{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,26]],"date-time":"2026-06-26T07:41:20Z","timestamp":1782459680106,"version":"3.54.5"},"reference-count":106,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,5,15]],"date-time":"2026-05-15T00:00:00Z","timestamp":1778803200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,5,15]],"date-time":"2026-05-15T00:00:00Z","timestamp":1778803200000},"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\u20102024\u201004494"],"award-info":[{"award-number":["RGPIN\u20102024\u201004494"]}],"id":[{"id":"10.13039\/501100000038","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["physoc.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Journal of Physiology"],"abstract":"<jats:sec>\n                    <jats:title>Abstract<\/jats:title>\n                    <jats:p>\n                      Cutaneous mechanoreceptor input from the foot sole is critical for balance and gait, yet ageing and disease can compromise skin sensitivity, increasing risks for falls and related injuries. Heating improves perceptual sensitivity, but its direct effects on afferent firing remain unknown. This study examined how topical heating alters A\u03b2 afferent responses to mechanical loads that closely emulate walking. Single tibial nerve afferents were recorded using microneurography and classified as fast\u2010adapting (FA) or slowly adapting (SA), type I or II. Ramp\u2010and\u2010hold compressions, normalized to regional stance pressure distributions, were applied over receptive fields using a 1 cm\n                      <jats:sup>2<\/jats:sup>\n                      probe before and after heating. Dynamic (ramp) and static (hold) firing rates were compared between temperatures. Thirty\u2010seven afferents (11 SAI, 13 SAII, 9 FAI, 4 FAII) were tested. Heating increased skin temperature by 9.5\u00b1 2.4\u00b0C (\n                      <jats:italic>P<\/jats:italic>\n                      \u00a0&lt;\u00a00.001) and significantly elevated SAI (\n                      <jats:italic>P<\/jats:italic>\n                      \u00a0=\u00a00.001) and SAII (\n                      <jats:italic>P<\/jats:italic>\n                      &lt; 0.001) static firing, as well as SAI (\n                      <jats:italic>P<\/jats:italic>\n                      \u00a0=\u00a00.03), SAII (\n                      <jats:italic>P<\/jats:italic>\n                      \u00a0=\u00a00.028) and FAI dynamic firing (\n                      <jats:italic>P<\/jats:italic>\n                      &lt; 0.001). These findings provide the first direct physiological evidence that heating enhances cutaneous mechanoreceptor activity, offering mechanistic basis for previously observed improvements in perceptual sensitivity. By using gait\u2010like compressions this work also demonstrates that the augmented encoding persists under functionally relevant conditions. Together these findings reveal that heating amplifies static and dynamic mechanoreceptor responses in ways essential to postural control, supporting heat\u2010based stimulation as a practical, non\u2010invasive strategy for improving plantar feedback and reducing fall risk.\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\/tjp70552-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 sole of the foot contains touch receptors that provide critical information for balance and walking, which decline with age and disease, increasing fall risk. Heating can improve touch perception, but the underlying mechanisms are unclear.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>Single nerve fibre activity from the foot sole was recorded in healthy young adults while applying natural, walking\u2010like loads before and after skin heating.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>Heating increased nerve activity in three classes of touch receptors, enhancing sensitivity during application of pressure (dynamic) and steady pressure (static). Static activity relays standing balance input, whereas dynamic activity relays gait and slip feedback.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>Notably in dynamic receptors increases in heat\u2010related nerve activity were more prevalent when loads were applied at faster rates.<\/jats:p>\n                        <\/jats:list-item>\n                        <jats:list-item>\n                          <jats:p>For the first time this work shows how heating differentially affects touch receptor classes and provides a physiological basis for using heat to support balance in future clinical studies.