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Touch sensitivity generally declines with age, contributing to loss of manual dexterity and tactile function. We investigated how touch changes over the lifespan, using different tests and on three body sites. We used a classical test of force detection sensitivity, where calibrated monofilaments were applied passively to the right index finger pad, forearm, and cheek. In addition, at the index, we used an active touch spatial discrimination task, developed by our group. Spatial discrimination was estimated through participants' ability to evaluate the distance between parallel bands printed on acrylic plates. Data were collected from 96 healthy women, aged 20-75 years. Force detection and tactile spatial discrimination on the index deteriorated significantly with age; however, no change was found for tactile detection on the forearm or cheek. Tactile detection on the cheek remained remarkably highly sensitive throughout life. There was a significant positive relationship between force detection and spatial discrimination on the index. Further, force detection on the forearm was significantly associated with detection on the index and cheek. Our results suggest a decrease in touch perception with age on the index finger pad, yet a preservation of tactile sensitivity in hairy skin. This opens discussion about the impact of daily activities upon the glabrous hand skin and on the function of hairs in tactile sensitivity. We highlight the need for new methods in evaluating tactile sensitivity on hairy skin., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Samain-Aupic, Dione, Ribot-Ciscar, Ackerley and Aimonetti.)

The human body is encompassed by a thin layer of tissue, the skin, which is heterogenous and highly specialized to protect the body and encode interactions with the external world. There is a fundamental scientific drive to understand its function, coupled with the need to preserve skin as we age, which impacts on our physiological and psychological well-being. In the present study, we aimed to define differences in touch perception between age groups and with skin cream application. We investigated touch on the finger, the forearm and cheek in younger (20-28 years, n = 22) and older (65-75 years, n = 22) females. We measured skin hydration, touch detection, finger spatial discrimination, forearm tactile pleasantness together with electrodermal activity, and perceptual ratings about cream use, skin dryness, and cosmetic habits. Glabrous finger skin became drier and touch performance was impaired with age, but these aspects were preserved in hairy skin. Skin moisturization immediately increased hydration levels, but did not significantly change touch perception. We also found that touch appreciation increased with age. We conclude that reduced finger capacity may impact self-evaluation of the skin and that long-term skin care strategies should focus on hydrating the hand to preserve touch capacities., (© 2023. Springer Nature Limited.)

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., Competing Interests: Declaration Of Competing Interest The authors declare no competing interests., (Copyright © 2023 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Tactile sensitivity generally decreases with aging and is associated with impairments in skin properties. Products that hydrate the skin can combat touch deficits and aromatic compounds have been shown to improve skin mechanical properties. Thus, we tested a base cosmetic oil against a perfumed oil, applied to the skin of females aged 40-60 years, on tactile sensitivity and skin properties after repeated application. Tactile detection thresholds were assessed using calibrated monofilaments applied at the index finger, palm, forearm, and cheek. Spatial discrimination on the finger was assessed using pairs of plates with different inter-band spaces. These tests were performed before and after 1 month of base or perfumed oil use. We found that tactile detection thresholds and spatial discrimination improved only in perfumed oil group. A complementary immunohistological study using human skin was conducted to estimate the expression of olfactory receptor OR2A4 and elastic fiber length. Further, the expression of OR2A4 intensity and the length of elastic fibers increased significantly with oil application, where larger effects were seen with the perfumed oil. We conclude that the application of a perfumed oil may be of additional benefit and could repair, and even prevent, tactile decline with aging by ameliorating skin condition., (© 2023. The Author(s).)

