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Touch plays an important role in the development of infants and children. In this review, we highlight the neural conditioning of affective touch and the related physiological responses, especially in the form of parasympathetic activation, pain suppression and stress relief. Based on recent studies, we show that the functionality of a population of C-tactile (CT) nerve fibers, hypothesized to provide the neurobiological substrate for the transmission of interpersonal touch, is already mature in the newborn and that the developmentally beneficial effects of such touch are already evident at this stage. We further aim to shed light on the role of nurturing touch in stimulating CT fibers. Based on this, we hypothesize that CT stimulation acts as a reinforcer that contributes to the development of newborns into social beings by means of conditioning.

The 'Virtual Touch Toolkit' is the exergame bundle of a newly developed smartphone application for mental health. The primary aim of the exercises is to promote self-discovery by focusing on social, affective touch. The necessity of acknowledging the undervalued properties of affective touch in this day and age is more important than ever as a global pandemic is provoking societal changes that are predicted to be detrimental to our mutual need for affective touch. The toolkit comprises multiple features and sets of exercises, which are designed to stimulate the user's oxytocinergic and reward systems while offering educational insights on affective touch.

This study explored whether a human-like feel of touch biases perceived pleasantness and whether such a bias depends on top-down cognitive and/or bottom-up sensory processes. In 2 experiments, 11 materials were stroked across the forearm at different velocities (bottom-up) and participants rated tactile pleasantness and humanness. Additionally, in Experiment 1, participants identified the materials (top-down), whereas in Experiment 2, they rated each material with respect to its somatosensory properties (bottom-up). Stroking felt most pleasant at velocities optimal for the stimulation of CT-afferents, a mechanosensory nerve hypothesized to underpin affective touch. A corresponding effect on perceived humanness was significant in Experiment 1 and marginal in Experiment 2. Whereas material identification was unrelated to both pleasantness and humanness, we observed a robust relation with the somatosensory properties. Materials perceived as smooth, slippery, and soft were also pleasant. A corresponding effect on perceived humanness was significant for the first somatosensory property only. Humanness positively predicted pleasantness and neither top-down nor bottom-up factors altered this relationship. Thus, perceiving gentle touch as human appears to promote pleasure possibly because this serves to reinforce interpersonal contact as a means for creating and maintaining social bonds. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

The sense of touch codes the detection and properties of physical objects on the body via mechanoreceptors within the skin. Technological advancements, such as ultrasonic haptic devices, are now able to ‘touch without touching’, claiming this is virtual touch. An initial aim of the study was to investigate subjective intensity and pleasantness ratings of ultrasound stimulation and the influence of top-down factors using the Touch Experience and Attitudes Questionnaire (TEAQ). Self-reported intensity and pleasantness ratings were measured in response to ultrasound stimuli. A second aim was to record from individual low threshold mechanoreceptors using the technique of microneurography in an attempt to determine which mechanoreceptors are activated by ultrasound stimulation of the skin. The major findings here were that microneurography found SAI and SAII units did not respond to ultrasound stimuli; intensity and pleasantness ratings were significantly different between age groups. Ultrasound can produce a variety of sensations with varying intensity and pleasantness ratings. A limitation of the study was the unexpected force difference generated between modulations. These findings have implications for mid-air haptics, somatosensory affective research, and virtual reality. Future research should focus on microneurography investigation of FA fibre responses to ultrasound.

Current theory divides the human mechanical sense into discriminative and affective systems. A discriminative system supports tactile exploration and manipulation via fast Aß signaling, whereas an affective system supports the pleasure of friendly interpersonal touch via slow CT signaling. To probe this system segregation, we recorded the electroencephalogram from participants being stroked and reporting stroke pleasantness. We observed a somatosensory negativity that was maximal for CT optimal as compared with sub-optimal velocities, that predicted subjective pleasantness, and showed only for stroking of hairy skin known to be CT innervated. Importantly, the latency of this negativity preceded C fiber input to the brain by several hundred milliseconds and is best explained by interactions between CT and Aß processes in the spinal cord. Our data challenge the divide between discriminative and affective touch implying instead that both fast Aß and slow CT signaling play an important role in tactile pleasure.

Pain is a universal, multidimensional experience with sensory, emotional, cognitive, and social components, which is fundamental to our environmental learning when functioning typically. Understanding pain processing in psychiatric conditions could provide unique insight into the underlying pathophysiology or psychiatric disease, especially given the psychobiological overlap with pain processing pathways. Studying pain in psychiatric conditions is likely to provide important insights, yet, there is a limited understanding beyond the work in depression and anxiety. This is a missed opportunity to describe psychiatric conditions in terms of neurobiological alterations. To examine the research into the pain experiences of these groups and the extent to which a-typicality is present, a systematic review was conducted. An electronic search strategy was developed and conducted in several databases. The current systematic review included 46 studies covering five Diagnostic and Statistical Manual of Mental Disorders (5th ed.; DSM-5) disorders: autism, attention-deficit hyperactivity disorder (ADHD), schizophrenia, personality disorder, and eating disorders, confirming tentative evidence of altered pain and touch processing. Specifically, hyposensitivity is reported in schizophrenia, personality disorder and eating disorder, hypersensitivity in ADHD, and mixed results for autism. Review of the research highlights a degree of methodological inconsistency in the utilization of comprehensive protocols, the lack of which fails to allow us to understand whether a-typicality is systemic or modality specific.

