Observed Touch on a Non-Human Face Is Not Remapped onto the Human Observer's Own Face

Visual remapping of touch (VRT) is a phenomenon in which seeing a human face being touched enhances detection of tactile stimuli on the observer''s own face, especially when the observed face expresses fear. This study tested whether VRT would occur when seeing touch on monkey faces and whether it would be similarly modulated by facial expressions. Human participants detected near-threshold tactile stimulation on their own cheeks while watching fearful, happy, and neutral human or monkey faces being concurrently touched or merely approached by fingers. We predicted minimal VRT for neutral and happy monkey faces but greater VRT for fearful monkey faces. The results with human faces replicated previous findings, demonstrating stronger VRT for fearful expressions than for happy or neutral expressions. However, there was no VRT (i.e. no difference between accuracy in touch and no-touch trials) for any of the monkey faces, regardless of facial expression, suggesting that touch on a non-human face is not remapped onto the somatosensory system of the human observer.     

Close to you: embodied simulation for peripersonal space in primary somatosensory cortex

Background

An increasing body of evidence has demonstrated that in contrast to the classic understanding the primary somatosensory cortex (SI) reflects merely seen touch (in the absence of any real touch on the own body). Based on these results it has been discussed that SI may play a role in understanding touch seen on other bodies. In order to further examine this understanding of observed touch, the current study aimed to test if mirror-like responses in SI are affected by the perspective of the seen touch. Thus, we presented touch on a hand and close to the hand either in first-person-perspective or in third-person-perspective.

Principal Findings

Results of functional magnetic resonance imaging (fMRI) revealed stronger vicarious brain responses in SI/BA2 for touch seen in first-person-perspective. Surprisingly, the third-person viewpoint revealed activation in SI both when subjects viewed a hand being stimulated as well as when the space close to the hand was being touched.

Conclusions/Significance

Based on these results we conclude that vicarious somatosensory responses in SI/BA2 are affected by the viewpoint of the seen hand. Furthermore, we argue that mirror-like responses in SI do not only reflect seen touch, but also the peripersonal space surrounding this body (in third-person-perspective). We discuss these findings with recent studies on mirror responses for action observation in peripersonal space.     

Proprioceptive alignment of visual and somatosensory maps in the posterior parietal cortex

A touch on one hand can enhance the response to a visual stimulus delivered at a nearby location [1, 2], improving our interactions with the external world. In order to keep such visual-tactile spatial interactions effective, the brain updates the continuous postural changes, like those typically accompanying hand actions, through proprioception, thus maintaining the somatosensory and visual maps in spatial register [2, 3]. The posterior parietal cortex (PPC) might be critical for such a spatial remapping [4]; nevertheless, a direct causal demonstration of its involvement is lacking. Here, we found that unattended touches to one hand enhanced visual sensitivity for phosphenes induced by occipital trancranial magnetic stimulation (TMS) [5] when the touched hand was spatially coincident to the reported location of the phosphenes in external space. Notably, this spatially specific crossmodal facilitation was maintained after hand crossing, suggesting an efficient visual-tactile remapping. Critically, after 1 Hz repetitive TMS interference [6] over the PPC, but not over the primary somatosensory cortex, phosphene detection was still enhanced by spatially coincident touches with uncrossed hands, but it was enhanced by spatially noncoincident touches after hand crossing. This is the first causal evidence in humans that the PPC constantly updates the representation of the body in space in order to facilitate crossmodal interactions.     