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

Slowly-adapting type II (SA-II, Ruffini) mechanoreceptive afferents respond well to pressure and stretch, and are regularly encountered in human microneurography studies. Despite an understanding of SA-II response properties, their role in touch perception remains unclear. Specific roles of different myelinated Aβ mechanoreceptive afferents in tactile perception have been revealed using single unit intraneural microstimulation (INMS), via microneurography, recording from and then electrically stimulating individual afferents. This method directly links single afferent artificial activation to perception, where INMS produces specific 'quantal' touch percepts associated with different mechanoreceptive afferent types. However, SA-II afferent stimulation has been ambiguous, producing inconsistent, vague sensations, or no clear percept. We physiologically characterized hundreds of individual Aβ mechanoreceptive afferents in the glabrous hand skin and examined the subsequent percepts evoked by trains of low amplitude INMS current pulses (<10 μA). We present 18 SA-II afferents where INMS resulted in a clear, electrically evoked sensation of large (∼36 mm 2 ) diffuse pressure, which was projected precisely to their physiologically-defined receptive field in the skin. This sensation was felt as natural, distinctive from other afferents, and showed no indications of multi-afferent stimulation. Stimulus frequency modulated sensation intensity and even brief stimuli (4 pulses, 60 ms) were perceived. These results suggest that SA-II afferents contribute to perceived tactile sensations, can signal this rapidly and precisely, and are relevant and important for computational models of touch sensation and artificial prosthetic feedback. KEY POINTS: Slowly adapting type II mechanoreceptors (SA-IIs) are primary sensory neurons in humans that respond to pressure and stretch applied to the skin. To date, no specific conscious correlate of touch has been linked to SA-II activation. Using microneurography and intraneural microstimulation to stimulate single sensory neurons in human subjects, we find a specific sensation linked to the activation of single SA-II afferents. This sensation of touch was reported as gentle pressure and subjects could detect this with a high degree of accuracy. Methods of artificial tactile sensory feedback and computational models of touch should include SA-IIs as meaningful contributors to the conscious sensation of touch., (© 2022 The Authors. The Journal of Physiology published by John Wiley & Sons Ltd on behalf of The Physiological Society.)

Humans rely on precise proprioceptive feedback from our muscles, which is important in both the acquisition and execution of movements, to perform daily activities. Somatosensory input from the body shapes motor learning through central processes, as demonstrated for tasks using the arm, under active (self-generated) and passive conditions. Presently, we investigated whether passive movement training of the ankle increased proprioceptive acuity (psychophysical experiment) and whether it changed the peripheral proprioceptive afferent signal (microneurography experiment). In the psychophysical experiment, the ankle of 32 healthy human participants was moved passively using pairs of ramp-and-hold movements in different directions. In a pretraining test, participants made judgements about the movement direction in a two-alternative forced choice paradigm. Participants then underwent passive movement training, but only half were cued for learning, where a reference position was signaled by a sound and the participant had to learn to recognize this position; they then completed a post-training test. In a paradigm using the same setup, nine healthy participants underwent microneurography recordings of Ia muscle afferents from the peroneal nerve, where all were cued during training. In the psychophysical experiment, proprioceptive acuity improved with training only in the cued group. In the microneurography experiment, we found that muscle afferent firing was modulated, via an increase in the dynamic index, after training. We suggest that changes in muscle afferent input from the periphery can contribute to and support central perceptual and motor learning, as shown under passive conditions using ankle movements, which may be exploited for movement rehabilitation., (Copyright © 2022 Ackerley et al.)

Many studies have investigated the perception of tactile pleasantness over a range of stroking velocities. On average, pleasantness is low at slow (e.g. 0.3 cm/s) and fast (e.g. 30 cm/s) stroking velocities, but is rated highest at velocities between 1 and 10 cm/s. On a group level, this results in an inverted-U shape pleasantness ratings curve, which is described statistically by a negative quadratic equation. We reanalyzed the data from five earlier studies to investigate whether the inverted-U shape pleasantness curve at the group level is also present at the level of the individual, - a precondition for using tactile pleasantness perception as a diagnostic marker. We pooled the data from five studies with a total of 127 participants. Each study included a 'standard condition' of stroking on the dorsal forearm over different velocities (0.3, 1, 3, 10, 30 cm/s) and participants rated the pleasantness. Factors other than stroking velocity were also varied in these studies. On the whole-group level and in each study, pleasantness ratings produced a significant negative quadratic pleasantness curve over the stroking velocities. In individual participants, ratings varied greatly and only 42% of the participants showed a significant negative quadratic curve. The steepness of the inverted-U correlated only moderately across other experimental conditions, showing that the experimental circumstances can influence pleasantness ratings. Our findings have important implications for future work, where differences in the tactile pleasantness curve should not be used to predict or diagnose issues at an individual level., (Copyright © 2020 IBRO. Published by Elsevier Ltd. All rights reserved.)

