Relations between tactile sensitivity of the finger, arm, and cheek skin over the lifespan showing decline only on the finger.

Touch sensitivity generally declines with age, contributing to loss of manual dexterity and tactile function. We investigated how touch changes over the lifespan, using different tests and on three body sites. We used a classical test of force detection sensitivity, where calibrated monofilaments were applied passively to the right index finger pad, forearm, and cheek. In addition, at the index, we used an active touch spatial discrimination task, developed by our group. Spatial discrimination was estimated through participants' ability to evaluate the distance between parallel bands printed on acrylic plates. Data were collected from 96 healthy women, aged 20-75 years. Force detection and tactile spatial discrimination on the index deteriorated significantly with age; however, no change was found for tactile detection on the forearm or cheek. Tactile detection on the cheek remained remarkably highly sensitive throughout life. There was a significant positive relationship between force detection and spatial discrimination on the index. Further, force detection on the forearm was significantly associated with detection on the index and cheek. Our results suggest a decrease in touch perception with age on the index finger pad, yet a preservation of tactile sensitivity in hairy skin. This opens discussion about the impact of daily activities upon the glabrous hand skin and on the function of hairs in tactile sensitivity. We highlight the need for new methods in evaluating tactile sensitivity on hairy skin.

Effects of skin moisturization on various aspects of touch showing differences with age and skin site.

The human body is encompassed by a thin layer of tissue, the skin, which is heterogenous and highly specialized to protect the body and encode interactions with the external world. There is a fundamental scientific drive to understand its function, coupled with the need to preserve skin as we age, which impacts on our physiological and psychological well-being. In the present study, we aimed to define differences in touch perception between age groups and with skin cream application. We investigated touch on the finger, the forearm and cheek in younger (20-28 years, n = 22) and older (65-75 years, n = 22) females. We measured skin hydration, touch detection, finger spatial discrimination, forearm tactile pleasantness together with electrodermal activity, and perceptual ratings about cream use, skin dryness, and cosmetic habits. Glabrous finger skin became drier and touch performance was impaired with age, but these aspects were preserved in hairy skin. Skin moisturization immediately increased hydration levels, but did not significantly change touch perception. We also found that touch appreciation increased with age. We conclude that reduced finger capacity may impact self-evaluation of the skin and that long-term skin care strategies should focus on hydrating the hand to preserve touch capacities.

Applying cosmetic oil with added aromatic compounds improves tactile sensitivity and skin properties.

Tactile sensitivity generally decreases with aging and is associated with impairments in skin properties. Products that hydrate the skin can combat touch deficits and aromatic compounds have been shown to improve skin mechanical properties. Thus, we tested a base cosmetic oil against a perfumed oil, applied to the skin of females aged 40-60 years, on tactile sensitivity and skin properties after repeated application. Tactile detection thresholds were assessed using calibrated monofilaments applied at the index finger, palm, forearm, and cheek. Spatial discrimination on the finger was assessed using pairs of plates with different inter-band spaces. These tests were performed before and after 1 month of base or perfumed oil use. We found that tactile detection thresholds and spatial discrimination improved only in perfumed oil group. A complementary immunohistological study using human skin was conducted to estimate the expression of olfactory receptor OR2A4 and elastic fiber length. Further, the expression of OR2A4 intensity and the length of elastic fibers increased significantly with oil application, where larger effects were seen with the perfumed oil. We conclude that the application of a perfumed oil may be of additional benefit and could repair, and even prevent, tactile decline with aging by ameliorating skin condition.

What are C-tactile afferents and how do they relate to "affective touch"?

Since their initial discovery in cats, low-threshold C-fiber mechanoreceptors have become a central interest of scientists studying the affective aspects of touch. Their pursuit in humans, here termed C-tactile (CT) afferents, has led to the establishment of a research field referred to as "affective touch", which is differentiated from "discriminative touch". Presently, we review these developments based on an automated semantic analysis of more than 1000 published abstracts as well as empirical evidence and the solicited opinions of leading experts in the field. Our review provides a historical perspective and update of CT research, it reflects on the meaning of "affective touch", and discusses how current insights challenge established views on the relation between CTs and affective touch. We conclude that CTs support gentle, affective touch, but that not every affective touch experience relies on CTs or must necessarily be pleasant. Moreover, we speculate that currently underappreciated aspects of CT signaling will prove relevant for the manner in which these unique fibers support how humans connect both physically and emotionally.