Affective sharing is a bottom-up process involving automatic processing of sensory inputs that facilitate vicarious experience of anothers emotional state. It is grounded directly in the prior experiences of the perceiver. In adults, vicarious ratings of affective touch match the known velocity tuning and hypothesised anatomical distribution of C-tactile afferents (CT), a subclass of C-fibre which respond preferentially to low force/velocity stroking touch, typically perceived as pleasant. Given the centrality of touch to early nurturing interactions, here we examined whether primary school aged childrens vicarious ratings of affective touch show the same anatomical and velocity specific patterns reported in adults. Forty-four children aged between 8 and 11 (mean age 9, 24 male) rated a sequence of video clips depicting one individual being touched by another on 5 different upper-body sites (palm, dorsal forearm, ventral forearm, upper-arm and back) at 3 different velocities (static, CT optimal, slow stroking and non-CT optimal, fast stroking). Immediately after viewing each clip, participants were asked to rate how pleasant they perceived the touch to be. While children rated the CT optimal velocity significantly higher than static or non-CT optimal touch, unlike adults their ratings did not vary across skin sites. This difference may reflect the fact childrens ratings are grounded in bottom-up affective resonance while adults also draw on top-down cognitive evaluation of the broader social context when rating the stimuli. Funding Agencies|Leverhulme TrustLeverhulme Trust [RPG-2013-058]; Liverpool John Moores University

Tactile sensitivities are common in Autism Spectrum Conditions (autism). Psychophysically, slow, gentle stroking touch is typically rated as more pleasant than faster or slower touch. Vicarious ratings of social touch results in a similar pattern of velocity dependent hedonic ratings as directly felt touch. Here we investigated whether adults and childrens vicarious ratings vary according to autism diagnosis and self-reported autistic traits. Adults scoring high on the AQ rated stroking touch on the palm as less pleasant than a Low AQ group. However, in contrast to our hypothesis, we did not find any effect of autism diagnosis on childrens touch ratings despite parental reports highlighting significant somatosensory sensitivities. These results are discussed in terms of underpinning sensory and cognitive factors. Funding Agencies|Linkoping University; Leverhulme TrustLeverhulme Trust [RPG-2013-058]; Liverpool John Moores University PhD studentship

Affective touch refers to the emotional and motivational facets of tactile sensation and has been linked to the activation of a specialised system of mechanosensory afferents (the CT system), that respond optimally to slow caress-like touch. Affective touch has been shown to play an important role in the building of the bodily self: the multisensory integrated global awareness of one’s own body. Here we investigated the effects of affective touch on subsequent tactile awareness and multisensory integration using the Somatic Signal Detection Task (SSDT). During the SSDT, participants were required to detect near-threshold tactile stimulation on their cheek, in the presence/absence of a concomitant light. Participants repeated the SSDT twice, before and after receiving a touch manipulation. Participants were divided into two groups: one received affective touch (CT optimal; n = 32), and the second received non-affective touch (non-CT optimal; n = 34). Levels of arousal (skin conductance levels, SCLs) and mood changes after the touch manipulation were also measured. Affective touch led to an increase in tactile accuracy, as indicated by less false reports of touch and a trend towards higher tactile sensitivity during the subsequent SSDT. Conversely, non-affective touch was found to induce a partial decrease in the correct detection of touch possibly due to a desensitization of skin mechanoreceptors. Both affective and non-affective touch induced a more positive mood and higher SCLs in participants. The increase in SCLs was greater after affective touch. We conclude that receiving affective touch enhances the sense of bodily self therefore increasing perceptual accuracy and awareness. Higher SCLs are suggested to be a possible mediator linking affective touch to a greater tactile accuracy. Clinical implications are discussed.

Recent evidence suggests that altered responses to affective touch - a pleasant interoceptive stimulus associated with activation of the C-Tactile (CT) system, may contribute to the aetiology and maintenance of mental conditions characterised by body image disturbances (e.g., Anorexia Nervosa). Here, we investigated whether tactile pleasantness and intensity differ across body sites, and if individual differences in dysmorphic appearance concerns and body awareness might be associated to touch perceptions across body sites. To this end, we measured perceived pleasantness and intensity of gentle, dynamic stroking touches applied to the palm, forearm, face, abdomen and back of 30 female participants (mean age: 25.87±1.17yrs) using CT-optimal (3 cm/s) and non-CT optimal (0.3 and 30 cm/s) stroking touch. As expected, participants rated CT-targeted touch as more pleasant compared to the two non-CT optimal stroking touch at all body sites. Nevertheless, CT-targeted touch applied to the abdomen elicited the lowest pleasantness ratings compared to all other body sites and to the two non-CT optimal stroking touch. Individual differences in body awareness and dysmorphic concerns significantly predicted preference for CT-optimal over non-CT optimal stroking touch applied to the forearm and the back. These findings begin to elucidate the link between CT sensitivity, dysmorphic appearance concerns and body awareness, which may have implications for future research looking to inform early interventions. Addressing impaired processing of affective interoceptive stimuli, such as CT-targeted touch, may be the key to current treatment approaches available for those populations at risk of disorders characterised by body image disturbance.

In the United Kingdom, the most common reasons for a child to come under the care of social services are neglect and abuse. Such early childhood adversity is a risk factor for social-isolation and poor mental health in adulthood. Touch is a key channel for nurturing interactions, and previous studies have shown links between early somatosensory input, experience dependent neural plasticity, and later life emotional functioning. The aim of the present study was to test the relationship between childhood neglect/abuse and later life experiences, attitudes, and hedonic ratings of affective touch. Here, affective touch is defined as low force, dynamic touch which C-Tactile afferents (CTs) respond optimally to. We hypothesized that a childhood lacking in early nurturing tactile stimulation would be associated with reduced sensitivity to socially relevant affective touch in adulthood. To test this, 19 care leavers (average 9.32 ± 3.70 years in foster care) and 32 non-care leavers were recruited through opportunity sampling (mean age = 21.25 ± 1.74 years). Participants completed a range of psychophysical somatosensory tests. First, they rated the pleasantness of CT-optimal (3 cm/s) and non-CT-optimal (0.3 and 30 cm/s) stroking touch applied to their forearm, both robotically and by an experimenter. They also made vicarious ratings of the anticipated pleasantness of social tactile interactions depicted in a series of videos. Finally, they filled in the Childhood Trauma Questionnaire (CTQ) and the Touch Experiences and Attitudes Questionnaire (TEAQ). As expected, care leavers reported significantly higher levels of childhood trauma than the control group. They also reported significantly lower levels of positive childhood touch compared to non-care leavers, but their attitudes and experiences of current intimate and affiliative touch did not differ. Across all psychophysical tests, care leavers showed specific reduction in sensitivity to the affective value of CT targeted 3 cm/s touch. The results of this study support the hypothesis that a lack of nurturing touch in early developmental periods leads to blunted sensitivity to the specific social value of affective touch. Future research should investigate the neural and physiological mechanisms underlying the observed effect.

