Role of medial prefrontal cortex and primary somatosensory cortex in self and other-directed vicarious social touch: a TMS study.

Conflicting evidence points to the contribution of several key nodes of the 'social brain' to the processing of both discriminatory and affective qualities of interpersonal touch. Whether the primary somatosensory cortex (S1) and the medial prefrontal cortex (mPFC), two brain areas vital for tactile mirroring and affective mentalizing, play a functional role in shared representations of C-tactile (CT) targeted affective touch is still a matter of debate. Here, we used offline continuous theta-burst transcranial magnetic stimulation (cTBS) to mPFC, S1 and vertex (control) prior to participants providing ratings of vicarious touch pleasantness for self and others delivered across several body sites at CT-targeted velocities. We found that S1-cTBS led to a significant increase in touch ratings to the self, with this effect being positively associated to levels of interoceptive awareness. Conversely, mPFC-cTBS reduced pleasantness ratings for touch to another person. These effects were not specific for CT-optimal (slow) stroking velocities, but rather they applied to all types of social touch. Overall, our findings challenge the causal role of the S1 and mPFC in vicarious affective touch and suggest that self- vs other-directed vicarious touch responses might crucially depend on the specific involvement of key social networks in gentle tactile interactions.

Depression severity is associated with reduced pleasantness of observed social touch and fewer current intimate touch experiences.

Depression is associated with loss of pleasure in previously enjoyed activities and withdrawal from social interactions. Depression alters the perception of social cues, but it is currently unclear whether this extends to social touch. In the current cross-sectional study, we explored the association between depression severity, perceived pleasantness of observed social touch, and general longing for touch. For observed touch, we contrasted videos of slow touch (1-10cm/s), which optimally activates C tactile afferent nerve fibres and generally feels pleasant, with 'non-CT-optimal' touch (i.e., outside the 1-10cm/s range, commonly rated more neutral). We predicted that greater depression severity would be related to lower pleasantness ratings specifically for CT-optimal touch, and less longing for touch. N = 226 adults completed self-report measures of depression severity and longing for touch, and rated touch pleasantness for six videos depicting social touch at three velocities (3cm/s in the CT-optimal range, 0.5 and 30cm/s outside this range) and at two locations varying in CT innervation (palm vs. arm). We controlled for general anhedonia and individual differences in touch experiences and attitudes. Across touch locations, greater depression severity was associated with lower perceived pleasantness of touch, especially for the fastest non-CT-optimal (rather than the CT-optimal) velocity, contrary to our prediction. However, when grouping participants into probable vs. no/minimal depression, the probable depression group rated both the fastest non-CT-optimal and the CT-optimal velocity as less pleasant than did the no/minimal depression group. Overall, while depression was associated with perceived pleasantness of observed touch, this was not specific to CT-optimal touch. Furthermore, touch longing was not associated with depression severity. Instead, variance in depression symptoms was better explained by reduced levels of current intimate touch. Though the direction of causality is unclear, greater depression severity is related to lower pleasantness of observed social touch, and lower levels of current intimate touch.

Hold me or stroke me? Individual differences in static and dynamic affective touch.

Low-threshold mechanosensory C-fibres, C-tactile afferents (CTs), respond optimally to sensations associated with a human caress. Additionally, CT-stimulation activates brain regions associated with processing affective states. This evidence has led to the social touch hypothesis, that CTs have a key role in encoding the affective properties of social touch. Thus, to date, the affective touch literature has focussed on gentle stroking touch. However, social touch interactions involve many touch types, including static, higher force touch such as hugging and holding. This study aimed to broaden our understanding of the social touch hypothesis by investigating relative preference for static vs dynamic touch and the influence of force on these preferences. Additionally, as recent literature has highlighted individual differences in CT-touch sensitivity, this study investigated the influence of affective touch experiences and attitudes, autistic traits, depressive symptomology and perceived stress on CT-touch sensitivity. Directly experienced, robotic touch responses were obtained through a lab-based study and vicarious touch responses through an online study where participants rated affective touch videos. Individual differences were determined by self-report questionnaire measures. In general, static touch was preferred over CT-non-optimal stroking touch, however, consistent with previous reports, CT-optimal stroking (velocity 1-10 cm/s) was rated most pleasant. However, static and CT-optimal vicarious touch were rated comparably for dorsal hand touch. For all velocities, 0.4N was preferred over 0.05N and 1.5N robotic touch. Participant dynamic touch quadratic terms were calculated for robotic and vicarious touch as a proxy CT-sensitivity measure. Attitudes to intimate touch significantly predict robotic and vicarious quadratic terms, as well as vicarious static dorsal hand touch ratings. Perceived stress negatively predicted robotic static touch ratings. This study has identified individual difference predictors of CT-touch sensitivity. Additionally, it has highlighted the context dependence of affective touch responses and the need to consider static, as well as dynamic affective touch.