Stroking trajectory shapes velocity effects on pleasantness and other touch percepts.

Research has identified an inverted u-shaped relationship between the pleasantness of arm stroking and stroking velocity. However, the generalizability of this relationship is questionable as much of the work relied on the rotary tactile stimulator (RTS), which strokes skin with force varying along an arc and confounds stimulus velocity with duration. We explored how these parameters shape the subjective evaluation of touch. In Study 1, one group of participants was stroked by the RTS, while two other groups were stroked by a new robot capable of different stroking trajectories. Participants were stroked at five velocities and rated pleasantness, humanness, intensity, and roughness. In Study 2, participants were stroked by the new robot imitating the trajectory of the RTS exactly, imitating it while controlling stimulus duration, or moving linearly or ovally with both constant force and duration. Participants rated pleasantness and humanness. Although stroke velocity was related to both pleasantness and humanness in an inverted u-shaped manner, stimulus trajectory modulated this relationship and the association between velocity and the other ratings. Together, our results clearly link stroking velocity to the perception of touch but highlight that this relationship is shaped by other physical parameters including touch duration and spatial pattern. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Slowly-adapting type II afferents contribute to conscious touch sensation in humans: Evidence from single unit intraneural microstimulation.

Slowly-adapting type II (SA-II, Ruffini) mechanoreceptive afferents respond well to pressure and stretch, and are regularly encountered in human microneurography studies. Despite an understanding of SA-II response properties, their role in touch perception remains unclear. Specific roles of different myelinated Aß mechanoreceptive afferents in tactile perception have been revealed using single unit intraneural microstimulation (INMS), via microneurography, recording from and then electrically stimulating individual afferents. This method directly links single afferent artificial activation to perception, where INMS produces specific 'quantal' touch percepts associated with different mechanoreceptive afferent types. However, SA-II afferent stimulation has been ambiguous, producing inconsistent, vague sensations, or no clear percept. We physiologically characterized hundreds of individual Aß mechanoreceptive afferents in the glabrous hand skin and examined the subsequent percepts evoked by trains of low amplitude INMS current pulses (<10 µA). We present 18 SA-II afferents where INMS resulted in a clear, electrically evoked sensation of large (∼36 mm2 ) diffuse pressure, which was projected precisely to their physiologically-defined receptive field in the skin. This sensation was felt as natural, distinctive from other afferents, and showed no indications of multi-afferent stimulation. Stimulus frequency modulated sensation intensity and even brief stimuli (4 pulses, 60 ms) were perceived. These results suggest that SA-II afferents contribute to perceived tactile sensations, can signal this rapidly and precisely, and are relevant and important for computational models of touch sensation and artificial prosthetic feedback. KEY POINTS: Slowly adapting type II mechanoreceptors (SA-IIs) are primary sensory neurons in humans that respond to pressure and stretch applied to the skin. To date, no specific conscious correlate of touch has been linked to SA-II activation. Using microneurography and intraneural microstimulation to stimulate single sensory neurons in human subjects, we find a specific sensation linked to the activation of single SA-II afferents. This sensation of touch was reported as gentle pressure and subjects could detect this with a high degree of accuracy. Methods of artificial tactile sensory feedback and computational models of touch should include SA-IIs as meaningful contributors to the conscious sensation of touch.

Passive Proprioceptive Training Alters the Sensitivity of Muscle Spindles to Imposed Movements.

Humans rely on precise proprioceptive feedback from our muscles, which is important in both the acquisition and execution of movements, to perform daily activities. Somatosensory input from the body shapes motor learning through central processes, as demonstrated for tasks using the arm, under active (self-generated) and passive conditions. Presently, we investigated whether passive movement training of the ankle increased proprioceptive acuity (psychophysical experiment) and whether it changed the peripheral proprioceptive afferent signal (microneurography experiment). In the psychophysical experiment, the ankle of 32 healthy human participants was moved passively using pairs of ramp-and-hold movements in different directions. In a pretraining test, participants made judgements about the movement direction in a two-alternative forced choice paradigm. Participants then underwent passive movement training, but only half were cued for learning, where a reference position was signaled by a sound and the participant had to learn to recognize this position; they then completed a post-training test. In a paradigm using the same setup, nine healthy participants underwent microneurography recordings of Ia muscle afferents from the peroneal nerve, where all were cued during training. In the psychophysical experiment, proprioceptive acuity improved with training only in the cued group. In the microneurography experiment, we found that muscle afferent firing was modulated, via an increase in the dynamic index, after training. We suggest that changes in muscle afferent input from the periphery can contribute to and support central perceptual and motor learning, as shown under passive conditions using ankle movements, which may be exploited for movement rehabilitation.