C-tactile afferents are hypothesized to form a distinct peripheral channel that encodes the affective nature of touch. Prevailing views indicate they project, as with other unmyelinated afferents, in lamina I-spinothalamic pathways that relay homeostatically relevant information from the body toward cortical regions involved in interoceptive processing. However, in a recent study, we found that spinothalamic ablation in humans, while profoundly impairing the canonical spinothalamic modalities of pain, temperature, and itch, had no effect on benchmark psychophysical affective touch metrics. These novel findings appear to indicate that perceptual judgments about the affective nature of touch pleasantness do not depend on the integrity of the lamina I-spinothalamic tract. In this commentary, we further discuss the implications of these unexpected findings. Intuitively, they suggest that signaling of emotionally relevant C-tactile mediated touch occurs in an alternative ascending pathway. However, we also argue that the deficits seen following interruption of a putative C-tactile lamina I-spinothalamic relay might be barely perceptible—a feature that would underline the importance of the C-tactile afferent in neurodevelopment.

Behaviors as ubiquitous as play, found throughout the animal kingdom, have clearly evolved to serve an evolutionary benefit to survival. From the perspective of a neuroethologist we can best understand such behaviors by teasing out the underlying neural mechanisms that drive them, and here we find that one sense in particular may well be playing an important role in the type of social bonding that play provides – the sense of touch. Recent research has identified a special class of touch nerves in the skin of the body, called the c-tactile afferent (CT), that code for the affective and emotional properties of touch, such as that experienced during play behaviors. The initial interest in CTs had focused on their role during early life nurturing care where tactile neglect is known to have a devastating effect on neurodevelopment. However, it appears there may be a second critical stage in brain development where CTs may also play a critical role – the play behaviors we see during adolescence.

We investigated the effects of non-informative vision of the body on exteroceptive multisensory integration and touch perception in participants presenting with different levels of eating disorder (ED) symptoms. The study employed a sample of women reporting low (low ED; n = 31) vs high (high ED; n = 34) levels of subclinical ED symptoms who undertook the Somatic Signal Detection task (SSDT). During the SSDT, participants are required to detect near-threshold tactile stimulation at their fingertip with and without a simultaneous light flash next to the stimulated fingertip. Previous research has found that participants have a tendency to erroneously report touch sensations in the absence of the stimulation, and especially when the light flash is presented. In this study, participants completed the SSDT under two conditions: while their hand was visible (non-informative vision), and while their hand was hidden from sight (no vision). Non-informative vision of the hand was found to have a different effect on SSDT performances according to participants’ levels of ED symptoms. High ED participants were better able to correctly detect the touch during the SSDT when their hand was visible. Conversely, for low ED participants, vision of the body was linked to a greater effect of the light in inducing false reports of touch. We suggest that in those with high ED symptoms, vision of the body may exacerbate a predisposition to focusing on external rather than internal bodily information.

During infancy relational experiences of body-to-body exchanges (i.e., embodied interactions) contribute to the infant's bodily perception. Early embodied interactions are based on countless multimodal reciprocal exchanges, in which mother and infant contribute to interpersonal rhythmic cycles of co-regulation (i.e., attunement). However, it remains unclear how infants and their mothers actually accomplish attunement in their exchanges. Interactions between mothers and their infants typically fluctuate between attuned and misattuned states and recovery attunement states by a process called 'reparation'. Here, we discuss recent neuroscientific evidence that provides insight into the mechanisms underpinning the concepts of attunement and misattunement in early embodied interactions. We propose that a process of embodied reparation might be achieved within the dyad through tactile contact behaviors (e.g., skin-to-skin, affectionate touch) and maternal interoceptive sensitivity (i.e., ability to perceive internal input about the state of one's own body). We describe how these elements that mothers provide during embodied interactions with their infants, might contribute not only to bodily attunement, but also to co-create the infant bodily-self.

Despite growing interest in the beneficial effects of positive touch experiences throughout our lives, and individual differences in how these experiences are perceived, there is not yet available a contemporary self-report measure of touch experiences and attitudes, for which the factor structure has been validated. This article describes four studies carried out during the construction and validation of the Touch Experiences and Attitudes Questionnaire (TEAQ). The original TEAQ, containing 117 items relating to positive touch experiences was systematically constructed. Principal component analysis reduced this measure to 57 items and identified six components relating to touch experiences during childhood and adult experiences relating to current intimate touch and touch with friends and family. Three attitudinal components were identified relating to attitude to intimate touch, touch with unfamiliar people, and self-care. The structure of this questionnaire was confirmed through confirmatory factor analysis carried out on data obtained from a second sample. Good concurrent and predictive validity of the TEAQ compared to other physical touch measures currently available was identified. Known-group validity in terms of gender, marital status and age was determined, with expected group differences identified. This study demonstrates the TEAQ to have good face validity, internal consistency, construct validity in terms of discriminant validity, known-group validity and convergent validity, and criterion-related validity in terms of predictive validity and concurrent validity. We anticipate this questionnaire will be a valuable tool for the field of physical touch research. Electronic supplementary material The online version of this article (10.1007/s10919-018-0281-8) contains supplementary material, which is available to authorized users.

