Somatosensory Representation Research Articles

Cortical representation of tympanic membrane movements due to pressure variation: An fMRI study

Middle ear sensory information has never been localized in the homunculus of the somatosensory cortex (S1). We investigated the somatosensory representation of the middle ear in 15 normal hearing subjects. We applied small air pressure variations to the tympanic membrane while performing a 3T-fMRI study. Unilateral stimulations of the right ear triggered bilateral activations in the caudal part of the postcentral gyrus in Brodmann area 43 (BA 43) and in the auditory associative areas 42 (BA 42) and 22 (BA 22). BA 43 has been found to be involved in activities accompanying oral intake and could be more largely involved in pressure activities in the oropharynx area. The tympanic membrane is indirectly related to the pharynx area through the action of tensor tympani, which is a Eustachian tube muscle. The Eustachian tube muscles have a role in pressure equalization in the middle ear and also have a role in the pharyngeal phase of swallowing. Activation of BA 42 and BA 22 could reflect activations associated with the bilateral acoustic reflex triggered prior to self-vocalization to adjust air pressure in the oropharynx during speech. We propose that BA 43, 42, and 22 are the cortical areas associated with middle ear function. We did not find representation of tympanic membrane movements due to pressure in S1, but its representation in the postcentral gyrus in BA 43 seems to suggest that at least part of this area conveys pure somatosensory information.

Social rejection shares somatosensory representations with physical pain

How similar are the experiences of social rejection and physical pain? Extant research suggests that a network of brain regions that support the affective but not the sensory components of physical pain underlie both experiences. Here we demonstrate that when rejection is powerfully elicited--by having people who recently experienced an unwanted break-up view a photograph of their ex-partner as they think about being rejected--areas that support the sensory components of physical pain (secondary somatosensory cortex; dorsal posterior insula) become active. We demonstrate the overlap between social rejection and physical pain in these areas by comparing both conditions in the same individuals using functional MRI. We further demonstrate the specificity of the secondary somatosensory cortex and dorsal posterior insula activity to physical pain by comparing activated locations in our study with a database of over 500 published studies. Activation in these regions was highly diagnostic of physical pain, with positive predictive values up to 88%. These results give new meaning to the idea that rejection "hurts." They demonstrate that rejection and physical pain are similar not only in that they are both distressing--they share a common somatosensory representation as well.

S33-03 - From mirror neurons to embodied simulation: A new neuroscientific perspective on intersubjectivity

Our seemingly effortless capacity of conceiving of the acting bodies inhabiting our social world as goal-oriented individuals like us depends on the constitution of a shared “we-centric” space. I have proposed that this shared manifold space can be characterized at the functional level as embodied simulation, a basic functional mechanism by means of which our brain/body system models its interactions with the world.The mirroring mechanism for action and other mirroring mechanisms in our brain represent sub-personal instantiations of embodied simulation. Embodied simulation provides a new empirically based notion of intersubjectivity, viewed first and foremost as intercorporeity. Embodied simulation challenges the notion that Folk-psychology is the sole account of interpersonal understanding. Before and below mind reading is intercorporeity as the main source of knowledge we directly gather about others.By means of embodied simulation we can map others’ actions onto our own motor representations, as well as others’ emotions and sensations onto our own viscero-motor and somatosensory representations. “Representation”, as used here, refers to a particular type of content, generated by the relations that our situated and inter-acting brain-body system instantiates with the world. Such content is pre-linguistic and pre-theoretical, but nevertheless has attributes normally and uniquely attributed to conceptual content.Social cognition is not only explicitly reasoning about the contents of someone else's mind. Embodied simulation, gives us a direct insight of other minds thus enabling our capacity to empathize with others.This proposal opens new perspectives on our understanding of autism and other psychopathological states such as schizophrenia.

Mislocalization of near‐threshold tactile stimuli in humans: a central or peripheral phenomenon?

