Gentle stroking elicits somatosensory ERP that differentiates between hairy and glabrous skin

Here we asked whether, similar to visual and auditory event-related potentials (ERPs), somatosensory ERPs reflect affect. Participants were stroked on hairy or glabrous skin at five stroking velocities (0.5, 1, 3, 10 and 20 cm/s). For stroking of hairy skin, pleasantness ratings related to velocity in an inverted u-shaped manner. ERPs showed a negativity at 400 ms following touch onset over somatosensory cortex contra-lateral to the stimulation site. This negativity, referred to as sN400, was larger for intermediate than for faster and slower velocities and positively predicted pleasantness ratings. For stroking of glabrous skin, pleasantness showed again an inverted u-shaped relation with velocity and, additionally, increased linearly with faster stroking. The sN400 revealed no quadratic effect and instead was larger for faster velocities. Its amplitude failed to significantly predict pleasantness. In sum, as was reported for other senses, a touch’s affective value modulates the somatosensory ERP. Notably, however, this ERP and associated subjective pleasantness dissociate between hairy and glabrous skin underscoring functional differences between the skin with which we typically receive touch and the skin with which we typically reach out to touch.     

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 3 cm/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.     

Visual perception of the arm manipulates the experienced pleasantness of touch

Touch, such as a caress, can be interpreted as very pleasant. The emotional valence assigned to touch is likely related to certain bottom-up factors, such as optimal activation of C-tactile (CT) afferents. It is however unclear if besides somatosensory input, contextual factors related to the own body also play a role in the perceived pleasantness of touch. To test this, we manipulated visual appearance of the participant’s arm (veridical vision, no vision, pixelated moving statistic projected onto the arm (i.e. crawling skin)). We used slow velocity stroking (CT optimal stroking) with a soft brush to induce pleasant touch, and fast velocity stroking as a control condition. After each visual condition we asked participants (N = 23) to rate the emotional valence of the stroking they felt. After slow velocity stroking ratings on perceived pleasantness (but not on perceived unpleasantness) were modulated by visual condition, with veridical vision of the arm resulting in higher pleasantness ratings than both no vision and pixelated vision. We conclude that contextual processes affect the perceived pleasantness of touch. These findings shed a new light on the underlying mechanisms of how humans experience pleasant touch and show that pleasant touch not solely dependents on bottom up information.     

The perception of affective touch in Parkinson's disease and its relation to small fibre neuropathy

Affective touch sensation is conducted by a sub‐class of C‐fibres in hairy skin known as C‐Tactile (CT) afferents. CT afferents respond maximally to gentle skin stroking at velocities between 1 and 10 cm/s. Parkinson's disease (PD) is characterised by markedly reduced cutaneous C‐fibres. It is not known if affective touch perception is influenced by C‐fibre density and if affective touch is impaired in PD compared to healthy controls. We predicted that perceived pleasantness to gentle stroking in PD would correlate with C‐afferent density and that affective touch perception would be impaired in PD compared to healthy controls. Twenty‐four PD patients and 27 control subjects rated the pleasantness of brush stroking at an optimum CT stimulation velocity (3 cm/s) and two sub‐optimal velocities (0.3 and 30 cm/s). PD patients underwent quantification of C‐fibre density using skin biopsies and corneal confocal microscopy. All participants rated a stroking velocity of 3 cm/s as the most pleasant with significantly lower ratings for 0.3 and 30 cm/s. There was a significant positive correlation between C‐fibre density and pleasantness ratings at 3 and 30 cm/s but not 0.3 cm/s. Mean pleasantness ratings were consistently higher in PD patients compared to control subjects across all three velocities. This study shows that perceived pleasantness to gentle touch correlates significantly with C‐fibre density in PD. The higher perceived pleasantness in PD patients compared to controls suggests central sensitisation to peripheral inputs, which may have been enhanced by dopamine therapy.     

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.     