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

Cutaneous foot receptors are important for balance control, and their activation during quiet standing depends on the speed and the amplitude of postural oscillations. We hypothesized that the transmission of cutaneous input to the cortex is reduced during prolonged small postural sways due to receptor adaptation during continued skin compression. Central mechanisms would trigger large sways to reactivate the receptors. We compared the amplitude of positive and negative post-stimulation peaks (P 50 N 90 ) somatosensory cortical potentials evoked by the electrical stimulation of the foot sole during small and large sways in 16 young adults standing still with their eyes closed. We observed greater P 50 N 90 amplitudes during large sways compared with small sways consistent with increased cutaneous transmission during large sways. Postural oscillations computed 200 ms before large sways had smaller amplitudes than those before small sways, providing sustained compression within a small foot sole area. Cortical source analyses revealed that during this interval, the activity of the somatosensory areas decreased, whereas the activity of cortical areas engaged in motor planning (supplementary motor area, dorsolateral prefrontal cortex) increased. We concluded that large sways during quiet standing represent self-generated functional behavior aiming at releasing skin compression to reactivate mechanoreceptors. Such balance motor commands create sensory reafference that help control postural sway., (© The Author(s) 2020. Published by Oxford University Press.)

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

Positive affective touch plays a central role in social and inter-personal interactions. Low-threshold mechanoreceptive afferents, including slowly-conducting C-tactile (CT) afferents found in hairy skin, transmit such signals from gentle touch to the brain. Tactile signals are processed, in part, by the posterior insula, where it is the thought to be the primary target for CTs. We used magnetoencephalography (MEG) to assess brain activity evoked by gentle, naturalistic stroking touch on the arm delivered by a new MEG-compatible brush robot. We aimed to use high temporal resolution MEG to allow us to distinguish between brain responses from fast-conducting Aβ and slowly-conducting CT afferents. Brush strokes were delivered to the left upper arm and left forearm of 15 healthy participants. We hypothesized that late brain responses, due to slow CT afference, would appear with a time shift between the two different locations on the arm. Our results show that gentle touch rapidly activated somatosensory, motor, and cingulate regions within the first 100 ms of skin contact, which was driven by fast-conducting mechanoreceptive afference, and that these responses were sustained during touch. Peak latencies in the posterior insula were shifted as a function of stimulus location and temporally-separate posterior insula activations were induced by Aβ and CT afference that may modulate the emotional processing of gentle touch on hairy skin. We conclude that the detailed information regarding temporal and spatial brain activity from MEG provides new insights into the central processing of gentle, naturalistic touch, which is thought to underpin affective tactile interactions., (Copyright © 2019 The Authors. Published by Elsevier Inc. All rights reserved.)

The sensation of touch in the glabrous skin of the human hand is conveyed by thousands of fast-conducting mechanoreceptive afferents, which can be categorised into four distinct types. The spiking properties of these afferents in the periphery in response to varied tactile stimuli are well-characterised, but relatively little is known about the spatiotemporal properties of the neural representations of these different receptor types in the human cortex. Here, we use the novel methodological combination of single-unit intraneural microstimulation (INMS) with magnetoencephalography (MEG) to localise cortical representations of individual touch afferents in humans, by measuring the extracranial magnetic fields from neural currents. We found that by assessing the modulation of the beta (13-30 Hz) rhythm during single-unit INMS, significant changes in oscillatory amplitude occur in the contralateral primary somatosensory cortex within and across a group of fast adapting type I mechanoreceptive afferents, which corresponded well to the induced response from matched vibrotactile stimulation. Combining the spatiotemporal specificity of MEG with the selective single-unit stimulation of INMS enables the interrogation of the central representations of different aspects of tactile afferent signalling within the human cortices. The fundamental finding that single-unit INMS ERD responses are robust and consistent with natural somatosensory stimuli will permit us to more dynamically probe the central nervous system responses in humans, to address questions about the processing of touch from the different classes of mechanoreceptive afferents and the effects of varying the stimulus frequency and patterning., (Copyright © 2019 The Authors. Published by Elsevier Inc. All rights reserved.)

