Diverse stimuli induce piloerection and yield varied autonomic responses in humans.

This research provides an in-depth exploration into the triggers and corresponding autonomic responses of piloerection, a phenomenon prevalent across various species. In non-human species, piloerection occurs in reaction to a variety of environmental changes, including social interactions and temperature shifts. However, its understanding in humans has been confined to emotional contexts. This is problematic because it reflects solely upon subjective experience rather than an objective response to the environment. Further, given our shared evolutionary paths, piloerection should function similarly in humans and other animals. I observed 1,198 piloerection episodes from eight participants while simultaneously recording multiple autonomic and body temperature indices, finding that piloerection in humans can be elicited by thermal, tactile, and audio-visual stimuli with equal effectiveness. The data also revealed variations in cardiac reactivity measures: audio-visual piloerection was associated with greater sympathetic arousal, while tactile piloerection was linked to greater parasympathetic arousal. Despite prevailing notions of piloerection as a vestigial response in humans, it does respond to decreases in skin temperature and is associated with a rise in skin temperature during episodes. This research underscores that piloerection in humans is not purely vestigial, nor is it solely an affective response to emotional stimuli. Rather, it is best understood as a reflexive response to environmental changes, suggesting a shared functional similarity with other species.

High frequency heart rate variability is associated with sensitivity to affective touch.

C-tactile afferents (CTs) are a class of unmyelinated, mechanosensitive nerve fibre that respond optimally to skin temperature, slow moving touch typical of a caress. They are hypothesised to signal the rewarding value of affiliative tactile interactions. While CT firing frequency is positively correlated with subjective ratings of touch pleasantness, trait differences in sensitivity to the specific hedonic value of CT targeted touch have been reported. Inter-individual differences in vagally mediated, high frequency heart rate variability (HF-HRV) have been linked to variation in visual social cognition. Thus, the aim of the present study was to examine the relationship between resting state HF-HRV and sensitivity to socially relevant CT targeted touch. 58 healthy participants first had a 5-minute electrocardiogram. They then rated the pleasantness of 5 randomly presented velocities of robotically delivered touch. Three velocities fell within (1, 3, 10 cm/s) and two outside (0.3, 30 cm/s) the CT optimal range. Each velocity was delivered twice. On a group level, affective touch ratings were described by a negative quadratic function, with CT optimal velocities rated as more pleasant than slower and faster speeds. Simple regression analysis confirmed participants' HF-HRV was significantly predicted by the quadratic curve fit of their touch ratings, with higher HF-HRV associated with a better quadratic fit. These findings indicate that, in line with previous observations that higher HF-HRV is associated with enhanced sensitivity to visual social cues, trait differences in autonomic control could account for previously reported individual differences in CT sensitivity.

Age-dependent small fiber neuropathy: Mechanistic insights from animal models.

Small fiber neuropathy (SFN) is a common and debilitating disease in which the terminals of small diameter sensory axons degenerate, producing sensory loss, and in many patients neuropathic pain. While a substantial number of cases are attributable to diabetes, almost 50% are idiopathic. An underappreciated aspect of the disease is its late onset in most patients. Animal models of human genetic mutations that produce SFN also display age-dependent phenotypes suggesting that aging is an important contributor to the risk of development of the disease. In this review we define how particular sensory neurons are affected in SFN and discuss how aging may drive the disease. We also evaluate how animal models of SFN can define disease mechanisms that will provide insight into early risk detection and suggest novel therapeutic interventions.

Determinants for positive and negative experiences of interpersonal touch: context matters.