Self-reported Pleasantness Ratings and Examiner-Coded Defensiveness in Response to Touch in Children with ASD: Effects of Stimulus Material and Bodily Location

Tactile defensiveness, characterized by behavioral hyperresponsiveness and negative emotional responses to touch, is a common manifestation of aberrant sensory processing in autism spectrum disorders (ASD) and other developmental disabilities (DD). Variations in tactile defensiveness with the properties of the stimulus and the bodily site of stimulation have been addressed in adults with self-report of perceived tactile pleasantness, but not in children. We presented three materials (pleasant, unpleasant, social) at three bodily sites and measured both examiner-coded defensiveness and self-reported pleasantness from a group of children with ASD and two comparison groups (one with DD, one with typical development (TD)). The main findings were: (1) children with ASD and DD showed significantly more defensiveness reactions and lower pleasantness ratings than the TD group, with higher variability, (2) there was a double dissociation for the effects of material and bodily site of stimulation: while bodily site predicted behavioral defensiveness, material predicted pleasantness rating. Additionally, it was noted that (3) the most pleasant material and the social touch conditions best distinguished ASD and DD from TD on defensiveness, and (4) within the ASD group, social impairment and defensiveness in bodily sites associated with social touch were positively correlated, suggesting a clinically relevant distinction between social and discriminative touch in ASD.     

Cognitive influences on the affective representation of touch and the sight of touch in the human brain

We show that the affective experience of touch and the sight of touch can be modulated by cognition, and investigate in an fMRI study where top-down cognitive modulations of bottom-up somatosensory and visual processing of touch and its affective value occur in the human brain. The cognitive modulation was produced by word labels, ‘Rich moisturizing cream’ or ‘Basic cream’, while cream was being applied to the forearm, or was seen being applied to a forearm. The subjective pleasantness and richness were modulated by the word labels, as were the fMRI activations to touch in parietal cortex area 7, the insula and ventral striatum. The cognitive labels influenced the activations to the sight of touch and also the correlations with pleasantness in the pregenual cingulate/orbitofrontal cortex and ventral striatum. Further evidence of how the orbitofrontal cortex is involved in affective aspects of touch was that touch to the forearm [which has C fiber Touch (CT) afferents sensitive to light touch] compared with touch to the glabrous skin of the hand (which does not) revealed activation in the mid-orbitofrontal cortex. This is of interest as previous studies have suggested that the CT system is important in affiliative caress-like touch between individuals.     

How the brain codes intimacy: The neurobiological substrates of romantic touch

Humans belong to a minority of mammalian species that exhibit monogamous pair‐bonds, thereby enabling biparental care of offspring. The high reward value of interpersonal closeness and touch in couples is a key proximate mechanism facilitating the maintenance of enduring romantic bonds. However, surprisingly, the neurobiological underpinnings mediating the unique experience of a romantic partner's touch remain unknown. In this randomized placebo (PLC)‐controlled, between‐group, pharmacofunctional magnetic resonance imaging (fMRI) study involving 192 healthy volunteers (96 heterosexual couples), we intranasally administered 24 IU of the hypothalamic peptide oxytocin (OXT) to either the man or the woman. Subsequently, we scanned the subjects while they assumed that they were being touched by their romantic partners or by an unfamiliar person of the opposite sex, although in reality an identical pattern of touch was always given by the same experimenter. Our results show that intranasal OXT compared to PLC selectively enhanced the subjective pleasantness of the partner's touch. Importantly, intranasal OXT selectively increased responses to partner touch in the nucleus accumbens (NAcc) and anterior cingulate cortex. Under OXT, NAcc activations to partner touch positively correlated with the subjects' evaluation of their relationship quality. Collectively, our results suggest that OXT may contribute to the maintenance of monogamous relationships in humans by concomitantly increasing the reward value of partner touch and diminishing the hedonic quality of stranger touch. Hum Brain Mapp 38:4525–4534, 2017. © 2017 Wiley Periodicals, Inc.     

Effect of Magnitude Estimation of Pleasantness and Intensity on fMRI Activation to Taste