We examined whether alterations in body perception in EDs extend to the integration of exteroceptive visual and tactile information. Moreover, we investigated the effect of self-focused attention on the ability to correctly detect tactile stimuli. Twenty-seven women reporting low ED symptoms, versus 26 women reporting high ED symptoms, undertook a modified version of the Somatic Signal Detection Task (SSDT), which involved detecting tactile stimuli on the cheek in the presence or absence of a concomitant light. The SSDT was completed while looking at a photograph of one’s own face, another female face, and a scrambled face. Heart rate and skin conductance were recorded continuously during the SSDT. Although ED symptoms were not associated with an overall increased tendency to misperceive touch in the presence of a light, High ED participants were differentially affected by self-focused attention. For the High ED group, physiological arousal, and tactile sensitivity (d′) were increased when self-focused attention was augmented. For the Low ED group, sensitivity (d′) and physiological arousal were higher in the control conditions. We suggest that in those with High ED symptoms, attention to the bodily self may exacerbate a predisposition to focusing on external rather than internal bodily information.

Recent evidence suggests that altered responses to affective touch—a pleasant interoceptive stimulus associated with activation of the C-Tactile (CT) system—may contribute to the aetiology and maintenance of mental conditions characterised by body image disturbances (e.g., Anorexia Nervosa). Here, we investigated whether tactile pleasantness and intensity differ across body sites, and if individual differences in dysmorphic appearance concerns and body and emotional awareness might be associated with touch perceptions across body sites. To this end, we measured perceived pleasantness and intensity of gentle, dynamic stroking touches applied to the palm, forearm, face, abdomen and back of 30 female participants (mean age: 25.87±1.17yrs) using CT-optimal (3 cm/s) and non-CT optimal (0.3 and 30 cm/s) stroking touch. As expected, participants rated CT-targeted touch as more pleasant compared to the two non-CT optimal stroking touch at all body sites. Regardless of stroking velocity, touch applied to the abdomen elicited the lowest pleasantness ratings. Lower levels of emotional awareness, greater levels of interoceptive sensibility and of dysmorphic concerns were associated with lower preference for CT-optimal stroking touch applied to the forearm and the back. These findings begin to elucidate the link between CT sensitivity, dysmorphic appearance concerns and body and emotional awareness, which may have implications for future research looking to inform early interventions. Addressing impaired processing of affective interoceptive stimuli, such as CT-targeted touch, may be the key to current treatment approaches available for those populations at risk of disorders characterised by body image disturbance.

Low intensity, non-noxious, stimulation of cutaneous somatosensory nerves has been shown to trigger oxytocin release and is associated with increased social motivation, plus reduced physiological and behavioural reactivity to stressors. However, to date, little attention has been paid to the specific nature of the mechanosensory nerves which mediate these effects. In recent years, the neuroscientific study of human skin nerves (microneurography studies on single peripheral nerve fibres) has led to the identification and characterisation of a class of touch sensitive nerve fibres named C-tactile afferents. Neither itch nor pain receptive, these unmyelinated, low threshold mechanoreceptors, found only in hairy skin, respond optimally to low force/velocity stroking touch. Notably, the speed of stroking which c-tactile afferents fire most strongly to is also that which people perceive to be most pleasant. The social touch hypothesis posits that this system of nerves has evolved in mammals to signal the rewarding value of physical contact in nurturing and social interactions. In support of this hypothesis, in this paper we review the evidence that cutaneous stimulation directly targeted to optimally activate c-tactile afferents reduces physiological arousal, carries a positive affective value and, under healthy conditions, inhibits responses to painful stimuli. These effects mirror those, we also review, which have been reported following endogenous release and exogenous administration of oxytocin. Taken together this suggests C-tactile afferent stimulation may mediate oxytocin release during affiliative tactile interactions.

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.

C-tactile afferents form a distinct channel that encodes pleasant tactile stimulation. Prevailing views indicate they project, as with other unmyelinated afferents, in lamina I-spinothalamic pathways. However, we found that spinothalamic ablation in humans, whilst profoundly impairing pain, temperature and itch, had no effect on pleasant touch perception. Only discriminative touch deficits were seen. These findings preclude privileged C-tactile-lamina I-spinothalamic projections and imply integrated hedonic and discriminative spinal processing from the body.

Negative, painful, somatosensory stimulation lengthens the perceived duration of time. However, to date, no research has explored the influence of positive, pleasant, somatosensory stimulation on temporal perception. Here we asked whether gentle stroking touch influences perceptions of duration. Pleasant (gentle) and mildly unpleasant (rough) tactile stimulation was delivered whilst participants estimated the duration of a neutral visual stimulus. Pleasant touch resulted in shorter estimates of duration than unpleasant touch. There was no difference in duration perception in the unpleasant and control conditions. Taken together with the results of previous research (Ogden, Moore, Redfern, & McGlone, 2015), the results of this study suggest that pleasant and painful somatosensory stimulation have opposing effects on temporal perception, and additionally that pleasant touch can alter aspects of perceptual and attentional processing outside the purely affective domain.

Theoretical models of time perception suggest a simple bottom-up relationship between physiological arousal and perceived duration. Increases in physiological arousal lengthen the perceived duration of events whereas decreases in physiological arousal reduce them. Whilst this relationship has been demonstrated for highly arousing negatively valenced stimuli, it has not been demonstrated for other classes of distorting stimuli (e.g. positively valenced or low arousal stimuli). The current study tested the effect of valence (positive and negative) and arousal level (high and low) on the relationship between physiological arousal and perceived duration. Sympathetic nervous system (SNS) and parasympathetic nervous system (PSNS) activity was measured during a verbal estimation task in which participants judged the duration of high and low arousal, positive, negative and neutrally valenced IAPS images. SNS and PSNS activity were indexed by measuring Pre-Ejection Period (PEP) and High Frequency Heart-rate Variability (HF-HRV) respectively. SNS reactivity was predicative of perceived duration, but only for high arousal negatively valenced stimuli, with decreases in PEP being associated with longer duration estimates. SNS and PSNS activity was not predictive of perceived duration for the low arousal negative stimuli or the low and high arousal positive stimuli. We therefore propose a new model suggesting that emotional distortions to time result from a combination of bottom-up (physiological arousal) and top-down (threat detection) factors.