The goal of the study was to determine which aspects of interpersonal touch interactions lead to a positive or negative experience. Previous research has focused primarily on physical characteristics. We suggest that this may not be sufficient to fully capture the complexity of the experience. Specifically, we examined how fulfilment of psychological needs influences touch experiences and how this relates to physical touch characteristics and situational factors.In two mixed-method studies, participants described their most positive and most negative interpersonal touch experience within a specific time frame. They reported fulfilment of nine needs, affect, intention, and reason for positivity/negativity, as well as the body part(s) touched, location, type of touch, interaction partner, and particular touch characteristics (e.g. humidity).Positive and negative touch experiences shared similar touch types, locations, and body parts touched, but differed in intended purpose and reasons. Overall, the valence of a touch experience could be predicted from fulfilment of relatedness, the interaction partner and initiator, and physical touch characteristics. Positive affect increased with need fulfilment, and negative affect decreased.The results highlight the importance of relatedness and reciprocity for the valence of touch, and emphasise the need to incorporate psychological needs in touch research.

Ghrelin is related to lower brain reward activation during touch.

The gut hormone ghrelin drives food motivation and increases food intake, but it is also involved in the anticipation of and response to rewards other than food. This pre-registered study investigated how naturally varying ghrelin concentrations affect the processing of touch as a social reward in humans. Sixty-seven volunteers received slow caressing touch (so-called CT-targeted touch) as a social reward and control touch on their shins during 3T functional imaging on two test days. On one occasion, participants were fasted, and on another, they received a meal. On each occasion, plasma ghrelin was measured at three time points. All touch was rated as more pleasant after the meal, but there was no association between ghrelin concentrations and pleasantness. CT-targeted touch was rated as the most pleasant and activated somatosensory and reward networks (whole brain). A region-of-interest in the right medial orbitofrontal cortex (mOFC) showed lower activation during all touches, the higher the ghrelin concentrations were. During CT-targeted touch, a larger satiety response (ghrelin decrease after the meal) was associated with higher mOFC activation, and this mOFC activation was associated with higher experienced pleasantness. Overall, higher ghrelin concentrations appear to be related to a lower reward value for touch. Ghrelin may reduce the value of social stimuli, such as touch, to promote food search and intake in a state of low energy. This suggests that the role of ghrelin goes beyond assigning value to food reward.

Oscillatory Responses to Tactile Stimuli of Different Intensity.

Tactile perception encompasses several submodalities that are realized with distinct sensory subsystems. The processing of those submodalities and their interactions remains understudied. We developed a paradigm consisting of three types of touch tuned in terms of their force and velocity for different submodalities: discriminative touch (haptics), affective touch (C-tactile touch), and knismesis (alerting tickle). Touch was delivered with a high-precision robotic rotary touch stimulation device. A total of 39 healthy individuals participated in the study. EEG cluster analysis revealed a decrease in alpha and beta range (mu-rhythm) as well as theta and delta increase most pronounced to the most salient and fastest type of stimulation. The participants confirmed that slower stimuli targeted to affective touch low-threshold receptors were the most pleasant ones, and less intense stimuli aimed at knismesis were indeed the most ticklish ones, but those sensations did not form an EEG cluster, probably implying their processing involves deeper brain structures that are less accessible with EEG.

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.

Cortical Response Variation with Social and Non-Social Affective Touch Processing in the Glabrous and Hairy Skin of the Leg: A Pilot fMRI Study.

