Human Somatosensory Cortex Research Articles

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.

31. The response of the human cerebral somatosensory cortex to mechanical stimulation

31. The response of the human cerebral somatosensory cortex to mechanical stimulation

Reward facilitates tactile judgments and modulates hemodynamic responses in human primary somatosensory cortex

Reinforcing effects of reward on action are well established, but possible effects on sensory function are less well explored. Here, using functional magnetic resonance imaging, we assessed whether reward can influence somatosensory judgments and modulate activity in human somatosensory cortex. Participants discriminated electrical somatosensory stimuli on an index finger with correct performance rewarded financially at trial end, at one of four different anticipated levels. Higher rewards improved tactile performance and led to increased hemodynamic signals from ventral striatum on rewarded trials. Remarkably, primary somatosensory cortex contralateral to the judged hand was reactivated at the point of reward delivery, despite the absence of concurrent somatosensory input at that time point. This side-specific reactivation of primary somatosensory cortex increased monotonically with level of reward. Moreover, the level of reward received on a particular trial influenced somatosensory performance and neural activity on the subsequent trial, with better discrimination and enhanced hemodynamic response in contralateral primary somatosensory cortex for trials that followed higher rewards. These results indicate that rewards can influence not only classical reward-related regions, but also early somatosensory cortex when a decision is required for that modality.

FMRI reflects functional connectivity of human somatosensory cortex

Unilateral sensory stimulation reliably elicits contralateral somatotopic activation of primary (SI) and secondary (SII) somatosensory cortex. There is an ongoing debate about the occurrence and nature of concomitant ipsilateral SI and SII activation. Here we used functional magnetic resonance imaging (fMRI) in healthy human subjects with unilateral tactile stimulation of fingers and lips, to compare somatosensory activation patterns from distal and proximal body parts. We hypothesized that fMRI in humans should reflect the functional connectivity of somatosensory cortex as predicted by animal studies. We show that both unilateral finger and lip stimulations activate contra- and ipsilateral SI and SII cortices with high detection frequency. Correlations of BOLD-signals to the applied hemodynamic reference function were significantly higher in contralateral as compared to ipsilateral SI and SII cortices for both finger and lip stimulation, reflecting strong contribution of contralateral thalamocortical input. Furthermore, BOLD-signal correlations were higher in SI than in SII activations on the contralateral but not on the ipsilateral side. While these asymmetries within and across hemispheres were consistent for finger and lip stimulations, indicating analogous underlying organizing principles, they were less prominent for lip stimulation. Somatotopic organization was detected in SI but not in SII representations of fingers and lips. These results qualitatively and quantitatively support the prevalent concepts of anatomical and functional connectivity in the somatosensory system and therefore may allow interpretation of sensory evoked fMRI signals in terms of normal human brain function. Thus, the assessment of human somatosensory function with fMRI may permit in the future investigations of pathological conditions.

Oscillatory activity in the human somatosensory cortex is modified by expectancy

Abstract Using magnetoencephalography (MEG), we examined region-specific alterations in oscillatory activity to investigate the neural basis of predictive responding in somatosensory processing when subjects were presented with simple patterns of tactile stimuli that varied in predictability. Here we showed event-related changes in beta rhythm in primary somatosensory cortex (SI) in response to a somatosensory stimulus accompanied by transient suppressions (desynchrony) followed by rebounds above baseline levels. We found that similar beta transient rebound occurred during a ‘gap period’, in the absence of a stimulus input, but only when the gap was part of a predictable pattern. Our results provide evidence that SI exhibits stimulus-driven and top–down oscillatory responses that can be modified by experience.

Discrimination of multiple sources in the somatosensory area of human finger based on 3-D SEF measurement

Abstract To discriminate multiple sources of human hand somatosensory cortex we recorded somatosensory evoked field after stimulation of three fingers (the thumb, fourth finger, and little finger), respectively. Time-frequency analysis (wavelet transforms, WT) was applied to the SEF data to examine the frequency component of each SEF data. Dominant power spectrum to the thumb (median nerve) stimulation appeared at higher frequency range than that of the little finger (ulnar nerve) stimulation (thumb: 15–20 Hz, little finger: 10–15 Hz). The frequency of dominant power spectrum to the fourth finger stimulation existed between that of the thumb and little finger stimulation. From the result of the coherence by the SEF data between the fourth finger and the other finger, band-pass filtering for the fourth finger was done with different frequency ranges (13–15 Hz: little finger and 16–18 Hz: the thumb). Source localization was done with these SEF data. It was confirmed that the proposed method, combining time-frequency analysis with the inverse problem of MEG (magnetoencephalogram), is useful for discriminating multiple sources of the SEF elicited by two nerve activities in the fourth finger overlapping in time.