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

Kinesthesia, the perception of our own body movements, relies on the integration of proprioceptive information arising mostly from muscle spindles, which are sensory receptors in skeletal muscles. We recently showed that emotions alter the proprioceptive messages from such muscle afferents, making them more sensitive to muscle lengthening when participants were listening sad music. Presently, we investigated whether these changes in proprioceptive feedback relating to emotional state may affect the perception of limb movements. Kinesthetic acuity was tested in 20 healthy, young adults by imposing ramp-and-hold movements that consisted of either plantar flexion or dorsiflexion movements of the ankle at 0.04°/s, or no movement. These were imposed during four emotional conditions (listening to neutral, sad, or happy music, or no music). The participants were asked to relax and focus on music (or nothing), and then they shifted their focus to the direction of an incoming movement. Once this had finished, they were asked its direction. Muscle activity, heart rate, and electrodermal activity were recorded during each trial, and after each music condition the participants rated the emotion felt on a visual analog scale. The rating of the emotional content of the music corroborated with changes in physiological measures. Kinesthetic acuity was also affected by the emotional state and found to be larger during the sad condition, as compared to the no music or neutral conditions. We conclude that emotion can shape our perception of movements, which we show here where feeling sadness significantly increase our kinesthetic acuity, this may be functionally relevant for the preparation of appropriate behavioral responses., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Touch is a common occurrence in our lives, where affective and inter-personal aspects of touch are important for our well-being. We investigated whether touch exposure affects hedonic and discriminative aspects of tactile perception. The perceived pleasantness and intensity of gentle forearm stroking, over different velocities, was assessed in individuals reporting to seldom receive inter-personal touch, and in controls who received touch often. The groups did not differ in their stroking intensity judgements, nor in tactile discrimination sensitivity; however, individuals with low touch exposure evaluated the pleasantness of touch differently. These individuals did not differentiate pleasantness over the stroking velocities in the same way as the control group. The pleasantness curve for the low touch exposure group was significantly flatter and they rated 3cm/s stroking as significantly less pleasant. Other physiological and questionnaire measures were obtained and the appreciation of touch from familiar persons was positively related to the pleasantness of touch in controls, but this was not found in low touch exposure individuals. This suggests that the association of human caresses from well-known individuals, with the pleasure derived, may depend on continued exposure to it., (Copyright © 2017 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Affective touch plays an important role in children's social interaction and is involved in shaping the development of the social brain. The positive affective component of touch is thought to be conveyed via a group of unmyelinated, low-threshold mechanoreceptive afferents, known as C-tactile fibers that are optimally activated by gentle, slow, stroking touch. Touch targeting these C-tactile fibers has been shown to decrease the heart rate in infants. The current study investigated the relationship between age and psychophysical ratings in response to affective touch. A total of n=43 participants (early childhood: aged 5-8 years, 9 girls, 12 boys; late childhood: aged 9-12 years, 12 girls, 10 boys) were presented with C-tactile optimal and sub-optimal stroking velocities and rated touch pleasantness on an affective pictorial scale. For both age groups, we found that children preferred C-tactile-targeted stimulation. A comparison with previously published data showed that the children's preference for C-tactile-targeted stimulation was similar to those obtained in adolescents and adults. We speculate that the effect of C-tactile-targeted touch, which is linked with pleasantness, shapes the children's preference for C-tactile over non-C-tactile-targeted stimulation, and that C-tactile afferent stimulation is important for social development., (Copyright © 2017 The Authors. Published by Elsevier Ltd.. All rights reserved.)