Despite the crucial role of touch in social development and its importance for social interactions, there has been very little functional magnetic resonance imaging (fMRI) research on brain mechanisms underlying social touch processing. Moreover, there has been very little research on the perception of social touch in the lower extremities in humans, even though this information could expand our understanding of the mechanisms of the c-tactile system. Here, variations in the neural response to stimulation by social and non-social affective leg touch were investigated using fMRI. Participants were subjected to slow a (at 3-5 cm/s) stroking social touch (hand, skin-to-skin) and a non-social touch (peacock feather) to the hairy skin of the shin and to the glabrous skin of the foot sole. Stimulation of the glabrous skin of the foot sole, regardless of the type of stimulus, elicited a much more widespread cortical response, including structures such as the medial segment of precentral gyri, left precentral gyrus, bilateral putamen, anterior insula, left postcentral gyrus, right thalamus, and pallidum. Stimulation of the hairy skin of the shin elicited a relatively greater response in the left middle cingulate gyrus, left angular gyrus, left frontal eye field, bilateral anterior prefrontal cortex, and left frontal pole. Activation of brain structures, some of which belong to the "social brain"-the pre- and postcentral gyri bilaterally, superior and middle occipital gyri bilaterally, left middle and superior temporal gyri, right anterior cingulate gyrus and caudate, left middle and inferior frontal gyri, and left lateral ventricle area, was associated with the perception of non-social stimuli in the leg. The left medial segment of pre- and postcentral gyri, left postcentral gyrus and precuneus, bilateral parietal operculum, right planum temporale, left central operculum, and left thalamus proper showed greater activation for social tactile touch. There are regions in the cerebral cortex that responded specifically to hand and feather touch in the foot sole region. These areas included the posterior insula, precentral gyrus; putamen, pallidum and anterior insula; superior parietal cortex; transverse temporal gyrus and parietal operculum, supramarginal gyrus and planum temporale. Subjective assessment of stimulus ticklishness was related to activation of the left cuneal region. Our results make some contribution to understanding the physiology of the perception of social and non-social tactile stimuli and the CT system, including its evolution, and they have clinical impact in terms of environmental enrichment.

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

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

Aß-CT Affective Touch: Touch Pleasantness Ratings for Gentle Stroking and Deep Pressure Exhibit Dependence on A-Fibers.

Gentle stroking of the skin is a common social touch behavior with positive affective consequences. A preference for slow versus fast stroking of hairy skin has been closely linked to the firing of unmyelinated C-tactile (CT) somatosensory afferents. Because the firing of CT afferents strongly correlates with touch pleasantness, the CT pathway has been considered a social-affective sensory pathway. Recently, ablation of the spinothalamic pathway- thought to convey all C-fiber sensations- in patients with cancer pain impaired pain, temperature, and itch, but not ratings of pleasant touch. This suggested integration of afferent A and CT fiber input in the spinal cord, or mechanoreceptive A-fiber contributions to computations of touch pleasantness in the brain. However, contribution of mechanoreceptive A-fibers to touch pleasantness, in humans without pain, remains unknown. In the current, single-blinded study, we performed two types of peripheral nerve blocks in healthy adults to temporarily eliminate the contribution of A-fibers to touch perception. Our findings show that when mechanoreceptive A-fiber function is greatly diminished, the perceived intensity and pleasantness of both gentle stroking and deep pressure are nearly abolished. These findings demonstrate that explicit perception of the pleasantness of CT-targeted brushing and pressure both critically depend on afferent A-fibers.

What Makes Touch Comfortable? An Examination of Touch Giving and Receiving in Two Cultures.

This study examined how touch role and culture shape affective touch experiences. Germans (N = 130) and Chinese (N = 130) were surveyed once as toucher and once as touchee. For different touch actions, they (a) provided free-text descriptions of what prompts touch, (b) indicated with whom touch feels comfortable, and (c) highlighted areas of touch comfort on a body outline. Overall, touch was prompted by affectionate feelings, was more comfortable with more closely bonded individuals, and when directed at the upper arms, shoulders, and upper back. Touch role mattered for the experiences prompting touch in that touchees felt less positive than touchers. Culture differentiated touch comfort topographies. Compared with Chinese, Germans felt more comfortable with more intimate touch to the torso and upper back and less comfortable with more public touch to the hands. Notably, however, examining touch role and culture revealed more overlap than divergence, ensuring mutual comfort as individuals physically connect.

Dynamic touch induces autonomic changes in preterm infants as measured by changes in heart rate variability.