Principles of brain function can be disclosed by studying their limits during performance. Tactile stimuli with near-threshold intensities have been used to assess features of somatosensory processing. When stimulating fingers of one hand using near-threshold intensities, localization errors are observed that deviate significantly from responses obtained by guessing - incorrectly located stimuli are attributed more often to fingers neighbouring the stimulated one than to more distant fingers. Two hypotheses to explain the findings are proposed. The 'central hypothesis' posits that the degree of overlap of cortical tactile representations depends on stimulus intensity, with representations less separated for near-threshold stimuli than for suprathreshold stimuli. The 'peripheral hypothesis' assumes that systematic mislocalizations are due to activation of different sets of skin receptors with specific thresholds. The present experiments were designed to decide between the two hypotheses. Taking advantage of the frequency tuning of somatosensory receptors, their contribution to systematic misclocalizations was studied. In the first experiment, mislocalization profiles were investigated using vibratory stimuli with frequencies of 10, 20 and 100 Hz. Unambiguous mislocalization effects were only obtained for the 10-Hz stimulation, precluding the involvement of Pacinian corpuscles in systematic mislocalization. In the second experiment, Pacinian corpuscles were functionally eliminated by applying a constant 100-Hz vibratory masking stimulus together with near-threshold pulses. Despite masking, systematic mislocation patterns were observed rendering the involvement of Pacinian corpuscles unlikely. The results of both experiments are in favor of the 'central hypothesis' assuming that the extent of overlap in somatosensory representations is modulated by stimulus intensity.

An Update on the Pathophysiology of Complex Regional Pain Syndrome

Complex regional pain syndrome (CRPS) is a neuropathic pain disorder with significant autonomic features. Few treatments have proven effective, in part, because of a historically poor understanding of the mechanisms underlying the disorder. CRPS research largely conducted during the past decade has substantially increased knowledge regarding its pathophysiologic mechanisms, indicating that they are multifactorial. Both peripheral and central nervous system mechanisms are involved. These include peripheral and central sensitization, inflammation, altered sympathetic and catecholaminergic function, altered somatosensory representation in the brain, genetic factors, and psychophysiologic interactions. Relative contributions of the mechanisms underlying CRPS may differ across patients and even within a patient over time, particularly in the transition from "warm CRPS" (acute) to "cold CRPS" (chronic). Enhanced knowledge regarding the pathophysiology of CRPS increases the possibility of eventually achieving the goal of mechanism-based CRPS diagnosis and treatment.

An implicit body representation underlying human position sense

Knowing the body's location in external space is a fundamental perceptual task. Perceiving the location of body parts through proprioception requires that information about the angles of each joint (i.e., body posture) be combined with information about the size and shape of the body segments between joints. Although information about body posture is specified by on-line afferent signals, no sensory signals are directly informative about body size and shape. Thus, human position sense must refer to a stored body model of the body's metric properties, such as body part size and shape. The need for such a model has long been recognized; however, the properties of this model have never been systematically investigated. We developed a technique to isolate and measure this body model. Participants judged the location in external space of 10 landmarks on the hand. By analyzing the internal configuration of the locations of these points, we produced implicit maps of the mental representation of hand size and shape. We show that this part of the body model is massively distorted, in a reliable and characteristic fashion, featuring shortened fingers and broadened hands. Intriguingly, these distortions appear to retain several characteristics of primary somatosensory representations, such as the Penfield homunculus.

