The influence of force on the encoding and perception of affective touch

Stroking touch has been shown to be most pleasant at intermediate velocities of 1-10 cm.s-1, which relates well to activity of C-low threshold mechanoreceptors (C-LTMRs), also called C-tactile afferents in humans, that have been implicated in encoding positive affective touch. This well-established 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-LTMRs. Presently, we investigated the perceptual pleasantness and intensity of stroking touch (0.3, 1, 3, 10, 30 cm.s-1) at different forces (0.2, 0.4, 0.8, 1.2, 1.6, 2.0 N) on hairy forearm skin and compared this to responses from C-LTMRs during stroking touch at the same velocities, but over fewer forces (0.05, 0.4, 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 shaped 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 five C-LTMRs 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, where tactile pleasantness and intensity appear to be a multi-faceted construct that are related to C-LTMR and A{beta}-LTMR activity, respectively, but that the firing in individual mechanoreceptor populations cannot fully account for percepts. Key Points SummaryO_LIThe skin is highly sensitive to small mechanical deformations, including dynamic touch and different forces. C_LIO_LIWe obtained perceptual ratings for touch pleasantness and intensity over different stroking velocities and forces applied to the forearm. This was also compared to microneurography recordings from C-low threshold mechanoreceptors (C-LTMRs). C_LIO_LIBuilding 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-LTMR firing showed that force differences were readily encoded, although the same force-velocity interaction was not seen. C_LIO_LIThis work expands the literature on affective touch to encompass the effects of force, which clearly modulates pleasantness. C_LIO_LIC-LTM force and velocity information likely combines with information from all types of mechanoreceptors, including those from the fascia, to produce the resulting percept. C_LI