Theories of human temporal perception suggest that changes in physiological arousal distort the perceived duration of events. Behavioural manipulations of sympathetic nervous system (SNS) activity support this suggestion, however the effects of behavioural manipulations of parasympathetic (PSNS) activity on time perception are unclear. The current study examined the effect of a paced respiration exercise known to increase PSNS activity on sub-second duration estimates. Participants estimated the duration of negatively and neutrally valenced images following a period of normal and paced breathing. PSNS and SNS activity were indexed by high-frequency heart-rate variability and pre-ejection period respectively. Paced breathing increased PSNS activity and reduced the perceived duration of the negative and neutrally valenced stimuli relative to normal breathing. The results show that manipulations of PSNS activity can distort time in the absence of a change in SNS activity. They also suggest that activities which increase PSNS activity may be effective in reducing the perceived duration of short events.

Observers can simulate aspects of other people's tactile experiences. We asked whether they do so when faced with full-body social interactions, whether emerging representations go beyond basic sensorimotor mirroring, and whether they depend on processing goals and inclinations. In an EEG/ERP study, we presented line-drawn, dyadic interactions with and without affectionate touch. In an explicit and an implicit task, participants categorized images into touch versus no-touch and same versus opposite sex interactions, respectively. Modulations of central Rolandic rhythms implied that affectionate touch displays engaged sensorimotor mechanisms. Additionally, the late positive potential (LPP) being larger for images with as compared to without touch pointed to an involvement of higher order socio-affective mechanisms. Task and sex modulated touch perception. Sensorimotor responding, indexed by Rolandic rhythms, was fairly independent of the task but appeared less effortful in women than in men. Touch induced socio-affective responding, indexed by the LPP, declined from explicit to implicit processing in women and disappeared in men. In sum, this study provides first evidence that vicarious touch from full-body social interactions entails shared sensorimotor as well as socio-affective experiences. Yet, mental representations of touch at a socio-affective level are more likely when touch is goal relevant and observers are female. Together, these results outline the conditions under which touch in visual media may be usefully employed to socially engage observers.

It has been demonstrated that nurturing and affiliative touch is essential for human emotional and physical well-being throughout our entire life. Within the last 30 years a system of low-threshold mechanosensitive C fibers innervating the hairy skin was discovered and described; this system is hypothesized to represent the neurobiological substrate for the affective and rewarding properties of touch. This discovery opens new perspectives for multidisciplinary research of the role of affiliative social touch in health and disease, and calls for establishing novel psychometric tools assessing individual differences in the domain of affective touch. The main objective of the study was to construct and validate a Russian version of the Touch Experiences and Attitudes Questionnaire (TEAQ), a self-report measure recently developed to quantify individual experience and attitude to social and affective touch. A pool of 117 items was translated into Russian and all the items were assessed for appropriateness for Russian culture (232 participants). After exploring the factor structure (468 participants), we composed a 37-item questionnaire (TEAQ-37 Rus) characterized by good reliability and a clear 5-factor structure, covering the aspects of attitude to intimate touch, attitude to friendly touch, attitude to self-care, current intimate touch experiences, and childhood touch experiences. Confirmatory factor analysis (551 participants) has demonstrated good consistency and reliability of the 5-factor structure of the TEAQ-37 Rus. Cross-validation research demonstrated moderate positive correlations between predisposition to social touch and emotional intelligence; positive correlations with extraversion and openness facets of the Big Five personality model were also found. As predicted, participants with higher TEAQ-37 Rus scores rated all observed kinds of touch as more pleasant, with a particular preference for slow touch. We anticipate that this questionnaire will be a valuable tool for researchers of social touch, touch perception abnormalities, and the importance of touch experiences for emotional and mental health.

Social touch is a powerful force in human development, shaping social reward, attachment, cognitive, communication, and emotional regulation from infancy and throughout life. In this review, we consider the question of how social touch is defined from both bottom-up and top-down perspectives. In the former category, there is a clear role for the C-touch (CT) system, which constitutes a unique submodality that mediates affective touch and contrasts with discriminative touch. Top-down factors such as culture, personal relationships, setting, gender, and other contextual influences are also important in defining and interpreting social touch. The critical role of social touch throughout the lifespan is considered, with special attention to infancy and young childhood, a time during which social touch and its neural, behavioral, and physiological contingencies contribute to reinforcement-based learning and impact a variety of developmental trajectories. Finally, the role of social touch in an example of disordered development –autism spectrum disorder—is reviewed. Keywords: Touch, Tactile, Social, Development, Affective, Discriminative, Autism, CT afferents

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 1-10 cm/sec. 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 (3cm/sec) and two sub-optimal velocities (0.3cm/sec & 30cm/sec). PD patients underwent quantification of C-fibre density using skin biopsies and corneal confocal microscopy. All participants rated stroking velocity of 3cm/sec as the most pleasant with significantly lower ratings for 0.3cm/sec and 30cm/sec. There was a significant positive correlation between C-fibre density and pleasantness ratings at 3cm/sec and 30cm/sec but not 0.3cm/sec. 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 correlate 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. This article is protected by copyright. All rights reserved.