P1.072 Mapping motor and somatosensory cortical flnger representation in focal hand dystonia

P1.072 Mapping motor and somatosensory cortical flnger representation in focal hand dystonia

An Easy and Practical Method for Routine, Bedside Testing of Somatosensory Systems in Extremely Low Birth Weight Infants

This study was set out to develop and describe a novel, simple, and safe method for routine bedside testing of somatosensory system in very early preterm infants. We recorded electroencephalogram (EEG) activity after tactile stimulation of hand (palm) and foot (sole) by a soft hairbrush stimulator in extremely low birth weight infants (n = 10; GA, 24-28, recording at conceptional age 30-32 wk) and compared with the raw EEG responses to those seen by one- or two-channel brain monitors. In every subject, single tactile stimuli produced prominent (100-350 microV) somatosensory evoked responses (SERs) that were readily identified in the ongoing EEG signal. The maximal SER was in the contralateral hemisphere at around the corresponding somatosensory representation areas. Conventional EEG filtering did significantly reduce the SERs, but they could still be identified in the routine brain monitor setting widely available in NICUs. The method described here is directly applicable to assessment of integrity of somatosensory system in the early preterm period. It needs minimal training and requires an EEG system or a brain monitor device that is available in most units. Thus, the technique is likely to open a novel window to neurologic assessment of these babies.

Differential tactile and motor recovery and cortical map alteration after C4–C5 spinal hemisection

After incomplete spinal cord injury (SCI), the adult central nervous system is spontaneously capable of substantial reorganizations that can underlie functional recovery. Most studies have focused on intraspinal reorganizations after SCI and not on the correlative cortical remodeling. Yet, differential studies of neural correlates of the recovery of sensory and motor abilities may be conducted by segregating motor and somatosensory representations in distinct and topologically organized primary cortical areas. This study was aimed at evaluating the effects of a cervical (C4–C5) spinal cord hemisection on sensorimotor performances and electrophysiological maps in primary somatosensory (S1) and motor (M1) cortices in adult rats. After SCI, an enduring loss of the affected forepaw tactile sensitivity was paralleled by the abolishment of somatosensory evoked responses in the deprived forepaw area within the S1 cortex. In contrast, severe motor deficits in unilateral forelimb were partially restored over the first postoperative month, despite remnant deficits in distal movement. The overall M1 map size was drastically reduced in SCI rats relative to intact rats. In the remaining M1 map, the shoulder and elbow movements were over-represented, consistent with the behavioral recovery of proximal joint movements in almost all rats. By contrast, residual wrist representations were observed in M1 maps of half of the rats that did not systematically correlate with a behavioral recovery of these joint movements. This study highlights the differential potential of ascending and descending pathways to reorganize after SCI.

Remapping of the somatosensory cortex after a photothrombotic stroke: dynamics of the compensatory reorganization

Although stroke can affect cerebral structure and function, the brain has a potential for plasticity thanks to which some degree of function can be restored. The pathways of such recovery are of great interest, since the dynamics of rewiring of the injured brain may become the basis for designing appropriate strategies of rehabilitation. We investigated the spontaneous plasticity of cortical somatosensory representations following a focal unilateral stroke in the barrel cortex of rats. Ischemic lesions were produced with the photothrombotic technique in the cortical representation of vibrissae. Functional activation of the brain in response to the stimulation of vibrissae with destroyed cortical representation was monitored through the 2 months post-stroke survival period with [14C] 2-deoxyglucose (2DG) autoradiographic brain mapping (1, 7, 28, 56 days after the stroke). 2DG uptake was measured on autoradiograms of tangential sections in several regions of somatosensory cortex and in motor, auditory and prefrontal cortex. Behavioral deficit was assessed by the gap-crossing test 3, 28, 56 days after the stroke. Changes in the activation pattern of the intact hemisphere and non-vibrissal somatosensory representations of the lesioned hemisphere evolved during the observation period. Full recovery of the behavioral function was reached 2 months after the stroke and at the same time, new foci of activation were observed in the lesioned hemisphere. At that time, hyperactivation of the somatosensory areas in the intact hemisphere subsided. The new activation foci located in representations of anterior vibrissae, front paw and hind paw were specific for the vibrissae stimulation and were most probably a new functional representation of the vibrissae. We demonstrated spatial and temporal remodeling of the brain induced by cortical stroke, leading to vicariation of function.

