{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,1]],"date-time":"2026-09-01T15:27:04Z","timestamp":1788276424461,"version":"build-2803163510"},"reference-count":106,"publisher":"Wiley","issue":"17","license":[{"start":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T00:00:00Z","timestamp":1785110400000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2026,7,27]],"date-time":"2026-07-27T00:00:00Z","timestamp":1785110400000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100000038","name":"Natural Sciences and Engineering Research Council of Canada","doi-asserted-by":"publisher","award":["RGPIN\u20102021\u201003980"],"award-info":[{"award-number":["RGPIN\u20102021\u201003980"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100004411","name":"Heart and Stroke Foundation of Canada","doi-asserted-by":"publisher","award":["NI\u201024\u20100038149"],"award-info":[{"award-number":["NI\u201024\u20100038149"]}],"id":[{"id":"10.13039\/100004411","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["physoc.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Journal of Physiology"],"published-print":{"date-parts":[[2026,9]]},"abstract":"\n Abstract<\/jats:title>\n \n How ventilation () matches gas exchange () while maintaining isocapnia during exercise remains unresolved. We measured , , arterialized\u2010venous (P\u0101CO\n 2<\/jats:sub>\n ) and hypercapnic ventilatory responsiveness (HCVR\u00a0=\u00a0\u2206\/\u2206) during four steady\u2010state exercise intensities below the respiratory compensation point (RCP) and used a mathematical model to characterize feedforward contributions to exercise hyperpnoea and test the hypothesis that CO\n 2<\/jats:sub>\n chemofeedback is inhibitory. Model assumptions were that the ventilatory response to exercise\u2010intensity changes (G\n SYS<\/jats:sub>\n \u00a0=\u00a0\u2206\/\u2206) equals the sum of a feedforward exercise stimulus proportional to (G\n EX<\/jats:sub>\n ) modified by the CO\n 2<\/jats:sub>\n chemoreflex (HCVR). Using experimental data G\n SYS<\/jats:sub>\n was quantified as \u2206\/\u2206 between exercise steps and G\n EX<\/jats:sub>\n equalled G\n SYS<\/jats:sub>\n minus chemoreflex feedback (i.e. HCVRx\u2206P\u0101CO\n 2<\/jats:sub>\n \/\u2206). Twenty healthy young participants completed a maximal cycling test and three submaximal step\u2010incremental protocols of two 8 min stages below the estimated lactate threshold (\u03b8\n LT<\/jats:sub>\n ; MOD\n 1\u00a0<\/jats:sub>\n and MOD\n 2<\/jats:sub>\n ) and two 12 min stages between \u03b8\n LT<\/jats:sub>\n and RCP (HVY\n 1<\/jats:sub>\n and HVY\n 2<\/jats:sub>\n ). During visits 2 and 3, three rebreathing tests performed at each intensity provided HCVR (average \u2013end\u2010tidal slope). During visit 4, heated hand vein blood provided P\u0101CO\n 2<\/jats:sub>\n at each intensity, and and were measured by metabolic cart. With higher steady\u2010state , one\u2010way repeated\u2010measures ANOVA showed that HCVR rose from 2.4 \u00b1 0.9 at MOD\n 1<\/jats:sub>\n to 2.8 \u00b1 0.8, 3.2 \u00b1 0.8 and 3.6 \u00b1 1.0 L\u2009\u00b7\u2009min\n \u22121<\/jats:sup>\n \u2009\u00b7\u2009mmHg\n \u22121<\/jats:sup>\n at MOD\n 2<\/jats:sub>\n , HVY\n 1<\/jats:sub>\n and HVY\n 2<\/jats:sub>\n , respectively; \u2206P\u0101CO\n 2<\/jats:sub>\n \/\u2206 decreased (\n P<\/jats:italic>\n \u00a0<\u00a00.001) from 0.2 \u00b1 4.3 at MOD\n 1<\/jats:sub>\n to \u22121.9 \u00b1 5.0, \u22125.5 \u00b1 4.5 and \u22128.0 \u00b1 4.6\u00a0mmHg\u2009\u00b7\u2009L\u2009\u00b7\u2009min\n \u22121<\/jats:sup>\n ; and G\n EX<\/jats:sub>\n increased (\n P<\/jats:italic>\n \u00a0<\u00a00.001) from 26.9 \u00b1 15.1 at MOD\n 1<\/jats:sub>\n to 34.5 \u00b1 20.3, 53.9 \u00b1 19.1 and 73.4 \u00b1 27.5 L\u2009\u00b7\u2009min\n \u22121<\/jats:sup>\n . Exercise hyperpnoea may reflect the balance of intensity\u2010dependent feedforward and inhibitory CO\n 2<\/jats:sub>\n chemofeedback drive.\n \n \n image<\/jats:alt-text>\n <\/jats:graphic>\n <\/jats:boxed-text>\n <\/jats:p>\n <\/jats:sec>\n \n Key points<\/jats:title>\n \n \n \n \n We aimed to quantify exercise intensity\u2010dependent changes in CO\n 2<\/jats:sub>\n chemoreflexes and interpret their role in exercise hyperpnoea.\n <\/jats:p>\n <\/jats:list-item>\n \n \n Ventilatory, gas exchange, arterialized\u2010venous (P\u0101CO\n 2<\/jats:sub>\n ) and hypercapnic ventilatory responses (HCVR) were measured during successive work\u2010to\u2010work cycling exercise transitions from low\u2010to\u2010moderate to heavy\u2010intensity work rates.\n <\/jats:p>\n <\/jats:list-item>\n \n \n The measured HCVR at each intensity and \u2206\/\u2206 and \u2206P\u0101CO\n 2<\/jats:sub>\n \/\u2206 from one work rate to the next were included in a mathematical model to characterize feedforward contributions to exercise hyperpnoea at four intensities.\n <\/jats:p>\n <\/jats:list-item>\n \n \n With increasing steady\u2010state , \u2206\/\u2206 and HCVR progressively increased, whereas \u2206P\u0101CO\n 2<\/jats:sub>\n \/\u2206 progressively became more negative.\n <\/jats:p>\n <\/jats:list-item>\n \n \n Assuming \u2206 is the sum of a feedforward gain\n plus<\/jats:italic>\n linear CO\n 2<\/jats:sub>\n chemofeedback gain responding to changes in and from rest to exercise, respectively, we estimated that feedforward gain rises with increasing exercise intensities, attaining a drive that exceeds requirements for regulation at its resting level.\n <\/jats:p>\n <\/jats:list-item>\n \n Exercise hyperpnoea, therefore,\u00a0requires a feedforward exercise stimulus constrained by inhibitory chemofeedback from rest to exercise to minimize hypocapnia .<\/jats:p>\n <\/jats:list-item>\n <\/jats:list>\n <\/jats:p>\n 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