Highlights • We propose an intra-neural microstimulation system for 7 T fMRI and MEG. • This custom-built system removes issues with existing equipment. • It provides efficient work-flow and improved participant comfort and safety. • Stimulating single mechanoreceptors evokes activity in 7 T fMRI and MEG. • Responses to unitary stimulation are shown for the first time in MEG., Background Intra-neural microstimulation (INMS) is a technique that allows the precise delivery of low-current electrical pulses into human peripheral nerves. Single unit INMS can be used to stimulate individual afferent nerve fibres during microneurography. Combining this with neuroimaging allows the unique monitoring of central nervous system activation in response to unitary, controlled tactile input, with functional magnetic resonance imaging (fMRI) providing exquisite spatial localisation of brain activity and magnetoencephalography (MEG) high temporal resolution. New method INMS systems suitable for use within electrophysiology laboratories have been available for many years. We describe an INMS system specifically designed to provide compatibility with both ultra-high field (7 T) fMRI and MEG. Numerous technical and safety issues are addressed. The system is fully analogue, allowing for arbitrary frequency and amplitude INMS stimulation. Results Unitary recordings obtained within both the MRI and MEG screened-room environments are comparable with those obtained in ‘clean’ electrophysiology recording environments. Single unit INMS (current

The multimodal properties of the human somatosensory system continue to be unravelled. There is mounting evidence that one of these submodalities—touch—has another dimension, providing not only its well-recognized discriminative input to the brain, but also an affective input. It has long been recognized that touch plays an important role in many forms of social communication and a number of theories have been proposed to explain observations and beliefs about the “power of touch.” Here, we propose that a class of low-threshold mechanosensitive C fibers that innervate the hairy skin represent the neurobiological substrate for the affective and rewarding properties of touch.

BACKGROUND/STUDY CONTEXT: Sensory function degrades with age, with well-established reductions in tactile spatial acuity, vibrotactile sensitivity, and thermosensation, to name but three aspects of perception. Such age-related losses might be partially stemmed by ongoing experience with tasks requiring high levels of manual dexterity or analogous tactile expertise; individuals who are highly expert in skills that have a fundamental tactile component can show improved tactile function as compared with nonexperts.Eighty individuals (17 males, 63 females) in the 18-58 age range were assessed on their tactile experience, as measured by self-assessment on a variety of tasks and competencies, each of which required a high level of skill with the hands. Tactile sensory performance, manual dexterity ("haptic efficiency"), and the subjective response to tactile stimulation were quantified.Degradation in tactile sensory acuity with age was confirmed, but no strong evidence was found for variations in acuity contingent on the tactile expertise of participants. In contrast to the performance measures, differences in tactile experience were associated with differences in the subjective response to touch. Greater tactile experience was associated with the provision of richer descriptions of textured materials manipulated with the digits.The range of tactile experience reported in a convenience sample of the population was apparently insufficient to preserve sensory function during aging.

The rewarding sensation of touch in affiliative interactions is hypothesized to be underpinned by a specialized system of nerve fibers called C-Tactile afferents (CTs), which respond optimally to slowly moving, gentle touch, typical of a caress. However, empirical evidence to support the theory that CTs encode socially relevant, rewarding tactile information in humans is currently limited. While in healthy participants, touch applied at CT optimal velocities (1-10cm/sec) is reliably rated as subjectively pleasant, neuronopathy patients lacking large myelinated afferents, but with intact C-fibres, report that the conscious sensation elicited by stimulation of CTs is rather vague. Given this weak perceptual impact the value of self-report measures for assessing the specific affective value of CT activating touch appears limited. Therefore, we combined subjective ratings of touch pleasantness with implicit measures of affective state (facial electromyography) and autonomic arousal (heart rate) to determine whether CT activation carries a positive affective value. We recorded the activity of two key emotion-relevant facial muscle sites (zygomaticus major-smile muscle, positive affect & corrugator supercilii-frown muscle, negative affect) while participants evaluated the pleasantness of experimenter administered stroking touch, delivered using a soft brush, at two velocities (CT optimal 3cm/sec & CT non-optimal 30cm/sec), on two skin sites (CT innervated forearm & non-CT innervated palm). On both sites, 3cm/sec stroking touch was rated as more pleasant and produced greater heart rate deceleration than 30cm/sec stimulation. However, neither self-report ratings nor heart rate responses discriminated stimulation on the CT innervated arm from stroking of the non-CT innervated palm. In contrast, significantly greater activation of the zygomaticus major (smiling muscle) was seen specifically to CT optimal, 3cm/sec, stroking on the forearm in comparison to all other stimuli. These results offer the first empirical evidence in humans that tactile stimulation that optimally activates CTs carries a positive affective valence that can be measured implicitly.

Executive functioning deficits are reported in ecstasy users. However research into mental set switching has been equivocal, with behavioural studies suggesting the function is preserved. The current study sought to address the issue of switching deficits in ecstasy users by combining behavioural performance with electrophysiological correlates (EEG). Twenty ecstasy polydrug users, 20 non-ecstasy polydrug users and 20 drug naive controls were\ud recruited. Participants completed questionnaires about their drug use, sleep quality, fluid intelligence and current mood state. Each participant completed a mental set switching task (the number-letter task) whilst EEG measures were recorded. ANOVA revealed no between group differences on performance of the task, however a regression suggested that ecstasy use was a significant predictor for performance, after controlling for cannabis use. Mixed\ud ANOVA revealed a significant effect of group on the P3, with significant differences between both drug groups and naives. There was also an interaction between electrode and\ud group on the P2 component, with ecstasy users differing from both other groups. On the P3 component the results suggest a reduction in positivity at parieto-occipital electrodes for drug users compared to controls. Furthermore a significant increase in negativity in ecstasy users compared to control groups could be observed in several occipito-parietal electrodes at an N2 component as well as observable atypicalities in early processing (P2) displayed by ecstasy users and polydrug controls. The present study provides evidence of atypical processing of attentional shifting in ecstasy and polydrug users. Deficits in this executive function could reflect cognitive inflexibility and paucity of rapid behavioural adjustment, which may be\ud problematic in real world situations.\ud \ud Keywords: Ecstasy; cannabis; executive function; stimulants; cannabis.