From maps to form to space: Touch and the body schema

Evidence from patients has shown that primary somatosensory representations are plastic, dynamically changing in response to central or peripheral alterations, as well as experience. Furthermore, recent research has also demonstrated that altering body posture results in changes in the perceived sensation and localization of tactile stimuli. Using evidence from behavioral studies with brain-damaged and healthy subjects, as well as functional imaging, we propose that the traditional concept of the body schema should be divided into three components. First are primary somatosensory representations, which are representations of the skin surface that are typically somatotopically organized, and have been shown to change dynamically due to peripheral (usage, amputation, deafferentation) or central (lesion) modifications. Second, we argue for a mapping from a primary somatosensory representation to a secondary representation of body size and shape ( body form representation). Finally, we review evidence for a third set of representations that encodes limb position and is used to represent the location of tactile stimuli relative to the subject using external, non-somatotopic reference frames ( postural representations).

The somatosensory representation of the human clitoris: An fMRI study

We studied the central representation of pudendal afferents arising from the clitoral nerves in 15 healthy adult female subjects using electrical dorsal clitoral nerve stimulation and fMRI. As a control body region, we electrically stimulated the right hallux in eight subjects. In a block design experiment, we applied bilateral clitoral stimulation and unilateral (right) hallux stimulation. Activation maps were calculated for the contrasts ‘electrical dorsal clitoral nerve stimulation versus rest’ and ‘electrical hallux stimulation versus rest’.A random-effect group analysis for the clitoral stimulation showed significant activations bilateral in the superior and inferior frontal gyri, insulae and putamen and in the postcentral, precentral and inferior parietal gyri (including the primary and secondary somatosensory cortices). No activation was found on the mesial surface of the postcentral gyrus. For the hallux, activations occurred in a similar neuronal network but the activation in the primary somatosensory cortex was localized in the inter-hemispheric fissure.The results of this study demonstrate that the central representation of pudendal afferents arising from the clitoral nerves and sensory inputs from the hallux can be studied and distinguished from each other by fMRI. From the somatotopic order described in the somatosensory homunculus one would expect for electrical clitoral nerve stimulation activation of the mesial wall of the postcentral gyrus. In contrast, we found activations on the lateral surface of the postcentral gyrus.

Is This Hand for Real? Attenuation of the Rubber Hand Illusion by Transcranial Magnetic Stimulation over the Inferior Parietal Lobule

In the rubber hand illusion (RHI), participants incorporate a rubber hand into a mental representation of one's body. This deceptive feeling of ownership is accompanied by recalibration of the perceived position of the participant's real hand toward the rubber hand. Neuroimaging data suggest involvement of the posterior parietal lobule during induction of the RHI, when recalibration of the real hand toward the rubber hand takes place. Here, we used off-line low-frequency repetitive transcranial magnetic stimulation (rTMS) in a double-blind, sham-controlled within-subjects design to investigate the role of the inferior posterior parietal lobule (IPL) in establishing the RHI directly. Results showed that rTMS over the IPL attenuated the strength of the RHI for immediate perceptual body judgments only. In contrast, delayed perceptual responses were unaffected. Furthermore, ballistic action responses as well as subjective self-reports of feeling of ownership over the rubber hand remained unaffected by rTMS over the IPL. These findings are in line with previous research showing that the RHI can be broken down into dissociable bodily sensations. The illusion does not merely affect the embodiment of the rubber hand but also influences the experience and localization of one's own hand in an independent manner. Finally, the present findings concur with a multicomponent model of somatosensory body representations, wherein the IPL plays a pivotal role in subserving perceptual body judgments, but not actions or higher-order affective bodily judgments.