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

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

The present case study details sensations elicited by electrical stimulation of peripheral nerve axons using an implanted nerve cuff electrode, in a participant with a transhumeral amputation. The participant uses an osseointegrated electromechanical interface, which enables skeletal attachment of the prosthesis and long-term, stable, bidirectional communication between the implanted electrodes and prosthetic arm. We focused on evoking somatosensory percepts, where we tracked and quantified the evolution of perceived sensations in the missing hand, which were evoked from electrical stimulation of the nerve, for over 2 yr. These sensations included small, pointlike areas of either vibration or pushing, to larger sensations over wider areas, indicating the recruitment of a few and many afferents, respectively. Furthermore, we used a two-alternative forced choice paradigm to measure the level of discrimination between trains of brief electrical stimuli, to gauge what the participant could reliably distinguish between. At best, the participant was able to distinguish a 0.5-Hz difference and on average acquired a 3.8-Hz just-noticeable difference at a more stringent psychophysical level. The current work shows the feasibility for long-term sensory feedback in prostheses, via electrical axonal stimulation, where small and relatively stable percepts were felt that may be used to deliver graded sensory feedback. This opens up opportunities for signaling feedback during movements (e.g., for precision grip), but also for conveying more complex cutaneous sensations, such as texture. NEW & NOTEWORTHY We demonstrate the long-term stability and generation of sensations from electrical peripheral nerve stimulation in an amputee, through an osseointegrated implant. We find that perceived tactilelike sensations could be generated for over 2 yr, in the missing hand. This is useful for prosthetic development and the implementation of feedback in artificial body parts.

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 (7T) 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 <7μA, 200μs pulses) of individual mechanoreceptive afferents produces appropriate and robust responses during fMRI and MEG., Comparison With Existing Method(s): This custom-built MRI- and MEG-compatible stimulator overcomes issues with existing INMS approaches; it allows well-controlled switching between recording and stimulus mode, prevents electrical shocks because of long cable lengths, permits unlimited patterns of stimulation, and provides a system with improved work-flow and participant comfort., Conclusions: We demonstrate that the requirements for an INMS-integrated system, which can be used with both fMRI and MEG imaging systems, have been fully met., (Copyright © 2017 The Author(s). Published by Elsevier B.V. All rights reserved.)

Emotions can evoke strong reactions that have profound influences, from gross changes in our internal environment to small fluctuations in facial muscles, and reveal our feelings overtly. Muscles contain proprioceptive afferents, informing us about our movements and regulating motor activities. Their firing reflects changes in muscle length, yet their sensitivity can be modified by the fusimotor system, as found in animals. In humans, the sensitivity of muscle afferents is modulated by cognitive processes, such as attention; however, it is unknown if emotional processes can modulate muscle feedback. Presently, we explored whether muscle afferent sensitivity adapts to the emotional situation. We recorded from single muscle afferents in the leg, using microneurography, and moved the ankle joint of participants, while they listened to evocative classical music to induce sad, neutral, or happy emotions, or sat passively (no music). We further monitored their physiological responses using skin conductance, heart rate, and electromyography measures. We found that muscle afferent firing was modified by the emotional context, especially for sad emotions, where the muscle spindle dynamic response increased. We suggest that this allows us to prime movements, where the emotional state prepares the body for consequent behaviour-appropriate reactions.