Preterm birth significantly increases the risk of developing various long-term health problems and developmental disabilities. While touch is a crucial component of many perinatal care strategies, the neurobiological underpinnings are rarely considered. C-tactile fibers (CTs) are unmyelinated nerve fibers that are activated by low-force, dynamic touch. Touch directed specifically at CTs activates the posterior insular cortex, consistent with an interoceptive function, and has been shown to reduce heart rate and increase oxygen saturation. The current research compared the effect of five minutes of CT optimal velocity stroking touch versus five minutes of static touch on autonomic markers of preterm infants between 28 and 37 weeks gestational age. CT touch induces a higher increase in heart rate variability metrics related to the parasympathetic system, which persisted for a 5-minute post-touch period. Conversely, there was no such increase in infants receiving static touch. The present findings confirmed that CTs signal the affective quality of nurturing touch, thereby arguing an additional neurobiological substrate for the evident valuable impacts of neonatal tactile interventions and improving the effectiveness of such interventions.

Perspectives on interpersonal touch are related to subjective sleep quality.

Affective touch has been reported for its calming effects; however, it is less clear whether touch is associated with sleep. Here, the relationship between different touch variables and self-reported sleep indicators was investigated. Data were extracted from the Touch Test, a cross-sectional survey conducted in 2020. Data from a sample of 15,049 healthy adults from the UK (mean age = 56.13, SD = 13.8; 75.4% female) were analysed. Participants were asked to attribute positive, negative, or no effects on sleep to hugs, strokes, massages, intimate touch, and sleep onset with and without touch. The time since last intentional touch, touch amount satisfaction, and childhood bed routine with hugs and kisses were assessed. Sleep quality, duration, latency, wake after sleep onset and diurnal preference were measured. Data were analysed using chi-square tests and logistic regressions. Affective touch before sleep was perceived to have positive effects on sleep. Touch recency emerged as a significant predictor for some sleep variables, with a longer timespan since the last intentional touch relating to improved sleep quality, longer sleep duration, and shorter and fewer instances of waking up after sleep onset in some participants. Experiencing too much touch was related to lower sleep quality and higher instances of waking up after sleep onset. These findings highlight the importance of interpersonal touch for subjective sleep quality.

Affective touch and regulation of stress responses.

Much has been documented on the association between stress and health. Both direct and indirect pathways have been identified and explored extensively, helping us understand trajectories from healthy individuals to reductions in well-being, and development of preclinical and disease states. Some of these pathways are well established within the field; physiology, affect regulation, and social relationships. The purpose of this review is to push beyond what is known separately about these pathways and provide a means to integrate them using one common mechanism. We propose that social touch, specifically affective touch, may be the missing active ingredient fundamental to our understanding of how close relationships contribute to stress and health. We provide empirical evidence detailing how affective touch is fundamental to the development of our stress systems, critical to the development of attachment bonds and subsequent social relationships across the life course. We will also explore how we can use this in applied contexts and incorporate it into existing interventions.

Gentle as a mother's touch: C-tactile touch promotes autonomic regulation in preterm infants.

Preterm infants are challenged to adapt to an extrauterine milieu, while their interoceptive system and autonomic regulation capacity is still immature. Caressing parental touch is known to foster parasympathetic regulation in infants by stimulating C-tactile (CT) afferents and in preterm infants, slow stroking stimulation also leads to a heart rate decrease. The particular impact of maternal stroking has not yet been investigated and factors influencing the maturation of the CT system in preterm infants remain unclear. We therefore analysed 53 standardized events in which preterm infants (24 to 36 weeks gestational age at birth) were stroked by their mothers. Video analysis revealed that mothers use CT optimal velocities to stroke their preterm child. Analysis of pulse oximetric data showed no effect of stroking on infantile blood oxygenation, but a significant decrease of the heart rate. Compared to term-born children, this decrease was delayed by about two minutes. Furthermore, our data suggested that more immature preterm infants benefited less from stroking than more mature ones. We conclude that maternal stroking touch targets CT afferents in preterm infants and that the preterm CT system is not yet mature.

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.

Osteopathic Manipulative Treatment Regulates Autonomic Markers in Preterm Infants: A Randomized Clinical Trial.