Racial Bias in Neural Empathic Responses to Pain

Recent studies have shown that perceiving the pain of others activates brain regions in the observer associated with both somatosensory and affective-motivational aspects of pain, principally involving regions of the anterior cingulate and anterior insula cortex. The degree of these empathic neural responses is modulated by racial bias, such that stronger neural activation is elicited by observing pain in people of the same racial group compared with people of another racial group. The aim of the present study was to examine whether a more general social group category, other than race, could similarly modulate neural empathic responses and perhaps account for the apparent racial bias reported in previous studies. Using a minimal group paradigm, we assigned participants to one of two mixed-race teams. We use the term race to refer to the Chinese or Caucasian appearance of faces and whether the ethnic group represented was the same or different from the appearance of the participant'' own face. Using fMRI, we measured neural empathic responses as participants observed members of their own group or other group, and members of their own race or other race, receiving either painful or non-painful touch. Participants showed clear group biases, with no significant effect of race, on behavioral measures of implicit (affective priming) and explicit group identification. Neural responses to observed pain in the anterior cingulate cortex, insula cortex, and somatosensory areas showed significantly greater activation when observing pain in own-race compared with other-race individuals, with no significant effect of minimal groups. These results suggest that racial bias in neural empathic responses is not influenced by minimal forms of group categorization, despite the clear association participants showed with in-group more than out-group members. We suggest that race may be an automatic and unconscious mechanism that drives the initial neural responses to observed pain in others.     

Seeing Touches Early in Life

The sense of touch provides fundamental information about the surrounding world, and feedback about our own actions. Although touch is very important during the earliest stages of life, to date no study has investigated infants’ abilities to process visual stimuli implying touch. This study explores the developmental origins of the ability to visually recognize touching gestures involving others. Looking times and orienting responses were measured in a visual preference task, in which participants were simultaneously presented with two videos depicting a touching and a no-touching gesture involving human body parts (face, hand) and/or an object (spoon). In Experiment 1, 2-day-old newborns and 3-month-old infants viewed two videos: in one video a moving hand touched a static face, in the other the moving hand stopped before touching it. Results showed that only 3-month-olds, but not newborns, differentiated the touching from the no-touching gesture, displaying a preference for the former over the latter. To test whether newborns could manifest a preferential visual response when the touched body part is different from the face, in Experiment 2 newborns were presented with touching/no-touching gestures in which a hand or an inanimate object—i.e., a spoon- moved towards a static hand. Newborns were able to discriminate a hand-to-hand touching gesture, but they did not manifest any preference for the object-to-hand touch. The present findings speak in favour of an early ability to visually recognize touching gestures involving the interaction between human body parts.     

Biological and bionic hands: natural neural coding and artificial perception

The first decade and a half of the twenty-first century brought about two major innovations in neuroprosthetics: the development of anthropomorphic robotic limbs that replicate much of the function of a native human arm and the refinement of algorithms that decode intended movements from brain activity. However, skilled manipulation of objects requires somatosensory feedback, for which vision is a poor substitute. For upper-limb neuroprostheses to be clinically viable, they must therefore provide for the restoration of touch and proprioception. In this review, I discuss efforts to elicit meaningful tactile sensations through stimulation of neurons in somatosensory cortex. I focus on biomimetic approaches to sensory restoration, which leverage our current understanding about how information about grasped objects is encoded in the brain of intact individuals. I argue that not only can sensory neuroscience inform the development of sensory neuroprostheses, but also that the converse is true: stimulating the brain offers an exceptional opportunity to causally interrogate neural circuits and test hypotheses about natural neural coding.     

Somatic and motor components of action simulation

Seminal studies in monkeys report that the viewing of actions performed by other individuals activates frontal and parietal cortical areas typically involved in action planning and execution. That mirroring actions might rely on both motor and somatosensory components is suggested by reports that action observation and execution increase neural activity in motor and in somatosensory areas. This occurs not only during observation of naturalistic movements but also during the viewing of biomechanically impossible movements that tap the afferent component of action, possibly by eliciting strong somatic feelings in the onlooker. Although somatosensory feedback is inherently linked to action execution, information on the possible causative role of frontal and parietal cortices in simulating motor and sensory action components is lacking. By combining low-frequency repetitive and single-pulse transcranial magnetic stimulation, we found that virtual lesions of ventral premotor cortex (vPMc) and primary somatosensory cortex (S1) suppressed mirror motor facilitation contingent upon observation of possible and impossible movements, respectively. In contrast, virtual lesions of primary motor cortex did not influence mirror motor facilitation. The reported double dissociation suggests that vPMc and S1 play an active, differential role in simulating efferent and afferent components of observed actions.     

