisher","DOI":"10.3758\/BF03202828"},{"key":"e_1_2_5_47_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cophys.2019.04.023"},{"key":"e_1_2_5_48_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.aai7984"},{"key":"e_1_2_5_49_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2586-16.2016"}],"container-title":["The Journal of Physiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/pdf\/10.1113\/JP287205","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/full-xml\/10.1113\/JP287205","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/pdf\/10.1113\/JP287205","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,26]],"date-time":"2026-06-26T21:34:56Z","timestamp":1782509696000},"score":1,"resource":{"primary":{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/10.1113\/JP287205"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,2,21]]},"references-count":48,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2025,3]]}},"alternative-id":["10.1113\/JP287205"],"URL":"https:\/\/doi.org\/10.1113\/jp287205","archive":["Portico"],"relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2024.06.27.599713","asserted-by":"object"}]},"ISSN":["0022-3751","1469-7793"],"issn-type":[{"value":"0022-3751","type":"print"},{"value":"1469-7793","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,2,21]]},"assertion":[{"value":"2024-07-02","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2025-01-31","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2025-02-21","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}]}}