BOLD signal in insula is differentially related to cardiac function during compassion meditation in experts vs. novices

The brain and the cardiovascular system influence each other during the processing of emotion. The study of the interactions of these systems during emotion regulation has been limited in human functional neuroimaging, despite its potential importance for physical health. We have previously reported that mental expertise in cultivation of compassion alters the activation of circuits linked with empathy and theory of mind in response to emotional stimuli. Guided by the finding that heart rate increases more during blocks of compassion meditation than neutral states, especially for experts, we examined the interaction between state (compassion vs. neutral) and group (novice, expert) on the relation between heart rate and BOLD signal during presentation of emotional sounds presented during each state. Our findings revealed that BOLD signal in the right middle insula showed a significant association with heart rate (HR) across state and group. This association was stronger in the left middle/posterior insula when experts were compared to novices. The positive coupling of HR and BOLD was higher within the compassion state than within the neutral state in the dorsal anterior cingulate cortex for both groups, underlining the role of this region in the modulation of bodily arousal states. This state effect was stronger for experts than novices in somatosensory cortices and the right inferior parietal lobule (group by state interaction). These data confirm that compassion enhances the emotional and somatosensory brain representations of others' emotions, and that this effect is modulated by expertise. Future studies are needed to further investigate the impact of compassion training on these circuits.

Axon Morphologies and Convergence Patterns of Projections from Different Sensory-Specific Cortices of the Anterior Ectosylvian Sulcus onto Multisensory Neurons in the Cat Superior Colliculus

Corticofugal projections to the thalamus reveal 2 axonal morphologies, each associated with specific physiological attributes. These determine the functional characteristics of thalamic neurons. It is not clear, however, whether such features characterize the corticofugal projections that mediate multisensory integration in superior colliculus (SC) neurons. The cortico-collicular projections from cat anterior ectosylvian sulcus (AES) are derived from its visual, auditory, and somatosensory representations and are critical for multisensory integration. Following tracer injections into each subdivision, 2 types of cortico-collicular axons were observed. Most were categorized as type I and consisted of small-caliber axons traversing long distances without branching, bearing mainly small boutons. The less frequent type II had thicker axons, more complex branching patterns, larger boutons, and more complex terminal boutons. Following combinatorial injections of 2 different fluorescent tracers into defined AES subdivisions, fibers from each were seen converging onto individual SC neurons and indicate that such convergence, like that in the corticothalamic system, is mediated by 2 distinct morphological types of axon terminals. Nevertheless, and despite the conservation of axonal morphologies across different subcortical systems, it is not yet clear if the concomitant physiological attributes described in the thalamus are directly applicable to multisensory integration.

Tactile co-activation improves detection of afferent spatial modulation.

Tactile co-activation, i.e., synchronous stimulation of a region of skin, has been reported to improve tactile spatial acuity and expand the corresponding somatosensory cortical representation. The current study aimed to clarify the nature of the changes resulting from tactile co-activation, using three measures of tactile sensitivity obtained with controlled mechanical stimulation. One was the grating orientation (GR/OR) discrimination task, where acuity is indexed by the threshold groove width required for 75% correct discrimination between two orthogonal orientations of a grating on the fingerpad. Since this task may be susceptible to intensity cues due to tactile anisotropy, another acuity measure, the 3-dot task, was also used. In this task, the acuity threshold corresponds to 75% correct discrimination of the direction of offset of the central dot in a 3-dot array. In Experiment 1, co-activation failed to induce significant improvement in acuity with either of these measures. Experiment 2 employed both the GR/OR task, and a third measure based on discriminating a grooved from a smooth surface (SM/GV). While the former task demands detailed spatial resolution, the latter requires only that spatial modulation in the afferent population be detected. This experiment also included a control group. GR/OR performance did not significantly improve for either the control or experimental groups. There was, however, a significant improvement in SM/GV performance following co-activation for the experimental but not the control group. These findings indicate that the SM/GV task may be better suited than the GR/OR or 3-dot tasks for measuring changes in tactile sensitivity following co-activation.