Dynamic Changes in Brain Activations and Functional Connectivity during Affectively Different Tactile Stimuli

In the present study, we compared brain activations produced by pleasant, neutral and unpleasant touch, to the anterior lateral surface of lower leg of human subjects. It was found that several brain regions, including the contralateral primary somatosensory area (SI), bilateral secondary somatosensory area (SII), as well as contralateral middle and posterior insula cortex were commonly activated under the three touch conditions. In addition, pleasant and unpleasant touch conditions shared a few brain regions including the contralateral posterior parietal cortex (PPC) and bilateral premotor cortex (PMC). Unpleasant touch specifically activated a set of pain-related brain regions such as contralateral supplementary motor area (SMA) and dorsal parts of bilateral anterior cingulated cortex, etc. Brain regions specifically activated by pleasant touch comprised bilateral lateral orbitofrontal cortex (OFC), posterior cingulate cortex (PCC), medial prefrontal cortex (mPFC), intraparietal cortex and left dorsal lateral prefrontal cortex (DLPFC). Using a novel functional connectivity model based on graph theory, we showed that a series of brain regions related to affectively different touch had significant functional connectivity during the resting state. Furthermore, it was found that such a network can be modulated between affectively different touch conditions.     

Modality-specific control of strategic spatial attention in parietal cortex

The neural basis of selective spatial attention presents a significant challenge to cognitive neuroscience. Recent neuroimaging studies have suggested that regions of the parietal and temporal cortex constitute a "supramodal" network that mediates goal-directed attention in multiple sensory modalities. Here we used transcranial magnetic stimulation (TMS) to determine which cortical subregions control strategic attention in vision and touch. Healthy observers undertook an orienting task in which a central arrow cue predicted the location of a subsequent visual or somatosensory target. To determine the attentional role of cortical subregions at different stages of processing, TMS was delivered to the right hemisphere during cue or target events. Results indicated a critical role of the inferior parietal cortex in strategic orienting to visual events, but not to somatosensory events. These findings are inconsistent with the existence of a supramodal attentional network and instead provide direct evidence for modality-specific attentional processing in parietal cortex.     

Comparison of cortical activation patterns by somatosensory stimulation on the palm and dorsum of the hand

Abstract

Objectives: Little is known about differences of cortical activation according to body location. We attempted to compare brain activation patterns by somatosensory stimulation on the palm and dorsum of the hand, using functional magnetic resonance imaging (fMRI).

Method: We recruited 15 healthy right-handed volunteers for this study. fMRI was performed during touch stimulation using a rubber brush on an area of the same size on the palm or dorsum of the hand. Regions of interest (ROIs) were drawn at the primary sensory–motor cortex (SM1), posterior parietal cortex, and secondary somatosensory cortex.

Results: Group analysis of fMRI data indicated that touch stimulation on the palm resulted in production of more activated voxels in the contralateral SM1 and posterior parietal cortex than on the dorsum of the hand. The most activated ROI was found to be the contralateral SM1 by stimulation of the palm or dorsum, and the number of activated voxels (5875) of SM1 by palm stimulation was more than 2 times that (2282) of dorsum stimulation. The peak activated value in the SM1 by palm stimulation (16.43) was also higher than that of the dorsum (5.52).

Conclusion: We found that stimulation of the palm resulted in more cortical activation in the contralateral SM1 than stimulation of the dorsum. Our results suggested that the palm of the hand might have larger somatotopy of somatosensory representation for touch in the cerebral cortex than the dorsum of the hand. Our results would be useful as a rehabilitation strategy when more or less somatosensory stimulation of the hand is necessary.     