The goal of the present study was to investigate whether the psychophysical evaluation of taste stimuli using magnitude estimation influences the pattern of cortical activation observed with neuroimaging. That is, whether different brain areas are involved in the magnitude estimation of pleasantness relative to the magnitude estimation of intensity. fMRI was utilized to examine the patterns of cortical activation involved in magnitude estimation of pleasantness and intensity during hunger in response to taste stimuli. During scanning, subjects were administered taste stimuli orally and were asked to evaluate the perceived pleasantness or intensity using the general Labeled Magnitude Scale (Green et al., Chem Senses, 21(3), 323-334, 1996; Bartoshuk et al., Physiol Behav, 82(1), 109-114, 2004). Image analysis was conducted using Analysis of Functional NeuroImage software. Magnitude estimation of intensity and pleasantness shared common activations in the insula, rolandic operculum, and the medio-dorsal nucleus of the thalamus. Globally, magnitude estimation of pleasantness produced significantly more activation than magnitude estimation of intensity. Areas differentially activated during magnitude estimation of pleasantness versus intensity included, e.g., the insula, the anterior cingulate gyrus, and putamen, suggesting that different brain areas were recruited when subjects made magnitude estimates of intensity and pleasantness. These findings demonstrate significant differences in brain activation during magnitude estimation of intensity and pleasantness to taste stimuli. An appreciation for the complexity of brain response to taste stimuli may facilitate a clearer understanding of the neural mechanisms underlying eating behavior and overconsumption.     

The affective and cognitive processing of touch, oral texture, and temperature in the brain

Some of the principles of the representation of affective touch in the brain are described. Positively affective touch and temperature are represented in parts of the orbitofrontal and pregenual cingulate cortex. The orbitofrontal cortex is implicated in some of the affective aspects of touch that may be mediated through C fibre touch afferents, in that it is activated more by light touch to the forearm (a source of C-tactile (CT) afferents) than by light touch to the glabrous skin of the hand. Oral somatosensory afferents implicated in sensing the texture of food including fat in the mouth also activate the orbitofrontal and pregenual cingulate cortex, as well as the insular taste cortex. Top-down cognitive modulation of the representation of affective touch produced by word labels is found in parietal cortex area 7, the insula and ventral striatum. The cognitive labels also influence activations to the sight of touch and also the correlations with pleasantness in the pregenual cingulate/orbitofrontal cortex and ventral striatum.     

Effects of acute tryptophan depletion on central processing of CT‐targeted and discriminatory touch in humans

C‐tactile afferents (CTs) are slowly conducting nerve fibres, present only in hairy skin. They are optimally activated by slow, gentle stroking touch, such as those experienced during a caress. CT stimulation activates affective processing brain regions, alluding to their role in affective touch perception. We tested a theory that CT‐activating touch engages the pro‐social functions of serotonin, by determining whether reducing serotonin, through acute tryptophan depletion, diminishes subjective pleasantness and affective brain responses to gentle touch. A tryptophan depleting amino acid drink was administered to 16 healthy females, with a further 14 receiving a control drink. After 4 h, participants underwent an fMRI scan, during which time CT‐innervated forearm skin and CT non‐innervated finger skin was stroked with three brushes of differing texture, at CT‐optimal force and velocity. Pleasantness ratings were obtained post scanning. The control group showed a greater response in ipsilateral orbitofrontal cortex to CT‐activating forearm touch compared to touch to the finger where CTs are absent. This differential response was not present in the tryptophan depleted group. This interaction effect was significant. In addition, control participants showed a differential primary somatosensory cortex response to brush texture applied to the finger, a purely discriminatory touch response, which was not observed in the tryptophan depleted group. This interaction effect was also significant. Pleasantness ratings were similar across treatment groups. These results implicate serotonin in the differentiation between CT‐activating and purely discriminatory touch responses. Such effects could contribute to some of the social abnormalities seen in psychiatric disorders associated with abnormal serotonin function.     

Effects of light fingertip touch on postural responses in subjects with diabetic neuropathy

OBJECTIVES: To investigate the potential benefits from lightly touching an external supporting device on automatic postural responses to support surface translations, in subjects with profound sensory neuropathy in the feet due to diabetes mellitus (DM-PN). METHODS: Eight subjects with DM-PN and 10 age matched controls were tested under randomly ordered conditions of no fingertip touch (NT), light touch (LT; <1 N), and heavy touch (HT, as needed) of a stationary touch plate, during three backward translation velocities of the support surface at 10, 20, and 30 cm/s. Dependent variables included response latencies, CoP velocity, and the slope of the relation between centre of pressure (CoP) velocity and translation velocity as a measure of response scaling. RESULTS: Postural response latencies were significantly longer and scaling of initial response magnitude in proportion to translation velocity was significantly smaller in the DM-PN subjects compared to the control subjects. LT had no significant effect on response latencies of the DM-PN patients. Fingertip touch increased the slope of the scaling of postural response magnitude in both groups. However, DM-PN subjects had to use HT to improve response scaling, whereas control subjects improved scaling with LT as well as HT. LT significantly increased rightward CoP velocity towards the touch plate in all subjects. CONCLUSIONS: LT did not reduce the latency or improve the scaling of automatic postural responses in subjects with peripheral neuropathy. The major effect of LT on the automatic postural responses of the DM-PN subjects was in increasing CoP velocity towards the side of the supporting device. HT in neuropathy subjects and LT in age matched control subjects increased the sensitivity of initial postural response scaling, suggesting that somatosensory substitution from a cane in the hand could be used to improve the magnitude of medium latency postural responses to slips and trips.     