Large Postural Sways Prevent Foot Tactile Information From Fading: Neurophysiological Evidence.

Cutaneous foot receptors are important for balance control, and their activation during quiet standing depends on the speed and the amplitude of postural oscillations. We hypothesized that the transmission of cutaneous input to the cortex is reduced during prolonged small postural sways due to receptor adaptation during continued skin compression. Central mechanisms would trigger large sways to reactivate the receptors. We compared the amplitude of positive and negative post-stimulation peaks (P50N90) somatosensory cortical potentials evoked by the electrical stimulation of the foot sole during small and large sways in 16 young adults standing still with their eyes closed. We observed greater P50N90 amplitudes during large sways compared with small sways consistent with increased cutaneous transmission during large sways. Postural oscillations computed 200 ms before large sways had smaller amplitudes than those before small sways, providing sustained compression within a small foot sole area. Cortical source analyses revealed that during this interval, the activity of the somatosensory areas decreased, whereas the activity of cortical areas engaged in motor planning (supplementary motor area, dorsolateral prefrontal cortex) increased. We concluded that large sways during quiet standing represent self-generated functional behavior aiming at releasing skin compression to reactivate mechanoreceptors. Such balance motor commands create sensory reafference that help control postural sway.

Individual Variability of Pleasantness Ratings to Stroking Touch Over Different Velocities.

Many studies have investigated the perception of tactile pleasantness over a range of stroking velocities. On average, pleasantness is low at slow (e.g. 0.3 cm/s) and fast (e.g. 30 cm/s) stroking velocities, but is rated highest at velocities between 1 and 10 cm/s. On a group level, this results in an inverted-U shape pleasantness ratings curve, which is described statistically by a negative quadratic equation. We reanalyzed the data from five earlier studies to investigate whether the inverted-U shape pleasantness curve at the group level is also present at the level of the individual, - a precondition for using tactile pleasantness perception as a diagnostic marker. We pooled the data from five studies with a total of 127 participants. Each study included a 'standard condition' of stroking on the dorsal forearm over different velocities (0.3, 1, 3, 10, 30 cm/s) and participants rated the pleasantness. Factors other than stroking velocity were also varied in these studies. On the whole-group level and in each study, pleasantness ratings produced a significant negative quadratic pleasantness curve over the stroking velocities. In individual participants, ratings varied greatly and only 42% of the participants showed a significant negative quadratic curve. The steepness of the inverted-U correlated only moderately across other experimental conditions, showing that the experimental circumstances can influence pleasantness ratings. Our findings have important implications for future work, where differences in the tactile pleasantness curve should not be used to predict or diagnose issues at an individual level.

Evidence for sparse C-tactile afferent innervation of glabrous human hand skin.

C-tactile (CT) afferents were long-believed to be lacking in humans, but these were subsequently shown to densely innervate the face and arm skin, and to a lesser extent the leg. Their firing frequency to stroking touch at different velocities has been correlated with ratings of tactile pleasantness. CT afferents were thought to be absent in human glabrous skin; however, tactile pleasantness can be perceived across the whole body, including glabrous hand skin. We used microneurography to investigate mechanoreceptive afferents in the glabrous skin of the human hand, during median and radial nerve recordings. We describe CTs found in the glabrous skin, with characteristics comparable with those in hairy arm skin, and detail recordings from three such afferents. CTs were infrequently encountered in the glabrous skin and we estimate that the ratio of recorded CTs relative to myelinated mechanoreceptors (1:80) corresponds to an absolute innervation density of around seven times lower than in hairy skin. This sparse innervation sheds light on discrepancies between psychophysical findings of touch perception on glabrous skin and hairy skin, although the role of these CT afferents in the glabrous skin remains subject to future work.NEW & NOTEWORTHY Human touch is encoded by low-threshold mechanoreceptors, including myelinated Aß afferents and unmyelinated C-tactile (CT) afferents. CTs are abundant in hairy skin and are thought to code gentle, stroking touch that signals positive affective interactions. CTs have never been described in human glabrous skin, yet we show evidence of their existence on the hand, albeit at a relatively low density. Glabrous skin CTs may provide modulatory reinforcement of gentle tactile interactions during touch using the hands.