Cortical Plasticity Induced by Spike-Triggered Microstimulation in Primate Somatosensory Cortex

Electrical stimulation of the nervous system for therapeutic purposes, such as deep brain stimulation in the treatment of Parkinson’s disease, has been used for decades. Recently, increased attention has focused on using microstimulation to restore functions as diverse as somatosensation and memory. However, how microstimulation changes the neural substrate is still not fully understood. Microstimulation may cause cortical changes that could either compete with or complement natural neural processes, and could result in neuroplastic changes rendering the region dysfunctional or even epileptic. As part of our efforts to produce neuroprosthetic devices and to further study the effects of microstimulation on the cortex, we stimulated and recorded from microelectrode arrays in the hand area of the primary somatosensory cortex (area 1) in two awake macaque monkeys. We applied a simple neuroprosthetic microstimulation protocol to a pair of electrodes in the area 1 array, using either random pulses or pulses time-locked to the recorded spiking activity of a reference neuron. This setup was replicated using a computer model of the thalamocortical system, which consisted of 1980 spiking neurons distributed among six cortical layers and two thalamic nuclei. Experimentally, we found that spike-triggered microstimulation induced cortical plasticity, as shown by increased unit-pair mutual information, while random microstimulation did not. In addition, there was an increased response to touch following spike-triggered microstimulation, along with decreased neural variability. The computer model successfully reproduced both qualitative and quantitative aspects of the experimental findings. The physiological findings of this study suggest that even simple microstimulation protocols can be used to increase somatosensory information flow.     

Sensory-motor coordination during grasping and manipulative actions.

Goal-directed grasping and manipulation of objects are human skills that depend on automatic sensory control in which predictive feed-forward mechanisms integrate somatosensory and visual signals with sensory-motor memory systems. Memory representations of physical and task-relevant properties of the object play a pivotal role. Anticipatory strategies are crucial when purposeful actions arise from learned relationships between afferent patterns and efferent commands. The development of even elementary precision grip skills is a protracted process not concluded until early adolescence. Not surprisingly, the neural control of manual actions engages most central nervous system areas known to be involved in motor control.     

Looking for Myself: Current Multisensory Input Alters Self-Face Recognition

How do I know the person I see in the mirror is really me? Is it because I know the person simply looks like me, or is it because the mirror reflection moves when I move, and I see it being touched when I feel touch myself? Studies of face-recognition suggest that visual recognition of stored visual features inform self-face recognition. In contrast, body-recognition studies conclude that multisensory integration is the main cue to selfhood. The present study investigates for the first time the specific contribution of current multisensory input for self-face recognition. Participants were stroked on their face while they were looking at a morphed face being touched in synchrony or asynchrony. Before and after the visuo-tactile stimulation participants performed a self-recognition task. The results show that multisensory signals have a significant effect on self-face recognition. Synchronous tactile stimulation while watching another person''s face being similarly touched produced a bias in recognizing one''s own face, in the direction of the other person included in the representation of one''s own face. Multisensory integration can update cognitive representations of one''s body, such as the sense of ownership. The present study extends this converging evidence by showing that the correlation of synchronous multisensory signals also updates the representation of one''s face. The face is a key feature of our identity, but at the same time is a source of rich multisensory experiences used to maintain or update self-representations.     

Changing reference frames during the encoding of tactile events

The mindless act of swatting a mosquito on the hand poses a remarkable challenge for the brain. Given that the primary somatosensory cortex maps skin location independently of arm posture [1, 2], the brain must realign tactile coordinates in order to locate the origin of the stimuli in extrapersonal space. Previous studies have highlighted the behavioral relevance of such an external mapping of touch, which results from combining somatosensory input with proprioceptive and visual cues about body posture [3-7]. However, despite the widely held assumption about the existence of this remapping process from somatotopic to external space and various findings indirectly suggesting its consequences [8-11], a demonstration of its changing time course and nature was lacking. We examined the temporal course of this multisensory interaction and its implications for tactile awareness in humans using a crossmodal cueing paradigm [12, 13]. What we show is that before tactile events are referred to external locations [12-15], a fleeting, unconscious image of the tactile sensation abiding to a somatotopic frame of reference rules performance. We propose that this early somatotopic "glimpse" arises from the initial feed-forward sweep of neural activity to the primary somatosensory cortex, whereas the later externally-based, conscious experience reflects the activity of a somatosensory network involving recurrent connections from association areas.     