Autistic traits modulate cortical responses to affective but not discriminative touch

The sense of touch is primarily considered a discriminative and exteroceptive sense, facilitating the detection, manipulation and exploration of objects, via an array of low‐threshold mechanoreceptors and fast conducting A‐beta (Aβ) afferents. However, a class of unmyelinated, low‐threshold mechanoreceptors identified in the hairy skin of mammals have been proposed to constitute a second, anatomically distinct system coding the affective qualities of touch. Unlike Aβs, which increase their firing rate linearly with the velocity of a stimulus moving across their receptive field, the response of these C‐tactile afferents (CTs) is described by an inverted ‘U’ curve fit, responding optimally to a skin temperature stimulus moving at between 1 and 10 cm/s. Given the distinct velocity tuning of these fast and slow touch fibres, here we used event‐related potentials to compare the time course of neural responses to 1st (fast) and 2nd (slow) touch systems. We identified a higher amplitude P300 in response to fast, Aβ‐targeted, versus slow CT‐targeted, stroking touch. In contrast, we identified a previously described, C‐fibre specific, ultra‐late potential (ULP) associated with CT‐targeted input. Of special note as regards the function of CTs is that the amplitude of the ULP was negatively correlated with self‐reported levels of autistic traits, which is consistent with the hypothesized affective and social significance of this response. Taken together, these findings provide further support for distinct discriminative and affective touch systems and suggests the temporal resolution of EEG provides an as yet underutilized tool for exploring individual differences in response sensitivity to CT‐targeted touch.     

Functional parcellation of the human primary somatosensory cortex to natural touch

Despite the significance of human touch, brain responses to interpersonal manual touch have been rarely investigated. We used functional magnetic resonance imaging to study brain activity in eight healthy adults whose left hand was touched by two individuals, in separate runs and in 20‐s blocks, either by holding, smoothing, or poking. Acceleration was measured from both the subject's and the touching person's hands for postimaging control of the stimuli. Independent component analysis of the functional magnetic resonance imaging data unraveled three functional networks involving the primary somatosensory cortex (SI). One network comprised the contralateral and another the ipsilateral Brodmann area 3. The third network included area 2 bilaterally, left‐hemisphere middle temporal gyrus and dorsolateral prefrontal regions, ventral prefrontal cortices bilaterally, and middle cingulate cortex. The response shapes and polarities varied between the three networks. The contralateral area 3 differentiated the responses between the three types of touch stimuli, and the response magnitudes depended on the variability of the touch within each block. However, the responses of the other two networks were strikingly similar to all stimuli. The subjects' reports on the pleasantness of the touch did not correlate with the characteristics of the SI responses. These findings imply area‐specific processing of the natural human touch in three networks including the SI cortex, with only area 2 connected to a functional network of brain areas that may support social interaction.     

Psychophysiological correlates of affects in human olfaction.

Pleasantness and arousal have been identified as the main dimensions of affective responses to environment. Pleasantness is defined as the degree of favorable feelings a subject can experience under given circumstances. Arousal is defined as the degree of excitement (general activation) the subject feels under these circumstances. In visual and auditory modalities, many studies using measures such as facial electromyographic (EMG) activity and skin conductance (SC) have found those parameters to vary as a function of either pleasantness or arousal: for example, facial corrugator EMG activity covaries with the pleasantness dimension, while SC increases together with arousal. The first objective of this research is to study the possible covariation between peripheral measures and pleasantness/arousal in olfaction. We also examined the effect of odor intensity on facial and autonomic variations. The second objective was to investigate whether odors could evoke verbally specific emotions (e.g. joy, anger, fear, surprise, disgust or sadness) and also induce specific patterns of peripheral responses. Participants were exposed to 12 different odors while their facial and autonomic parameters were recorded, and estimated their intensity, pleasantness, and arousal capacity. Then, they chose between seven words for emotions (fear, anger, sadness, surprise, neutral, joy or disgust) to describe their reaction to the odor. As in vision, olfactory pleasantness covaries (negatively) with facial activity of the corrugator muscle, and arousal (positively) with skin conductance. No relationships were observed between physiological changes and variations in perceived intensity. Results also showed that emotions of "disgust" and "joy" were more frequently evoked verbally than any other emotions, and that only facial EMG activity distinguishes them (e.g. "disgust" vs. "joy" and "neutral state"). The results are discussed in terms of possible existence of two brain systems (defensive and appetitive), each of them being able to vary in metabolic arousal.     