<\/jats:p>\n                        <\/jats:list-item>\n                      <\/jats:list>\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1113\/jp290676","type":"journal-article","created":{"date-parts":[[2026,5,15]],"date-time":"2026-05-15T16:39:08Z","timestamp":1778863148000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Low\u2010threshold cutaneous mechanoreceptors on the foot sole have enhanced sensitivity when skin temperature is elevated"],"prefix":"10.1113","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4476-9431","authenticated-orcid":false,"given":"Erika E.","family":"Howe","sequence":"first","affiliation":[{"name":"Human Health Sciences University of Guelph  Guelph Ontario Canada"},{"name":"Kinesiology and Health Sciences University of Waterloo  Waterloo Ontario Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Apollinaro","sequence":"additional","affiliation":[{"name":"Human Health Sciences University of Guelph  Guelph Ontario Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0938-2570","authenticated-orcid":false,"given":"Joanna","family":"Reeves","sequence":"additional","affiliation":[{"name":"Human Health Sciences University of Guelph  Guelph Ontario Canada"},{"name":"School of Health &amp; Society University of Salford  Salford UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Christopher","family":"Nester","sequence":"additional","affiliation":[{"name":"School Kingdom of Allied Health Professions and Pharmacy University of Keele  Keele Staffordshire UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8951-8783","authenticated-orcid":false,"given":"Leah R.","family":"Bent","sequence":"additional","affiliation":[{"name":"Human Health Sciences University of Guelph  Guelph Ontario Canada"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,5,15]]},"reference":[{"key":"e_1_2_5_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2013.07.051"},{"key":"e_1_2_5_3_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature05910"},{"key":"e_1_2_5_4_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00608.2012"},{"key":"e_1_2_5_5_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.mvr.2015.04.004"},{"key":"e_1_2_5_6_1","doi-asserted-by":"crossref","unstructured":"Bhuiyan S. A. Nagi S. S. Sankaranarayanan I. Semizoglou E. Usoskin D. Yang L. Yu H. Arendt\u2010Tranholm A. Bertels Z. Bhatia P. Bouchatta O. Boyer K. Cervantes A. Chalif J. Chintalapudi H. Cicalo A. Copits B. Cronin C. Curatolo M. \u2026 NIH PRECISION Human Pain Network. (2025).A reference atlas of the human dorsal root ganglion.BioRxiv 2025.11.05.686654.https:\/\/doi.org\/10.1101\/2025.11.05.686654","DOI":"10.1101\/2025.11.05.686654"},{"key":"e_1_2_5_7_1","doi-asserted-by":"publisher","DOI":"10.1054\/foot.1999.0538"},{"key":"e_1_2_5_8_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.1968.31.6.833"},{"key":"e_1_2_5_9_1","doi-asserted-by":"publisher","DOI":"10.1016\/0006-8993(72)90471-4"},{"key":"e_1_2_5_10_1","doi-asserted-by":"publisher","DOI":"10.1038\/39807"},{"key":"e_1_2_5_11_1","doi-asserted-by":"publisher","DOI":"10.1113\/expphysiol.1972.sp002177"},{"key":"e_1_2_5_12_1","doi-asserted-by":"publisher","DOI":"10.1098\/rsif.2023.0052"},{"key":"e_1_2_5_13_1","doi-asserted-by":"publisher","DOI":"10.3390\/jcm10020227"},{"key":"e_1_2_5_14_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00313.2020"},{"key":"e_1_2_5_15_1","doi-asserted-by":"publisher","DOI":"10.1002\/j.2040-4603.2020.tb00131.x"},{"key":"e_1_2_5_16_1","doi-asserted-by":"publisher","DOI":"10.1119\/1.19253"},{"key":"e_1_2_5_17_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.64679"},{"key":"e_1_2_5_18_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-023-44895-w"},{"key":"e_1_2_5_19_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00628.2004"},{"key":"e_1_2_5_20_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.1991.65.3.657"},{"key":"e_1_2_5_21_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0966-6362(03)00095-X"},{"key":"e_1_2_5_22_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00359.2005"},{"key":"e_1_2_5_23_1","doi-asserted-by":"publisher","DOI":"10.3390\/jcm10194609"},{"key":"e_1_2_5_24_1","doi-asserted-by":"publisher","DOI":"10.1111\/joa.12983"},{"key":"e_1_2_5_25_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00414.2001"},{"key":"e_1_2_5_26_1","doi-asserted-by":"publisher","DOI":"10.3758\/BF03214234"},{"issue":"4","key":"e_1_2_5_27_1","first-page":"327","article-title":"The