An improved method for isolating a section of the rat neocortex]

Neocortical island chronically isolated from surrounding cortical and subcortical structures with preserved pial blood supply has long been a model for research into the mechanisms of cortex functioning. To fully cut the cortex we improved the type of knife by using a retractable tungsten wire. The tip of a syringe needle was bent and cut away all but the beginning of the bend. In anesthetized rats the somatosensory cortex was exposed, the guide needle was lowered down to the desired depth into the cortex avoiding blood vessels. The wire then was pulled out through the curved needle tip until the tip of the wire touched the pia mater. The device was then raised, lowered, rotated to achieve complete separation of the cortical island from the surrounding tissues. The wire was retracted into the needle before withdrawal of the device. Analysis of neocortical slices 8 weeks later showed lesions of the white matter and transcortical cuts.     

对侧外周神经移位到损伤手臂引起的体感皮层功能动态重组

单侧肢体的外周神经损伤通常导致对侧体感皮层的功能重组. 然而,接受了对侧颈 7 (C7) 外周神经移位手术治疗单侧手臂臂丛全撕脱的病人,在术后早期当其患手被触摸时,只在其健手产生感觉. 在术后晚期,病人才逐渐恢复其患手和健手的正常、独立的功能. 我们在模拟对侧颈 7 (C7) 外周神经移位手术病例的大鼠模型上,用记录体感诱发电位的方法研究了患手和健手的体感代表区. 患手的体感和运动功能由于 C7 神经的再生而逐渐恢复. 术后第 5 个月始, 13 只大鼠患手的体感代表区只出现在其同侧的皮层,同时患手和健手的代表区在该皮层内是高度重叠的 (除掉一个例外),虽然刺激它们产生的体感诱发电位的潜伏期和反应幅度有很大的不同. 结果表明,移位到患手的对侧外周神经能够导致同侧体感皮层动态的功能重组,提示身体另侧感觉输入的介入激发了大脑显著的可塑性.     

Observation-based learning for brain-machine interfaces

Canonically, 'mirror neurons' are cells in area F5 of the ventral premotor cortex that are active during both observation and execution of goal-directed movements. Recently, cells with similar properties have been observed in a number of other areas in the motor system, including the primary motor cortex. Mirror neurons are a part of a system whose function is thought to involve the prediction and interpretation of the sensory consequences of our own actions as well as the actions of others. Mirror-like responses are relevant to the development of brain-machine interfaces (BMIs) because they provide a robust way to map neural activity to behavior, and because they represent high-level information about goals and intentions that may have utility in future BMI applications.     

Spatio‐temporal regulation of the formation of the somatosensory system

The somatosensory system in the brain has been widely used for investigating the mechanisms underlying neural circuit formation and developmental neural plasticity. In the primary somatosensory cortex (S1) of rodents, there are discrete cytoarchitectonic units called barrels. Reverse genetic analyses using knockout mice have revealed molecules that control spatial pattern formation of barrels in S1. Glutamatergic receptors such as the NMDA receptor and mGluR5, and molecules related to serotonin such as serotonin transporter and monoamine oxidase A are essential for the formation of barrels. In addition to the mechanisms of spatial pattern formation, those regulating the timing of developmental processes were uncovered recently. Barrels are formed soon after the birth of newborn mouse pups from their mothers, and it was shown that the timing of barrel formation was determined by the timing of the birth of mouse pups. The mechanisms downstream of birth were also examined. It would be intriguing to examine if the mechanisms found using the somatosensory system are applicable to other brain regions.