Infants discriminate the source of social touch at stroking speeds eliciting maximal firing rates in CT-fibers

The evaluation of interpersonal touch is heavily influenced by its source. For example, a gentle stroke from a loved one is generally more pleasant than the same tactile stimulation from a complete stranger. Our study tested the early ontogenetic roots of humans’ sensitivity to the source of interpersonal touch. We measured the heart rate of three groups of nine-month-olds while their legs were stroked with a brush. The participants were stroked at a different speed in each group (0.3 cm/s, 3 cm/s, 30 cm/s). Depending on the Identity condition (stranger vs. parent), the person who acted as if she was stroking the infant’s leg was either an unfamiliar experimenter or the participant’s caregiver. In fact, the stimulation was always delivered by a second experimenter blind to the Identity condition. Infants’ heart rate decreased more in reaction to strokes when their caregiver rather than a stranger acted as the source of the touch. This effect was found only for tactile stimulations whose velocity (3 cm/s) is known to elicit maximal mean firing rates in a class of afferents named C-tactile fibers (CTs). Thus, the infants’ reaction to touch is modulated not just by its mechanical properties but also by its social source.     

Genetic and Environmental Contributions to Perceived Intensity and Pleasantness of Androstenone Odor: An International Twin Study

We estimated the genetic and environmental components of variation in perceived intensity and pleasantness of androstenone, an odorous compound showing specific anosmia, by modeling twin data from Finland, Denmark, the UK, and Australia. The pooled data comprised 917 twin individuals (338 are male and 579 are female; aged from 10 to 83years) including 126 complete monozygous and 264 dizygous twin pairs as well as 137 twin individuals without their co-twin. They rated intensity and pleasantness of androstenone and citronellal (control) odors using nine categories. Additive genetic effects (heritability) contributed 28 and 21% to the variation in the perceived intensity and pleasantness of androstenone, respectively, but negligibly to variations in citronellal perception. A strong genetic correlation existed between the intensity and pleasantness of androstenone, whereas the environmental correlation was negligible. These results suggest that both intensity and pleasantness of androstenone are moderately influenced by genetic factors and that the traits are modified by an overlapping set of genes.     

Is perceived angular displacement the time integral of perceived angular velocity?

Estimates of rotational self-displacement and self-velocity have been used interchangeably in vestibular psychophysics to characterize vestibular ego-motion perception. However, the assumption underlying this indiscriminate use has never been tested. The assumption holds that the two estimates are equivalent, with the displacement estimates reflecting the time integral of the signal underlying the velocity estimate. We tested this hypothesis by directly comparing displacement and velocity estimates. Two groups of healthy young subjects (2 × N = 15) were presented with the same vestibular stimuli (horizontal whole body rotations in the dark in the form of velocity steps of 5, 10, 20, and 40°/s with 1, 2, 4, 8, and 16 s duration, yielding position ramps of 5, 10, 20, 40, 80, 160, and 320° total displacement). The first subject group estimated peak velocity, and the second group estimated total displacement, both groups using a comparable psychophysical procedure (Stevens' magnitude estimation). The experimentally obtained velocity estimates were used to predict the displacement estimates. To this end, the velocity signal was assumed to decay exponentially from the reported peak value (reflecting the dynamics of peripheral and early central vestibular mechanisms) and was mathematically integrated. Predicted and measured displacement estimates were similar when a time constant of 20 s was assumed, which is in good agreement with earlier studies. We conclude that vestibular displacement estimates can, indeed, be considered equivalent to vestibular velocity estimates, at least for the stimulus parameters used.