Can We Share a Pain We Never Felt? Neural Correlates of Empathy in Patients with Congenital Insensitivity to Pain

Theories of empathy differ regarding the relative contributions of automatic resonance and perspective taking in understanding others' emotions. Patients with the rare syndrome of congenital insensitivity to pain cannot rely on "mirror matching" (i.e., resonance) mechanisms to understand the pain of others. Nevertheless, they showed normal fMRI responses to observed pain in anterior mid-cingulate cortex and anterior insula, two key regions of the so-called "shared circuits" for self and other pain. In these patients (but not in healthy controls), empathy trait predicted ventromedial prefrontal responses to somatosensory representations of others' pain and posterior cingulate responses to emotional representations of others' pain. These findings underline the major role of midline structures in emotional perspective taking and understanding someone else's feeling despite the lack of any previous personal experience of it--an empathic challenge frequently raised during human social interactions.

Somatosensory evoked potentials and high-frequency oscillations in athletes

Objective Athletes perform skilled movements during games and daily training. We hypothesized that the cortical representation in athletes differs from that in non-athletes. Methods Somatosensory evoked potentials (SEPs) and high-frequency oscillations (HFOs) were recorded from seven healthy football players, seven healthy racquet players and seven healthy non-athletes. Electrical stimuli were delivered to the posterior tibial nerves and the median nerves, bilaterally. Cortical and spinal SEPs and sensory nerve action potentials (SNAPs) were recorded. SEPs were recorded by 0.3–3000 Hz filter. HFOs were separated by 400–800 Hz band-pass filtering. SNAPs were recorded by 20–2000 Hz filter. Results The P37–N45 amplitude in football players and the N20–P25 amplitude in racquet players were significantly larger than those in non-athletes. The number of negative peaks of HFOs from the posterior tibial nerve in football players and the HFO amplitudes from the median nerve in racquet players were significantly larger than those in non-athletes. The earlier an individual started playing football, the larger the P37–N45 amplitude. Neither spinal SEPs nor SNAPs differed significantly among the three groups. Conclusions Daily long-term training brings about plastic excitation in the somatosensory cortex representation of the trained limbs in athletes. Significance Plastic changes in the somatosensory cortex are induced specifically by physical training.

Postura da mão e imagética motora: um estudo sobre reconhecimento de partes do corpo

OBJECTIVE: Recognition of body parts activates specific somatosensory representations in a way that is similar to motor imagery. These representations are implicitly activated to compare the body with the stimulus. In the present study, we investigate the influence of proprioceptive information relating to body posture on the recognition of body parts (hands). It proposes that this task could be used for rehabilitation of neurological patients. METHODS: Ten right-handed volunteers participated in this experiment. The task was to recognize the handedness of drawings of a hand that were presented in different perspectives and several orientations. For drawings of a right hand, the volunteers pressed the right key, and for drawings of a left hand, they pressed the left key. The volunteers underwent two sessions: one with their hands in a prone posture and the other with their hands in a supine posture. RESULTS: The manual reaction time was longer for perspectives and orientations for which the real movement was difficult to achieve. This showed that, during the task, motor representations were activated to compare the body with the stimulus. Furthermore, the subject's posture had an influence in relation to specific perspectives and orientations. CONCLUSIONS: These results showed that motor representations are activated to compare the body with the stimulus, and that the position of the hand influences this resonance between the stimulus and the body part.

Relationship between the types of behavioral reaction in a conditioned shake-off reflex and neocortical lateralization

The nature of the spike activity and interhemisphere interactions of neurons in the parietal cortex (somatosensory representation of the ear) during the intersignal period before active shake-off and passive (freezing) reactions to stimuli was studied in rabbits after acquisition of a conditioned defensive escape reflex. Before passive reactions, in contrast to the situation before active shake-off, there were increases in right-sided influences on cells in the left hemisphere with delays of up to 100 msec, leading to asymmetry in interhemisphere interactions with right-sided dominance. Passive reactions were preceded by increases in the extent of frequencies in the delta range in the spike activity of individual neurons and in interactions between pairs of cells. These data provide evidence that the existence and nature of interhemisphere asymmetry before stimulation is one of the factors determining the passive or active type of behavioral reaction and reflecting the level of activation of the cortex.