Osteopathic manipulative treatment (OMT) has been found to be effective in the context of premature infants. Nonetheless, no studies have investigated the immediate effects of OMT on heart rate variability (HRV). As altered HRV reflects poor or worsening newborn's clinical conditions and neurodevelopment, should OMT improve HRV fluctuations, it could become a relevant intervention for improving the care of preterm newborns. Therefore, this study aimed to evaluate whether OMT could affect HRV. The study was carried out at the Buzzi Hospital in Milan. From the neonatal intensive care unit, ninety-six preterm infants (41 males) were enrolled and were randomly assigned to one of two treatment groups: OMT or Static Touch. The infants were born at 33.5 weeks (±4.3) and had a mean birth weight of 2067 g (±929). The study had as primary outcome the change in the beat-to-beat variance in heart rate measured through root mean square of consecutive RR interval differences (RMSSD); other metrics were used as secondary and exploratory analyses. Despite the lack of statistically significant results regarding the primary outcomeand some study limitations, compared to static touch, OMT seemed to favor a parasympathetic modulation and improved HRV, which could reflect improvement in newborn's clinical conditions and development.

Understanding sex differences in affective touch: Sensory pleasantness, social comfort, and precursive experiences.

Although previous research revealed sex differences in affective touch, the implicated processes and the manner in which men and women differ have been left uncertain. Here we addressed this issue in two studies examining sensory pleasure, interpersonal comfort, and touch motivators. Study 1 comprised a series of lab-based experiments in which a robot stroked 214 participants (half female) at five different velocities modulating the activity of C-tactile afferents thought to support tactile pleasantness. Average pleasantness ratings followed velocity with the typical inverted u-shape similarly in both sexes. In Study 2, 260 participants (half female) completed an online survey. Here, women were more likely than men to express touch comfort with less familiar or unknown individuals, had a greater preference for touch with other women, and felt more comfortable giving and receiving touch to the forearm. Additionally, when describing how their own experiences might motivate others to touch them affectively, women produced more negative descriptions than men. Together, these results show that, while the sexes compare in a touch's sensory pleasantness, they differ in their preceding affective experiences and how they value touch at a higher-order social level. This agrees with extant research on negative affect and stress and suggests that affective touch may be a more relevant coping mechanism for women than for men.

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.

C-low threshold mechanoafferent targeted dynamic touch modulates stress resilience in rats exposed to chronic mild stress.

Affiliative tactile interactions buffer social mammals against neurobiological and behavioral effects of stress. The aim of this study was to investigate the cutaneous mechanisms underlying such beneficial consequences of touch by determining whether daily stroking, specifically targeted to activate a velocity/force tuned class of low-threshold c-fiber mechanoreceptor (CLTM), confers resilience against established markers of chronic unpredictable mild stress (CMS). Adult male Sprague Dawley rats were exposed to 2 weeks of CMS. Throughout the CMS protocol, some rats were stroked daily, either at CLTM optimal velocity (5 cm/s) or outside the CLTM optimal range (30 cm/s). A third CMS exposed group did not receive any tactile stimulation. The effect of CMS on serum corticosterone levels, anxiety- and depressive-like behaviors in these three groups was assessed in comparison to a control group of non-CMS exposed rats. While stroking did not mitigate the effects of CMS on body weight gain, CLTM optimal velocity stroking did significantly reduce CMS-induced elevations in corticosterone following an acute forced-swim. Rats receiving CLTM optimal stroking also showed significantly fewer anxiety-like behaviors (elevated plus-maze) than the other CMS exposed rats. In terms of depressive-like behavior, whereas the same velocity-specific resilience was observed in a forced-swim test and social interaction test both groups of stroked rats spent significantly less time interacting than control rats, though they also spent significantly less time in the corner than non-stroked CMS rats. Together, these findings support the theory CLTMs play a functional role in regulating the physiological condition of the body.