effect of skin temperature on the perception of roughness","volume":"3","author":"Green B. G.","year":"1979","journal-title":"Sensory Processes"},{"key":"e_1_2_5_28_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.01033.2003"},{"key":"e_1_2_5_29_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1995.sp021075"},{"key":"e_1_2_5_30_1","doi-asserted-by":"publisher","DOI":"10.3233\/THC-140877"},{"key":"e_1_2_5_31_1","doi-asserted-by":"publisher","DOI":"10.1016\/0167-9457(92)90015-4"},{"key":"e_1_2_5_32_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00424-014-1651-7"},{"key":"e_1_2_5_33_1","doi-asserted-by":"publisher","DOI":"10.1016\/B978-0-08-045396-5.00222-0"},{"key":"e_1_2_5_34_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41586-019-1345-6"},{"key":"e_1_2_5_35_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00230848"},{"key":"e_1_2_5_36_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnins.2024.1329832"},{"key":"e_1_2_5_37_1","doi-asserted-by":"publisher","DOI":"10.1152\/japplphysiol.00533.2023"},{"key":"e_1_2_5_38_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00176.2014"},{"key":"e_1_2_5_39_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1978.sp012411"},{"key":"e_1_2_5_40_1","doi-asserted-by":"publisher","DOI":"10.1016\/0166-2236(83)90011-5"},{"key":"e_1_2_5_41_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.1979.sp013048"},{"key":"e_1_2_5_42_1","doi-asserted-by":"publisher","DOI":"10.1002\/j.2040-4603.2014.tb00541.x"},{"key":"e_1_2_5_43_1","doi-asserted-by":"publisher","DOI":"10.1111\/bph.12414"},{"key":"e_1_2_5_44_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.isci.2022.105874"},{"key":"e_1_2_5_45_1","doi-asserted-by":"publisher","DOI":"10.1097\/00001756-199810050-00021"},{"key":"e_1_2_5_46_1","doi-asserted-by":"publisher","DOI":"10.1152\/japplphysiol.01071.2005"},{"key":"e_1_2_5_47_1","doi-asserted-by":"publisher","DOI":"10.1113\/jphysiol.2001.013087"},{"key":"e_1_2_5_48_1","doi-asserted-by":"publisher","DOI":"10.1113\/EP092348"},{"key":"e_1_2_5_49_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1748-1716.1970.tb04783.x"},{"key":"e_1_2_5_50_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.jdiacomp.2017.05.014"},{"key":"e_1_2_5_51_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1748-1716.1987.tb08065.x"},{"key":"e_1_2_5_52_1","doi-asserted-by":"publisher","DOI":"10.1123\/mc.2021-0124"},{"key":"e_1_2_5_53_1","doi-asserted-by":"publisher","DOI":"10.1046\/j.1523-1747.2000.00055.x"},{"key":"e_1_2_5_54_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gaitpost.2021.01.016"},{"key":"e_1_2_5_55_1","doi-asserted-by":"publisher","DOI":"10.1111\/srt.12094"},{"key":"e_1_2_5_56_1","doi-asserted-by":"publisher","DOI":"10.1038\/nature05662"},{"key":"e_1_2_5_57_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1440-1681.2005.04143.x"},{"key":"e_1_2_5_58_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.foot.2021.101887"},{"key":"e_1_2_5_59_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gaitpost.2023.03.013"},{"key":"e_1_2_5_60_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gaitpost.2017.07.004"},{"key":"e_1_2_5_61_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1475-097X.2012.01137.x"},{"key":"e_1_2_5_62_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.exger.2017.09.007"},{"key":"e_1_2_5_63_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00221-006-0568-6"},{"key":"e_1_2_5_64_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00376.2020"},{"key":"e_1_2_5_65_1","article-title":"Passive heating of the skin softens skin across the foot sole","author":"Nolan B.","journal-title":"In Prep"},{"key":"e_1_2_5_66_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0268-0033(01)00090-0"},{"key":"e_1_2_5_67_1","doi-asserted-by":"publisher","DOI":"10.1002\/cne.21262"},{"key":"e_1_2_5_68_1","doi-asserted-by":"publisher","DOI":"10.1115\/1.2800860"},{"key":"e_1_2_5_69_1","doi-asserted-by":"publisher","DOI":"10.1126\/science.1073140"},{"key":"e_1_2_5_70_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neulet.2005.08.060"},{"key":"e_1_2_5_71_1","doi-asserted-by":"publisher","DOI":"10.1093\/gerona\/63.6.595"},{"key":"e_1_2_5_72_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00339.2016"},{"key":"e_1_2_5_73_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