{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T14:00:03Z","timestamp":1790172003286,"version":"4.1.0"},"reference-count":38,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T00:00:00Z","timestamp":1790121600000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100001665","name":"Agence Nationale de la Recherche","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100001665","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100000323","name":"Pain Relief Foundation","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100000323","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100004097","name":"Fondation ARC pour la Recherche sur le Cancer","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004097","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["physoc.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The Journal of Physiology"],"abstract":"\n Abstract<\/jats:title>\n \n Stroking touch has been shown to be most pleasant at intermediate velocities of 1\u201310 cm\u00a0s\n \u22121<\/jats:sup>\n , which relates well to the activity of C\u2010low\u2010threshold mechanoreceptors (C\u2010LTMRs), also called C\u2010tactile afferents in humans, that have been implicated in encoding positive affective touch. This well\u2010established finding has been demonstrated at gentle stroking forces (typically, 0.4 N peak normal force), yet few studies have investigated the effect of force on the perception of stroking touch or on the activity of C\u2010LTMRs. We investigated the pleasantness and intensity of stroking touch (0.3, 1, 3, 10 and 30 cm\u00a0s\n \u22121<\/jats:sup>\n ) at different forces (0.2, 0.4, 0.8, 1.2, 1.6 and 2.0 N) on hairy forearm skin and compared this with responses from C\u2010LTMRs during stroking touch at the same velocities, but over fewer forces (0.05, 0.4 and 1.5 N). We found significant effects of stroking velocity and force for both tactile pleasantness and intensity ratings. Pleasantness showed the typical inverted U\u2010shaped relationship over stroking velocities, but this was modified by force; higher forces significantly decreased pleasantness at faster velocities. Conversely, tactile intensity increased linearly with both increasing velocity and force. Recordings from seven C\u2010LTMRs showed that their firing frequency changed with stroking velocity, but that they were strongly modulated by force. Overall, we demonstrate the profound effect that force has on the perception of stroking touch, whereby tactile pleasantness and intensity appear to be a multi\u2010faceted construct that is related to C\u2010LTMR and A\u03b2\u2010LTMR activity, respectively, but that the firing in individual mechanoreceptor populations cannot account fully for percepts.\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 The skin is highly sensitive to small mechanical deformations, including dynamic touch and different forces.<\/jats:p>\n <\/jats:list-item>\n \n We obtained perceptual ratings for touch pleasantness and intensity over different stroking velocities and forces applied to the forearm. This was also compared with microneurography recordings from C\u2010low\u2010threshold mechanoreceptors (C\u2010LTMRs).<\/jats:p>\n <\/jats:list-item>\n \n Building on previous work, we find that it is not only stroking velocity that affects pleasantness and intensity perception, but also stroking force. Pleasantness ratings decreased more at higher forces when the stroking was faster. C\u2010LTMR firing showed that differences in force were readily encoded, although the same force\u2013velocity interaction was not seen.<\/jats:p>\n <\/jats:list-item>\n \n This work expands the literature on affective touch to encompass the effects of force, which clearly modulates pleasantness.<\/jats:p>\n <\/jats:list-item>\n \n C\u2010LTMR force and velocity information is likely to combine with information from all types of mechanoreceptors, including those from the fascia, to produce the resulting percept.<\/jats:p>\n <\/jats:list-item>\n <\/jats:list>\n <\/jats:p>\n <\/jats:sec>","DOI":"10.1113\/jp290946","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T13:05:00Z","timestamp":1790168700000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["The influence of force on the encoding and perception of affective touch"],"prefix":"10.1113","author":[{"ORCID":"https:\/\/orcid.org\/0009-0005-2242-2559","authenticated-orcid":false,"given":"Sophia","family":"Faresse","sequence":"first","affiliation":[{"name":"Aix Marseille Univ, CNRS CRPN (Centre de Recherche en Psychologie et Neurosciences \u2013 UMR 7077) Marseille France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-9814-367X","authenticated-orcid":false,"given":"Luidgi","family":"Thrace","sequence":"additional","affiliation":[{"name":"Aix Marseille Univ, CNRS CRPN (Centre de Recherche en Psychologie et Neurosciences \u2013 UMR 7077) Marseille France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4744-4963","authenticated-orcid":false,"given":"S. Hasan","family":"Ali","sequence":"additional","affiliation":[{"name":"Institute of Population Health, Department of Psychological Sciences University of Liverpool Liverpool UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2192-3862","authenticated-orcid":false,"given":"Adarsh","family":"Makdani","sequence":"additional","affiliation":[{"name":"Research Centre for Brain & Behaviour, School of Psychology Liverpool John Moores University Liverpool UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Francis","family":"McGlone","sequence":"additional","affiliation":[{"name":"Faculty of Science & Engineering, School of Life Sciences Manchester Metropolitan University Manchester UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8273-7089","authenticated-orcid":false,"given":"Andrew G.","family":"Marshall","sequence":"additional","affiliation":[{"name":"Institute of Life Course and Medicine Sciences, Pain Research Institute University of Liverpool Liverpool UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2667-011X","authenticated-orcid":false,"given":"Paula D.","family":"Trotter","sequence":"additional","affiliation":[{"name":"Research Centre for Brain & Behaviour, School of Psychology Liverpool John Moores University Liverpool UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4621-7929","authenticated-orcid":false,"given":"Rochelle","family":"Ackerley","sequence":"additional","affiliation":[{"name":"Aix Marseille Univ, CNRS CRPN (Centre de Recherche en Psychologie et Neurosciences \u2013 UMR 7077) Marseille France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,23]]},"reference":[{"key":"e_1_2_5_2_1","doi-asserted-by":"crossref","first-page":"95","DOI":"10.1016\/j.cobeha.2021.08.012","article-title":"C\u2010tactile (CT) afferents: Evidence of their function from microneurography studies in humans","volume":"43","author":"Ackerley R.","year":"2022","journal-title":"Current Opinion in Behavioral Sciences"},{"key":"e_1_2_5_3_1","doi-asserted-by":"crossref","first-page":"54","DOI":"10.3389\/fnbeh.2014.00054","article-title":"Touch perceptions across skin sites: Differences between sensitivity, direction discrimination and pleasantness","volume":"8","author":"Ackerley R.","year":"2014","journal-title":"Frontiers in Behavioral Neuroscience"},{"issue":"8","key":"e_1_2_5_4_1","doi-asserted-by":"crossref","first-page":"2879","DOI":"10.1523\/JNEUROSCI.2847-13.2014","article-title":"Human C\u2010tactile afferents are tuned to the temperature of a skin\u2010stroking caress","volume":"34","author":"Ackerley R.","year":"2014","journal-title":"Journal of Neuroscience"},{"key":"e_1_2_5_5_1","volume-title":"Somatosensory research methods","author":"Ackerley R.","year":"2022"},{"issue":"4","key":"e_1_2_5_6_1","doi-asserted-by":"crossref","first-page":"1885","DOI":"10.1152\/jn.00940.2017","article-title":"Differential effects of radiant and mechanically applied thermal stimuli on human C\u2010tactile afferent firing patterns","volume":"120","author":"Ackerley R.","year":"2018","journal-title":"Journal of Neurophysiology"},{"issue":"5","key":"e_1_2_5_7_1","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0281253","article-title":"Hold me or stroke me? 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