Summary Background The sensation of itch and the scratch response elicited by application of histamine are enhanced by itch-related visual cues in people with existing skin conditions and, to a lesser extent, in healthy controls. Objectives We tested whether visual cues alone could generate feelings of itch and provoke a scratch response in healthy volunteers. A secondary aim was to assess whether the content of some pictures evoked these sensations more effectively. Methods Thirty participants viewed static images that were either itch-related (e.g. ants, fleas or skin conditions) or neutral (e.g. butterflies or healthy skin). These were further separated by picture type into ‘skin contact’ (e.g. ants crawling on the hand or a butterfly on a finger), ‘skin response’ (e.g. scratching an insect bite or washing the hands) or ‘context only’ (e.g. viewing midges or birds flying). Results The sensation of itch was successfully generated using itch-related pictures, with higher self-reports of itch in answer to the questions ‘How itchy do you feel?’ and ‘How itchy do you think the person in the picture feels?’, compared with viewing neutral pictures (P

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 hours, participants underwent an fMRI scan, during which time CT-innervated forearm skin and CT non-innervated finger skin was stroked with 3 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. Additionally, 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 comparable 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. This article is protected by copyright. All rights reserved.

Laboratory research suggests that the processing of painful stimuli can be modulated by selective attention to a particular sensory modality. However, alternative accounts for previous findings remain possible in terms of task-switching and spatial attention effects. In the present study, we examined whether attention can be selectively directed to the pain modality in order to facilitate the processing of the sensory-discriminative aspects of painful laser heat stimuli when these alternatives were ruled out. Participants made speeded spatial discrimination responses to an unpredictable sequence of painful laser heat and visual stimuli presented on the left forearm. On each trial, a symbolic cue predicted the likely modality for the upcoming target on the majority of trials. Participants responded more rapidly when the target was presented in the expected as opposed to the unexpected modality, demonstrating that selective attention can modulate the processing of painful stimuli. These findings are discussed in relation to contemporary theories of crossmodal attention and multisensory information-processing.

Tactile perception is inhibited during movement execution, a phenomenon known as tactile suppression. Here, we investigated whether the type of movement determines whether or not this form of sensory suppression occurs. Participants performed simple reaching or exploratory movements. Tactile discrimination thresholds were calculated for vibratory stimuli delivered to participants' wrists while executing the movement, and while at rest (a tactile discrimination task, TD). We also measured discrimination performance in a same vs. different task for the explored materials during the execution of the different movements (a surface discrimination task, SD). The TD and SD tasks could either be performed singly or together, both under active movement and passive conditions. Consistent with previous results, tactile thresholds measured at rest were significantly lower than those measured during both active movement and passive touch (that is, tactile suppression was observed). Moreover, SD performance was significantly better under conditions of single-tasking, active movements, as well as exploratory movements, as compared to conditions of dual-tasking, passive movements, and reaching movements, respectively. Therefore, the present results demonstrate that when active hand movements are made with the purpose of gaining information about the surface properties of different materials an enhanced perceptual performance is observed. As such, it would appear that tactile suppression occurs for irrelevant tactual features during both reaching and exploratory movements, but not for those task-relevant features that result from action execution during tactile exploration. Taken together, then, these results support a context-dependent modulation of tactile suppression during movement execution.

Pleasant touch sensations may begin with neural coding in the periphery by specific afferents. We found that during soft brush stroking, low-threshold unmyelinated mechanoreceptors (C-tactile), but not myelinated afferents, responded most vigorously at intermediate brushing velocities (1-10 cm s(-1)), which were perceived by subjects as being the most pleasant. Our results indicate that C-tactile afferents constitute a privileged peripheral pathway for pleasant tactile stimulation that is likely to signal affiliative social body contact.

Grooming behaviours, whether directed at the self or at others, are ubiquitous within the animal kingdom—from bees to bonobos. Having evolved for hygienic purposes, it is in primates that grooming behaviours supersede their original role to one in which there is a clear social element: nurture, control of dominance relationships, facilitating group cohesion, etc. In humans, grooming has been seen traditionally as providing a functional, plus an aesthetic, benefit—we keep clean and we look good. However, there may be another factor driving these impulses to groom ourselves: one which is less overt and a consequence of grooming that actually makes us feel good as well. It is axiomatic that grooming involves touching and, as the previous chapters in this book have described, there exists in the skin of the body a population of unmyelinated mechanosensory nerves that respond optimally to precisely the kinds of touch that typify many grooming behaviours—gentle moving touch.

Somatic sensation comprises four main modalities relaying tactile, thermal, painful, or pruritic (itch) information to the central nervous system. These input channels can be further classified as sub-serving sensory functions, such as spatial and temporal discrimination, and the provision of essential information for controlling and guiding exploratory manual behaviours, or affective functions that include the provision of the subjective experience of affective or emotional pleasurable touch. Signalling in fast-conducting myelinated peripheral nerve fibres (Aβ afferents) is important for the discriminative properties of tactile sensations, whereas signalling in unmyelinated peripheral nerve fibres, C-tactile (CT) afferents seems to be important for the rewarding, emotional properties of touch. CT afferents have specific biophysical, electrophysiological, neurobiological and anatomical properties to drive the temporally delayed affective somatic system. This chapter explores step by step the differences between the discriminative and affective touch systems, from the first stage of encoding touch in the skin to the neural pathways in the brain. The below quote from Bentley (Am J Psychol 11:405–425, 1900) reiterates the complexity of the skin and the wonder in the phenomenon of somatosensation: ‘The skin is burdened with offices. One of the surprises of physiology is the revelation of the multitude of functions performed by this apparently simple organ. As a rind it is not only the container, but the warder-off, and also the go-between for the organism and its world; tegument, buckler, interagent. It is small wonder that its work is represented in mental process; that many of our most worn and useful perceptions are made up of cutaneous sensations.’