Translation, validity, and reliability of the European Portuguese version of the Touch Experiences and Attitudes Questionnaire.

Background: Positive touch experiences have proved to be extremely important throughout our lifespan, with cascading effects on our social life. However, few questionnaires are available to measure attitudes and experiences of touch in the Portuguese population. This study aimed to translate and validate the European Portuguese version of the Touch Experiences and Attitudes Questionnaire (TEAQ), as a reliable and valid instrument to measure different aspects of affective touch experiences and attitudes. Methods: Therefore, an online sample of 384 (299 females and 85 males) participants, aged between 18 and 75 years (M = 24.59; SD = 9.56) was collected. Multidimensional Rasch model and confirmatory factor analysis were carried out, and also reliability and convergent and discriminant validity were determined. In addition, we examined sex differences in attitudes and experiences of touch. Results: Results showed good fit indexes for the 52-item six-factor model structure (friends and family touch, current intimate touch, childhood touch, attitudes to self-care, attitudes to intimate touch, and attitudes to unfamiliar touch). This instrument also showed good reliability and acceptable convergent and discriminant validity. Significant sex differences were found, with female participants reporting more positive touch experiences (including childhood touch, friends and family touch, and current intimate touch) and a more favourable attitude to self-care, with males showing a more positive attitude towards unfamiliar touch. Regarding attitudes towards the intimate touch, scores for both groups were comparable. Conclusion: Overall, the European Portuguese version of the TEAQ presented good psychometric properties and appears to be a reliable and valid self-report measure, being a useful and beneficial instrument in research and clinical settings.

Acute tryptophan depletion alters affective touch perception.

RATIONALE: Affiliative tactile interactions help regulate physiological arousal and confer resilience to acute and chronic stress. C-tactile afferents (CTs) are a population of unmyelinated, low threshold mechanosensitive cutaneous nerve fibres which respond optimally to a low force stimulus, moving at between 1 and 10 cm/s. As CT firing frequencies correlate positively with subjective ratings of touch pleasantness, they are hypothesised to form the first stage of encoding affiliative tactile interactions. Serotonin is a key modulator of social responses with known effects on bonding. OBJECTIVES: The aim of the present study was to determine the effect of acutely lowering central serotonin levels on perceptions of CT-targeted affective touch. METHODS: In a double blind, placebo-controlled design, the effect of acute tryptophan depletion (ATD) on 25 female participants' ratings of directly and vicariously experienced touch was investigated. Psychophysical techniques were used to deliver dynamic tactile stimuli; some velocities were targeted to optimally activate CTs (1-10 cm/s), whereas other, faster and slower strokes fell outside the CT optimal range. Discriminative tactile function, cold pain threshold and tolerance were also measured. RESULTS: ATD significantly increased pleasantness ratings of both directly and vicariously experienced affective touch, increasing discrimination of the specific hedonic value of CT targeted velocities. While ATD had no effect on either tactile or cold pain thresholds, there was a trend for reduced tolerance to cold pain. CONCLUSIONS: These findings are consistent with previous reports that depletion of central serotonin levels modulates neural and behavioural responsiveness to appetitive sensory signals.

Affective touch topography and body image.

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.

Vicarious ratings of social touch the effect of age and autistic traits.

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 children's 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 children's touch ratings despite parental reports highlighting significant somatosensory sensitivities. These results are discussed in terms of underpinning sensory and cognitive factors.

Children's vicarious ratings of social touch are tuned to the velocity but not the location of a caress.

Affective sharing is a bottom-up process involving automatic processing of sensory inputs that facilitate vicarious experience of another's 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 children's 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 children's 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.

Effects of Acute Tryptophan Depletion on Human Taste Perception.

Taste perception has been reported to vary with changes in affective state. Distortions of taste perception, including blunted recognition thresholds, intensity, and hedonic ratings have been identified in those suffering from depressive disorders. Serotonin is a key neurotransmitter implicated in the etiology of anxiety and depression; systemic and peripheral manipulations of serotonin signaling have previously been shown to modulate taste detection. However, the specific effects of central serotonin function on taste processing have not been widely investigated. Here, in a double-blind placebo-controlled study, acute tryptophan depletion was used to investigate the effect of reduced central serotonin function on taste perception. Twenty-five female participants aged 18-28 attended the laboratory on two occasions at least 1 week apart. On one visit, they received a tryptophan depleting drink and on the other, a control drink was administered. Approximately, 6 h after drink consumption, they completed a taste perception task which measured detection thresholds and supra-threshold perceptions of the intensity and pleasantness of four basic tastes (sweet, sour, bitter, and salt). While acutely reducing central levels of serotonin had no effect on the detection thresholds of sweet, bitter, or sour tastes, it significantly enhanced detection of salt. For supra-threshold stimuli, acutely reduced serotonin levels significantly enhanced the perceived intensity of both bitter and sour tastes and blunted pleasantness ratings of bitter quinine. These findings show manipulation of central serotonin levels can modulate taste perception and are consistent with previous reports that depletion of central serotonin levels enhances neural and behavioral responsiveness to aversive signals.