C-mechanoreceptors in humans comprise a population of unmyelinated afferents exhibiting a wide range of mechanical sensitivities. C-mechanoreceptors are putatively divided into those signaling gentle touch (C-tactile afferents, CTs) and nociception (C-mechanosensitive nociceptors, CMs), giving rise to positive and negative affect, respectively. We sought to distinguish, compare, and contrast the properties of a population of human C-mechanoreceptors to see how fundamental the divisions between these putative subpopulations are. We used microneurography to record from individual afferents in humans and applied electrical and mechanical stimulation to their receptive fields. We show that C-mechanoreceptors can be distinguished unequivocally into two putative populations, comprising CTs and CMs, by electrically evoked spike latency changes (slowing). After both natural mechanical stimulation and repetitive electrical stimulation there was markedly less latency slowing in CTs compared with CMs. Electrical receptive field stimulation, which bypasses the receptor end organ, was most effective in classifying C-mechanoreceptors, as responses to mechanical receptive field stimulation overlapped somewhat, which may lead to misclassification. Furthermore, we report a subclass of low-threshold CM responding to gentle mechanical stimulation and a potential subclass of CT afferent displaying burst firing. We show that substantial differences exist in the mechanisms governing axonal conduction between CTs and CMs. We provide clear electrophysiological "signatures" (extent of latency slowing) that can be used in unequivocally identifying populations of C-mechanoreceptors in single-unit and multiunit microneurography studies and in translational animal research into affective touch. Additionally, these differential mechanisms may be pharmacologically targetable for separate modulation of positive and negative affective touch information. NEW & NOTEWORTHY Human skin encodes a plethora of touch interactions, and affective tactile information is primarily signaled by slowly conducting C-mechanoreceptive afferents. We show that electrical stimulation of low-threshold C-tactile afferents produces markedly different patterns of activity compared with high-threshold C-mechanoreceptive nociceptors, although the populations overlap in their responses to mechanical stimulation. This fundamental distinction demonstrates a divergence in affective touch signaling from the first stage of sensory processing, having implications for the processing of interpersonal touch., (Copyright © 2017 the American Physiological Society.)

Our perception of skin wetness is generated readily, yet humans have no known receptor (hygroreceptor) to signal this directly. It is easy to imagine the sensation of water running over our hands or the feel of rain on our skin. The synthetic sensation of wetness is thought to be produced from a combination of specific skin thermal and tactile inputs, registered through thermoreceptors and mechanoreceptors, respectively. The present review explores how thermal and tactile afference from the periphery can generate the percept of wetness centrally. We propose that the main signals include information about skin cooling, signaled primarily by thinly myelinated thermoreceptors, and rapid changes in touch, through fast-conducting, myelinated mechanoreceptors. Potential central sites for integration of these signals, and thus the perception of skin wetness, include the primary and secondary somatosensory cortices and the insula cortex. The interactions underlying these processes can also be modeled to aid in understanding and engineering the mechanisms. Furthermore, we discuss the role that sensing wetness could play in precision grip and the dexterous manipulation of objects. We expand on these lines of inquiry to the application of the knowledge in designing and creating skin sensory feedback in prosthetics. The addition of real-time, complex sensory signals would mark a significant advance in the use and incorporation of prosthetic body parts for amputees in everyday life. NEW & NOTEWORTHY Little is known about the underlying mechanisms that generate the perception of skin wetness. Humans have no specific hygroreceptor, and thus temperature and touch information combine to produce wetness sensations. The present review covers the potential mechanisms leading to the perception of wetness, both peripherally and centrally, along with their implications for manual function. These insights are relevant to inform the design of neuroengineering interfaces, such as sensory prostheses for amputees., (Copyright © 2017 the American Physiological Society.)

Using ultra-high field 7 Tesla (7T) functional magnetic resonance imaging (fMRI), we map the cortical and perceptual responses elicited by intraneural microstimulation (INMS) of single mechanoreceptive afferent units in the median nerve, in humans. Activations are compared to those produced by applying vibrotactile stimulation to the unit's receptive field, and unit-type perceptual reports are analyzed. We show that INMS and vibrotactile stimulation engage overlapping areas within the topographically appropriate digit representation in the primary somatosensory cortex. Additional brain regions in bilateral secondary somatosensory cortex, premotor cortex, primary motor cortex, insula and posterior parietal cortex, as well as in contralateral prefrontal cortex are also shown to be activated in response to INMS. The combination of INMS and 7T fMRI opens up an unprecedented opportunity to bridge the gap between first-order mechanoreceptive afferent input codes and their spatial, dynamic and perceptual representations in human cortex.