Previous functional magnetic resonance imaging studies in two rare patients, together with microneurography and psychophysical observations in healthy subjects, have demonstrated a system of mechanosensitive C-fiber tactile (CT) afferents sensitive to slowly moving stimuli. They project to the posterior insular cortex and signal pleasant aspects of touch. Importantly, CTs have not been found in the glabrous skin of the hand, yet it is commonly observed that glabrous skin touch is also perceived as pleasant. Here we asked if the brain processing of pleasant touch differs between hairy and glabrous skin by stroking the forearm and glabrous skin of the hand during positron emission tomography. The data showed that, when contrasting slow brush stroking on the forearm with slow brush stroking on the palm, there were significant activations of the posterior insular cortex and mid-anterior orbitofrontal cortex. The opposite contrast showed a significant activation of the somatosensory cortices. Although concurrent psychophysical ratings showed no differences in intensity or pleasantness ratings, a subsequent touch questionnaire in which subjects used a newly developed 'touch perception task' showed significant difference for the two body sites. Emotional descriptors received higher ratings on the forearm and sensory descriptors were rated more highly on the palm. The present findings are consistent with the hypothesis that pleasant touch from hairy skin, mediated by CT afferents, is processed in the limbic-related cortex and represents an innate non-learned process. In contrast, pleasant touch from glabrous skin, mediated by A-beta afferents, is processed in the somatosensory cortex and represents an analytical process dependent on previous tactile experiences.

Human brain imaging studies demonstrate distributed activation of limbic, paralimbic and sensory systems to food and food-associated cues. Activity in this circuit may be modulated by internal factors, such as hunger, and cognitive factors. Anticipation to eat is one such factor, which likely impacts consummatory behavior. Here, the neural substrates of food expectancy were identified in 10 healthy male participants who underwent two whole-brain functional Magnetic Resonance Imaging scans on separate days. Fasted subjects viewed images of food and scenery, in two counterbalanced states. During one condition, subjects were 'expecting' to eat right after the scan and during the other they were 'not expecting' to eat for 1 h after the scan. Food pictures compared with scenery yielded bilateral activation in visual areas as well as in the left insula and amygdala in both conditions. The left dorsolateral prefrontal cortex, hippocampus and putamen were additionally activated in the 'not expecting' condition while right orbitofrontal cortex activity was enhanced in the 'expecting' condition. These data suggest that cognitive manipulations affect the response to food cues in the prefrontal cortex, in areas involved in the planning and control of motivated behaviors, while the amygdala and insula responded equally in both conditions, consistent with a more basic role in homeostatically driven appetitive behavior.

The tactile sensitivity of the teeth, and associated periodontium, serves important sensory and motor functions. Microneurographic recordings from human periodontal ligament mechanoreceptor (PDLM) nerves, in response to tooth loading, reveal discharge patterns with sole slowly adapting (SA) II-type characteristics, highlighting the unique role of PDLMs in oral sensory processes. Here we investigate these receptors' properties, psychophysically and with neuroimaging (fMRI), in response to varying frequencies of dynamic (vibrotactile) stimulation. The finding of increased activity in primary (SI) and secondary (SII) somatosensory cortices (SI and SII) at low frequencies of stimulation (20 Hz) as compared with higher frequencies (50 and 100 Hz), shows an increased entrainment of the PDLMs at this lower frequency in line with expected SA II-type response properties. At the highest frequency (100 Hz), no significant activity was found in SI or SII, suggesting this frequency is outside the range of activity of PDLMs. An activation matrix is mapped that includes SI, SII, insular, inferior frontal gyrus, inferior parietal lobe and supplementary motor area as well as middle frontal gyrus and cerebellum. We compared the responses to tooth stimulation with those produced by identical vibrotactile stimulation of the finger. The results strongly suggest that the PDLMs play a significant role in the specification of the forces used to hold and manipulate food between teeth, and in these respects, the masticatory system appears analogous to fine finger-control mechanisms used during precision manipulation of small objects. Because fMRI reveals activations in posterior insular cortex, we also speculate that PDLMs, and SA II-type receptors in general, may be involved in one aspect of the feeling of body ownership.

The hedonic attributes of tactile stimulation are important to one's quality of life, yet they have rarely been studied scientifically. The earliest experimental investigations suggested soft and smooth materials as pleasant, those that were stiff, rough, or coarse as unpleasant. More recent studies conducted by the authors and described herein obtained ratings of pleasantness of different textured materials stroked across the skin of multiple body sites at controlled velocities and forces of application. Statistically significant interactions between materials, sites, velocities, forces and subject sex attest to the complexity of the percept. Less pleasant percepts arose from stimuli that were rougher. However, the difficulty in making further general statements regarding hedonic touch raises questions as to whether the body surface can be mapped affectively in a meaningful manner with a single stimulus and indeed whether pleasantness-to-touch can be viewed as a unidimensional construct.

Here we report two experiments that investigated the tactile perception of one's own skin (intrapersonal touch) versus the skin of other individuals (interpersonal touch). In the first experiment, thirteen female participants rated, along four perceptual attributes, the skin of their own palm and volar forearm, then that of several of the other participants. Ratings were made using visual analogue scales for perceived smoothness, softness, stickiness, and pleasantness. One's own skin was rated less pleasant than the skin of others. For both intra- and interpersonal touch, the forearm skin was rated smoother, softer, less sticky and more pleasant than the palmar skin. In the second experiment, ten pairs of female participants rated each other's palm and volar forearm skin, with the skin of the touched individual being assessed before and after the application of skin emollients that alter skin feel. As in the first experiment, the untreated skin of others was rated more pleasant than the participants' own skin, and the forearm versus palm differences were replicated. However, the emollient had generally larger effects on self-assessments than the assessments of others, and the site effect showed greater positive sensory and pleasantness increases for palm versus volar forearm. The disparate results of the two experiments suggest that attention, influenced by the ecological importance of the stimulus, is more important to assessment of touched skin than ownership of the skin or the contribution to self-touch made by the additional receptors in the passively touched skin. In both experiments, the pleasantness of touched skin was associated with the skin's perceived smoothness and softness, with weak trends toward negative associations with its perceived stickiness, consistent with prior research using inanimate surfaces (e.g., textiles and sandpapers).