Cutaneous warmth, but not touch, increases muscle sympathetic nerve activity during a muscle fatigue hand-grip task.

In homeostasis, somatosensory C fibre afferents are hypothesised to mediate input to the brain about interactions with external stimuli and sympathetic efference provides the output that regulates bodily functions. We aimed to test this hypothesis and whether different types of innocuous somatosensory input have differential effects. Healthy volunteers performed a muscle fatigue (hand-grip) task to exhaustion, which produces increased muscle sympathetic nerve activity (MSNA), as measured through microneurography. Participants completed the muscle fatigue task without concurrent cutaneous sensory stimulation (control) or we applied skin warming (heat pack) as a C fibre stimulation, slow brush stroking as C and Aß fibre stimulation, or vibration as Aß fibre stimulation, to the participant's forearm. We also measured heart rate, the duration of the hand-grip task, and ratings of pain at the end of the task. Concurrent skin warming showed increased MSNA compared to the other conditions. Tactile stimuli (brushing, vibration) were not significantly different to the control (no intervention) condition. Warming increased the pain from the muscle contraction, whereas the tactile stimuli did not. We interpret the effect of warming on MSNA as providing relevant afferent information during muscle contraction, which needed to be counteracted via vasoconstriction to maintain homeostasis. Brushing and vibration were less homeostatically relevant stimuli for the muscle contraction and hence had no significant effect. The findings add sensory specificity to our current understanding of homeostatic regulation through somatosensory afferent and sympathetic efferent pathways.

Spatio-temporal profile of brain activity during gentle touch investigated with magnetoencephalography.

Positive affective touch plays a central role in social and inter-personal interactions. Low-threshold mechanoreceptive afferents, including slowly-conducting C-tactile (CT) afferents found in hairy skin, transmit such signals from gentle touch to the brain. Tactile signals are processed, in part, by the posterior insula, where it is the thought to be the primary target for CTs. We used magnetoencephalography (MEG) to assess brain activity evoked by gentle, naturalistic stroking touch on the arm delivered by a new MEG-compatible brush robot. We aimed to use high temporal resolution MEG to allow us to distinguish between brain responses from fast-conducting Aß and slowly-conducting CT afferents. Brush strokes were delivered to the left upper arm and left forearm of 15 healthy participants. We hypothesized that late brain responses, due to slow CT afference, would appear with a time shift between the two different locations on the arm. Our results show that gentle touch rapidly activated somatosensory, motor, and cingulate regions within the first 100 ms of skin contact, which was driven by fast-conducting mechanoreceptive afference, and that these responses were sustained during touch. Peak latencies in the posterior insula were shifted as a function of stimulus location and temporally-separate posterior insula activations were induced by Aß and CT afference that may modulate the emotional processing of gentle touch on hairy skin. We conclude that the detailed information regarding temporal and spatial brain activity from MEG provides new insights into the central processing of gentle, naturalistic touch, which is thought to underpin affective tactile interactions.

Seeing Your Foot Move Changes Muscle Proprioceptive Feedback.

Multisensory effects are found when the input from single senses combines, and this has been well researched in the brain. Presently, we examined in humans the potential impact of visuo-proprioceptive interactions at the peripheral level, using microneurography, and compared it with a similar behavioral task. We used a paradigm where participants had either proprioceptive information only (no vision) or combined visual and proprioceptive signals (vision). We moved the foot to measure changes in the sensitivity of single muscle afferents, which can be altered by the descending fusimotor drive. Visual information interacted with proprioceptive information, where we found that for the same passive movement, the response of muscle afferents increased when the proprioceptive channel was the only source of information, as compared with when visual cues were added, regardless of the attentional level. Behaviorally, when participants looked at their foot moving, they more accurately judged differences between movement amplitudes, than in the absence of visual cues. These results impact our understanding of multisensory interactions throughout the nervous system, where the information from different senses can modify the sensitivity of peripheral receptors. This has clinical implications, where future strategies may modulate such visual signals during sensorimotor rehabilitation.