Pleasant, affective touch provides various health benefits, including stress and depression relief. There is a dichotomy between mechanoreceptive afferents that predominantly signal discriminative (myelinated A-beta) and affective (unmyelinated C-tactile) aspects of touch. It is well documented that discriminative abilities of touch decline with age. However, a thorough investigation of how the pleasant aspects of touch develop with age has not been previously attempted. Here, we investigated the relationship between age and psychophysical ratings in response to gentle stroking touch. One hundred twenty participants (60 males, 60 females) ages 13-82 years were presented with C-tactile optimal and suboptimal stroking velocities, and rated pleasantness and intensity. Moreover, to examine the specificity of age effects on touch perception, we used olfactory stimuli as a cross-sensory comparison. For all ages, we found that C-tactile optimal stimuli were rated significantly more pleasant than C-tactile suboptimal stimuli. Although, both touch and olfactory intensity ratings were negatively correlated with age, a positive correlation between pleasantness ratings of touch (but not olfactory stimuli) and age was found. We conclude that the affective, but not the discriminative, aspects of touch are enhanced with increasing age. The increase of pleasantness of all touch stimuli in late adulthood is discussed in relation to cognitive modulations., ((c) 2016 APA, all rights reserved).)

The present review focusses on how tactile somatosensory afference is encoded and processed, and how this information is interpreted and acted upon in terms of motor control. We relate the fundamental workings of the sensorimotor system to the rehabilitation of amputees using modern prosthetic interventions. Our sense of touch is central to our everyday lives, from allowing us to manipulate objects accurately to giving us a sense of self-embodiment. There are a variety of specialised cutaneous mechanoreceptive afferents, which differ in terms of type and density according to the skin site. In humans, there is a dense innervation of our hands, which is reflected in their vast over-representation in somatosensory and motor cortical areas. We review the accumulated evidence from animal and human studies about the precise interplay between the somatosensory and motor systems, which is highly integrated in many brain areas and often not separable. The glabrous hand skin provides exquisite, discriminative detail about touch, which is useful for refining movements. When these signals are disrupted, such as through injury or amputation, the consequences are considerable. The development of sensory feedback in prosthetics offers a promising avenue for the full integration of a missing body part. Real-time touch feedback from motor intentions aids in grip control and the ability to distinguish different surfaces, even introducing the possibility of pleasure in artificial touch. Thus, our knowledge from fundamental research into sensorimotor interactions should be used to develop more realistic and integrative prostheses., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

It is not known how changes in skin mechanics affect the responses of cutaneous mechanoreceptors in the finger pads to compression forces. We used venous occlusion to change the stiffness of the fingers and investigated whether this influenced the firing of low-threshold mechanoreceptors to surfaces of differing stiffness. Unitary recordings were made from 10 slowly adapting type I (SAI), 10 fast adapting type I (FAI) and 9 slowly adapting type II (SAII) units via tungsten microelectrodes inserted into the median nerve at the wrist. A servo-controlled stimulator applied ramp-and-hold forces (1, 2, and 4 N) at a constant loading and unloading rate (2 N/s) via a flat 2.5-cm-diameter silicone disk over the center of the finger pad. Nine silicone disks (objects), varying in compliance, were used. Venous occlusion, produced by inflating a sphygmomanometer cuff around the upper arm to 40 ± 5 mmHg, was used to induce swelling of the fingers and increase the compliance of the finger pulp. Venous occlusion had no effect on the firing rates of the SAI afferents, nor on the slopes of the relationship between mean firing rate and object compliance at each amplitude, but did significantly reduce the slopes for the FAI afferents. Although the SAII afferents possess a poor capacity to encode changes in object compliance, mean firing rates were significantly lower during venous occlusion. The finding that venous occlusion had no effect on the firing properties of SAI afferents indicates that these afferents preserve their capacity to encode changes in object compliance, despite changes in skin mechanics., (Copyright © 2015 the American Physiological Society.)