Childhood Adversity and Affective Touch Perception: A Comparison of United Kingdom Care Leavers and Non-care Leavers.

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.

A positive touch: C-tactile afferent targeted skin stimulation carries an appetitive motivational value.

The rewarding sensation of touch in affiliative interactions is hypothesised to be underpinned by an unmyelinated system of nerve fibres called C-tactile afferents (CTs). CTs are velocity tuned, responding optimally to slow, gentle touch, typical of a caress. Here we used evaluative conditioning to examine whether CT activation carries a positive affective value. A set of neutral faces were paired with robotically delivered touch to the forearm. With half the faces touch was delivered at a CT optimal velocity of 3cm/s (CT touch) and with the other half at a faster, non-CT optimal velocity of 30cm/s (Control touch). Heart-rate and skin conductance responses (SCRs) were recorded throughout. Whilst rated equally approachable pre-conditioning, post-conditioning faces paired with CT touch were judged significantly more approachable than those paired with Control touch. CT touch also elicited significantly greater heart-rate deceleration and lower amplitude SCRs than Control touch. The results indicate CT touch carries a positive affective value, which can be acquired by socially relevant stimuli it is associated with.

C-tactile afferents: Cutaneous mediators of oxytocin release during affiliative tactile interactions?

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, 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.

Vicarious ratings of social touch reflect the anatomical distribution & velocity tuning of C-tactile afferents: A hedonic homunculus?

A subclass of C-fibres, C-tactile afferents (CTs), have been discovered which respond preferentially to low force/velocity stroking touch, that is typically perceived as pleasant. Molecular genetic visualization of these low-threshold mechanosensitive C-fibres (CLTMs) in mice revealed a denser distribution in dorsal than ventral thoracic sites, scattered distal limb innervation and a complete absence from glabrous paw skin (Liu et al., 2007). Here we used third-party ratings to examine whether affective responses to social touch reflect the anatomical distribution and velocity tuning of CTs. Participants viewed and rated a sequence of video clips depicting one individual being touched by another at different skin sites and at 3 different velocities (static, 3cm/s, 30cm/s). Immediately after viewing each clip participants were asked to rate how pleasant they perceived the touch to be. Vicarious preferences matched the previously reported anatomical innervation density of rodent CLTMs, with touch on the back being rated significantly more pleasant than any other location. Furthermore, in contrast to all other skin sites, CT optimal (3cm/s) touch on the palm of the hand was not preferred to static touch, consistent with the anatomical absence of CTs in glabrous skin. Our findings demonstrate that humans recognise the specific rewarding value of CT optimal caressing touch and their preferences reflect the hypothesised anatomical distribution of CTs.

Enhanced subgenual cingulate response to altruistic decisions in remitted major depressive disorder.

BACKGROUND: Major depressive disorder (MDD) is associated with functional abnormalities in fronto-meso-limbic networks contributing to decision-making, affective and reward processing impairments. Such functional disturbances may underlie a tendency for enhanced altruism driven by empathy-based guilt observed in some patients. However, despite the relevance of altruistic decisions to understanding vulnerability, as well as everyday psychosocial functioning, in MDD, their functional neuroanatomy is unknown. METHODS: Using a charitable donations experiment with fMRI, we compared 14 medication-free participants with fully remitted MDD and 15 demographically-matched control participants without MDD. RESULTS: Compared with the control group, the remitted MDD group exhibited enhanced BOLD response in a septal/subgenual cingulate cortex (sgACC) region for charitable donation relative to receiving simple rewards and higher striatum activation for both charitable donation and simple reward relative to a low level baseline. The groups did not differ in demographics, frequency of donations or response times, demonstrating only a difference in neural architecture. CONCLUSIONS: We showed that altruistic decisions probe residual sgACC hypersensitivity in MDD even after symptoms are fully remitted. The sgACC has previously been shown to be associated with guilt which promotes altruistic decisions. In contrast, the striatum showed common activation to both simple and altruistic rewards and could be involved in the so-called "warm glow" of donation. Enhanced neural response in the depression group, in areas previously linked to altruistic decisions, supports the hypothesis of a possible association between hyper-altruism and depression vulnerability, as shown by recent epidemiological studies.