Imaging human cortical responses to intraneural microstimulation using magnetoencephalography.

The sensation of touch in the glabrous skin of the human hand is conveyed by thousands of fast-conducting mechanoreceptive afferents, which can be categorised into four distinct types. The spiking properties of these afferents in the periphery in response to varied tactile stimuli are well-characterised, but relatively little is known about the spatiotemporal properties of the neural representations of these different receptor types in the human cortex. Here, we use the novel methodological combination of single-unit intraneural microstimulation (INMS) with magnetoencephalography (MEG) to localise cortical representations of individual touch afferents in humans, by measuring the extracranial magnetic fields from neural currents. We found that by assessing the modulation of the beta (13-30 Hz) rhythm during single-unit INMS, significant changes in oscillatory amplitude occur in the contralateral primary somatosensory cortex within and across a group of fast adapting type I mechanoreceptive afferents, which corresponded well to the induced response from matched vibrotactile stimulation. Combining the spatiotemporal specificity of MEG with the selective single-unit stimulation of INMS enables the interrogation of the central representations of different aspects of tactile afferent signalling within the human cortices. The fundamental finding that single-unit INMS ERD responses are robust and consistent with natural somatosensory stimuli will permit us to more dynamically probe the central nervous system responses in humans, to address questions about the processing of touch from the different classes of mechanoreceptive afferents and the effects of varying the stimulus frequency and patterning.

Exposure shapes the perception of affective touch.

Touch is a common occurrence in our lives, where affective and inter-personal aspects of touch are important for our well-being. We investigated whether touch exposure affects hedonic and discriminative aspects of tactile perception. The perceived pleasantness and intensity of gentle forearm stroking, over different velocities, was assessed in individuals reporting to seldom receive inter-personal touch, and in controls who received touch often. The groups did not differ in their stroking intensity judgements, nor in tactile discrimination sensitivity; however, individuals with low touch exposure evaluated the pleasantness of touch differently. These individuals did not differentiate pleasantness over the stroking velocities in the same way as the control group. The pleasantness curve for the low touch exposure group was significantly flatter and they rated 3cm/s stroking as significantly less pleasant. Other physiological and questionnaire measures were obtained and the appreciation of touch from familiar persons was positively related to the pleasantness of touch in controls, but this was not found in low touch exposure individuals. This suggests that the association of human caresses from well-known individuals, with the pleasure derived, may depend on continued exposure to it.

Gentle touch perception: From early childhood to adolescence.

Affective touch plays an important role in children's social interaction and is involved in shaping the development of the social brain. The positive affective component of touch is thought to be conveyed via a group of unmyelinated, low-threshold mechanoreceptive afferents, known as C-tactile fibers that are optimally activated by gentle, slow, stroking touch. Touch targeting these C-tactile fibers has been shown to decrease the heart rate in infants. The current study investigated the relationship between age and psychophysical ratings in response to affective touch. A total of n=43 participants (early childhood: aged 5-8 years, 9 girls, 12 boys; late childhood: aged 9-12 years, 12 girls, 10 boys) were presented with C-tactile optimal and sub-optimal stroking velocities and rated touch pleasantness on an affective pictorial scale. For both age groups, we found that children preferred C-tactile-targeted stimulation. A comparison with previously published data showed that the children's preference for C-tactile-targeted stimulation was similar to those obtained in adolescents and adults. We speculate that the effect of C-tactile-targeted touch, which is linked with pleasantness, shapes the children's preference for C-tactile over non-C-tactile-targeted stimulation, and that C-tactile afferent stimulation is important for social development.

Emotions can alter kinesthetic acuity.

Kinesthesia, the perception of our own body movements, relies on the integration of proprioceptive information arising mostly from muscle spindles, which are sensory receptors in skeletal muscles. We recently showed that emotions alter the proprioceptive messages from such muscle afferents, making them more sensitive to muscle lengthening when participants were listening sad music. Presently, we investigated whether these changes in proprioceptive feedback relating to emotional state may affect the perception of limb movements. Kinesthetic acuity was tested in 20 healthy, young adults by imposing ramp-and-hold movements that consisted of either plantar flexion or dorsiflexion movements of the ankle at 0.04°/s, or no movement. These were imposed during four emotional conditions (listening to neutral, sad, or happy music, or no music). The participants were asked to relax and focus on music (or nothing), and then they shifted their focus to the direction of an incoming movement. Once this had finished, they were asked its direction. Muscle activity, heart rate, and electrodermal activity were recorded during each trial, and after each music condition the participants rated the emotion felt on a visual analog scale. The rating of the emotional content of the music corroborated with changes in physiological measures. Kinesthetic acuity was also affected by the emotional state and found to be larger during the sad condition, as compared to the no music or neutral conditions. We conclude that emotion can shape our perception of movements, which we show here where feeling sadness significantly increase our kinesthetic acuity, this may be functionally relevant for the preparation of appropriate behavioral responses.