0140-6736(03)14470-4"},{"key":"e_1_2_5_74_1","doi-asserted-by":"publisher","DOI":"10.1002\/ana.20670"},{"key":"e_1_2_5_75_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.gaitpost.2007.06.009"},{"key":"e_1_2_5_76_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.91041.2008"},{"key":"e_1_2_5_77_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00221-022-06402-x"},{"key":"e_1_2_5_78_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnagi.2024.1387136"},{"key":"e_1_2_5_79_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neulet.2009.07.006"},{"key":"e_1_2_5_80_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.clinph.2009.06.010"},{"key":"e_1_2_5_81_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cnp.2016.12.005"},{"key":"e_1_2_5_82_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0021-9290(97)00065-1"},{"key":"e_1_2_5_83_1","doi-asserted-by":"publisher","DOI":"10.2522\/ptj.20060083"},{"key":"e_1_2_5_84_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.ceca.2014.02.015"},{"key":"e_1_2_5_85_1","doi-asserted-by":"publisher","DOI":"10.3758\/BF03206232"},{"key":"e_1_2_5_86_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00647.2016"},{"key":"e_1_2_5_87_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00524.2015"},{"key":"e_1_2_5_88_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00848.2017"},{"key":"e_1_2_5_89_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M302561200"},{"key":"e_1_2_5_90_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00933.2017"},{"key":"e_1_2_5_91_1","doi-asserted-by":"publisher","DOI":"10.1136\/jnnp.44.7.600"},{"key":"e_1_2_5_92_1","doi-asserted-by":"publisher","DOI":"10.1016\/0304-3940(82)90190-2"},{"key":"e_1_2_5_93_1","doi-asserted-by":"publisher","DOI":"10.1121\/1.394047"},{"key":"e_1_2_5_94_1","doi-asserted-by":"publisher","DOI":"10.1080\/089902203100105163"},{"key":"e_1_2_5_95_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M208277200"},{"key":"e_1_2_5_96_1","doi-asserted-by":"publisher","DOI":"10.1113\/JP282873"},{"key":"e_1_2_5_97_1","doi-asserted-by":"publisher","DOI":"10.1037\/h0059426"},{"key":"e_1_2_5_98_1","doi-asserted-by":"publisher","DOI":"10.1093\/gerona\/58.8.B680"},{"key":"e_1_2_5_99_1","doi-asserted-by":"publisher","DOI":"10.1007\/BF00236209"},{"issue":"197","key":"e_1_2_5_100_1","article-title":"Intradermal microdialysis: An approach to investigating novel mechanisms of microvascular dysfunction in humans","volume":"202","author":"Williams A. C.","year":"2023","journal-title":"Journal of Visualized Experiments"},{"issue":"1","key":"e_1_2_5_101_1","first-page":"92","article-title":"Temperature\u2010dependent mechanical behaviors of skin tissue","volume":"35","author":"Xu F.","year":"2008","journal-title":"IAENG International Journal of Computer Science"},{"key":"e_1_2_5_102_1","doi-asserted-by":"publisher","DOI":"10.3389\/fnmol.2022.982202"},{"key":"e_1_2_5_103_1","doi-asserted-by":"publisher","DOI":"10.1007\/978-3-642-54215-2_11"},{"key":"e_1_2_5_104_1","doi-asserted-by":"publisher","DOI":"10.3390\/brainsci10120966"},{"key":"e_1_2_5_105_1","doi-asserted-by":"publisher","DOI":"10.1186\/2052-1847-6-33"},{"key":"e_1_2_5_106_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0301-0082(98)00081-1"},{"key":"e_1_2_5_107_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1910213116"}],"container-title":["The Journal of Physiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/pdf\/10.1113\/JP290676","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/full-xml\/10.1113\/JP290676","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/pdf\/10.1113\/JP290676","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,15]],"date-time":"2026-06-15T12:27:36Z","timestamp":1781526456000},"score":1,"resource":{"primary":{"URL":"https:\/\/physoc.onlinelibrary.wiley.com\/doi\/10.1113\/JP290676"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,5,15]]},"references-count":106,"alternative-id":["10.1113\/JP290676"],"URL":"https:\/\/doi.org\/10.1113\/jp290676","archive":["Portico"],"relation":{},"ISSN":["0022-3751","1469-7793"],"issn-type":[{"value":"0022-3751","type":"print"},{"value":"1469-7793","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,5,15]]},"assertion":[{"value":"2025-12-17","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-04-06","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-05-15","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"JP290676"}}