Microneurography as a tool to study the function of individual C-fiber afferents in humans: responses from nociceptors, thermoreceptors, and mechanoreceptors.

The technique of microneurography-recording neural traffic from nerves in awake humans-has provided us with unrivaled insights into afferent and efferent processes in the peripheral nervous system for over 50 years. We review the use of microneurography to study single C-fiber afferents and provide an overview of the knowledge gained, with views to future investigations. C-fibers have slowly conducting, thin-diameter, unmyelinated axons and make up the majority of the fibers in peripheral nerves (~80%). With the use of microneurography in humans, C-fiber afferents have been differentiated into discrete subclasses that encode specific qualities of stimuli on the skin, and their functional roles have been investigated. Afferent somatosensory information provided by C-fibers underpins various positive and negative affective sensations from the periphery, including mechanical, thermal, and chemical pain (C-nociceptors), temperature (C-thermoreceptors), and positive affective aspects of touch (C-tactile afferents). Insights from microneurographic investigations have revealed the complexity of the C-fiber system, methods for delineating fundamental C-fiber populations in a translational manner, how C-fiber firing can be used to identify nerve deficits in pathological states, and how the responses from C-fibers may be modified to change sensory percepts, including decreasing pain. Understanding these processes may lead to future medical interventions to diagnose and treat C-fiber dysfunction. NEW & NOTEWORTHY The technique of microneurography allows us to directly investigate the functional roles of single C-fiber afferents in awake human beings. Here we outline and discuss the current field of C-fiber research on this heterogeneous population of afferents in healthy subjects, in pathological states, and from a translational perspective. We cover C-fibers encoding touch, temperature, and pain and provide perspectives on the future of C-fiber microneurography investigations in humans.

Differential effects of radiant and mechanically applied thermal stimuli on human C-tactile afferent firing patterns.

C-tactile (CT) afferents respond to gentle tactile stimulation, but only a handful of studies in humans and animals have investigated whether their firing is modified by temperature. We describe the effects of radiant thermal stimuli, and of stationary and very slowly moving mechanothermal stimuli, on CT afferent responses. We find that CT afferents are primarily mechanoreceptors, as they fired little during radiant thermal stimuli, but they exhibited different patterns of firing during combined mechano-cool stimulation compared with warming. CTs fired optimally to gentle, very slowly moving, or stationary mechanothermal stimuli delivered at neutral temperature (~32°C, normal skin temperature), but they responded with fewer spikes (median 67% decrease) and at significantly lower rates (47% decrease) during warm (~42°C) tactile stimuli. During cool tactile stimuli (~18°C), their mean instantaneous firing frequency significantly decreased by 35%, but they often fired a barrage of afterdischarge spikes at a low frequency (~5 Hz) that outlasted the mechanical stimulus. These effects were observed under a variety of stimulus conditions, including during stationary and slowly moving touch (0.1 cm/s), and we complemented these tactile approaches using a combined electrical-thermal stimulation experiment where we found a suppression of spiking during warming. Overall, CT afferents are exquisitely sensitive to tactile events, and we show that their firing is modulated with touch temperatures above and below neutral skin temperature. Warm touch consistently decreased their propensity to fire, whereas cool touch produced lower firing rates but afterdischarge spiking. NEW & NOTEWORTHY C-tactile (CT) afferents are thought to underpin pleasant touch, and previous work has shown that they respond optimally to a slow caress delivered at typical (neutral) skin temperature. Here, we show that, although CTs are primarily mechanoreceptive afferents, they are modified by temperature: warm touch decreases their firing, whereas cool touch produces lower firing rates but long-lasting spiking, frequently seen as afterdischarges. This has implications for the encoding of affective sensory events in human skin.