Studying the human somatosensory hand area: A new way to compare fMRI and MEG

Valid localization is a prerequisite to study plasticity of the somatosensory cortex in humans. We compared the localizations of left and right thumb and little finger in the primary somatosensory cortex obtained with fMRI and MEG. Representations were investigated in 11 healthy right-handed subjects using echoplanar fMRI and 122-channel MEG together with electric finger stimulation. Activation observed with fMRI was based on an increase in the BOLD signal. Most of the activation clusters (71.1%) were located on the lateral surface of the postcentral gyrus. Representations of thumb and little finger were 17mm apart on average and consistently showed a somatotopic arrangement with the thumb representation inferior, lateral, and anterior to the representation of the little finger. Activation observed with MEG was modelled by equivalent current dipoles. Dipole localization was compatible with an assumed origin of activation within the posterior wall of the central sulcus. The Euclidian distance between corresponding dipoles was 11.5mm on average with deviations from the expected spatial arrangement of 35, 30, and 20% in the x-, y- und z-direction, respectively. Our study demonstrates how relative localization of somatosensory activations can serve as an indicator for localization validity when comparing different methods or studying somatosensory plasticity.

LTP‐like changes induced by paired associative stimulation of the primary somatosensory cortex in humans: source analysis and associated changes in behaviour

Paired associative stimulation (PAS), which combines repetitive peripheral nerve stimulation with transcranial magnetic stimulation (TMS), may induce neuroplastic changes in somatosensory cortex (S1), possibly by long-term potentiation-like mechanisms. We used multichannel median nerve somatosensory evoked potential (MN-SSEP) recordings and two-point tactile discrimination testing to examine the location and behavioural significance of these changes. When TMS was applied to S1 near-synchronously to an afferent signal containing mechanoreceptive information, MN-SSEP changes (significant at 21-31 ms) could be explained by a change in a tangential source located in Brodmann area 3b, with their timing and polarity suggesting modification of upper cortical layers. PAS-induced MN-SSEP changes between 28 and 32 ms were linearly correlated with changes in tactile discrimination. Conversely, when the near-synchronous afferent signal contained predominantly proprioceptive information, PAS-induced MN-SSEP changes (20-29 ms) were shifted medially, and tactile performance remained stable. With near-synchronous mechanoreceptive stimulation subtle differences in the timing of the two interacting signals tended to influence the direction of tactile performance changes. PAS performed with TMS delivered asynchronously to the afferent pulse did not change MN-SSEPs. Hebbian interaction of mechanoreceptive afferent signals with TMS-evoked activity may modify synaptic efficacy in superficial cortical layers of Brodmann area 3b and is associated with timing-dependent and qualitatively congruent behavioural changes.

Spatiotemporal integration of tactile information in human somatosensory cortex

BackgroundOur goal was to examine the spatiotemporal integration of tactile information in the hand representation of human primary somatosensory cortex (anterior parietal somatosensory areas 3b and 1), secondary somatosensory cortex (S2), and the parietal ventral area (PV), using high-resolution whole-head magnetoencephalography (MEG). To examine representational overlap and adaptation in bilateral somatosensory cortices, we used an oddball paradigm to characterize the representation of the index finger (D2; deviant stimulus) as a function of the location of the standard stimulus in both right- and left-handed subjects.ResultsWe found that responses to deviant stimuli presented in the context of standard stimuli with an interstimulus interval (ISI) of 0.33s were significantly and bilaterally attenuated compared to deviant stimulation alone in S2/PV, but not in anterior parietal cortex. This attenuation was dependent upon the distance between the deviant and standard stimuli: greater attenuation was found when the standard was immediately adjacent to the deviant (D3 and D2 respectively), with attenuation decreasing for non-adjacent fingers (D4 and opposite D2). We also found that cutaneous mechanical stimulation consistently elicited not only a strong early contralateral cortical response but also a weak ipsilateral response in anterior parietal cortex. This ipsilateral response appeared an average of 10.7 ± 6.1 ms later than the early contralateral response. In addition, no hemispheric differences either in response amplitude, response latencies or oddball responses were found, independent of handedness.ConclusionOur findings are consistent with the large receptive fields and long neuronal recovery cycles that have been described in S2/PV, and suggest that this expression of spatiotemporal integration underlies the complex functions associated with this region. The early ipsilateral response suggests that anterior parietal fields also receive tactile input from the ipsilateral hand. The lack of a hemispheric difference in responses to digit stimulation supports a lack of any functional asymmetry in human somatosensory cortex.

Localization of Palatal Area in Human Somatosensory Cortex

To determine the 'hard palate representing' area in the primary somatosensory cortex, we recorded somatosensory-evoked magnetic fields from the cortex in ten healthy volunteers, using magnetoencephalography. Following electrical stimulation of 3 sites on the hard palate (the first and third transverse palatine ridges, and the greater palatine foramen), magnetic responses showed peak latencies of 15, 65, and 125 ms. Equivalent current dipoles for early magnetic responses were found along the posterior wall of the inferior part of the central sulcus. These dipoles were localized anterior-inferiorly, compared with locations for the hand area in the cortex. However, there were no significant differences in three-dimensional locations among the 3 selected regions for hard palate stimulation. These results demonstrated the precise location of palatal representation in the primary somatosensory cortex, the actual area being small.

Neural activities in human somatosensory cortical areas evoked by acupuncture stimulation

To investigate neural representation evoked by acupuncture from human somatosensory cortices, especially from primary (SI) and secondary (SII) somatosensory areas. Neuroimaging study - Blood-oxygenation-level-dependent (BOLD) functional MRI was performed during acupuncture on LI4 (n=12 healthy participants). Sham acupuncture and innocuous tactile stimulation were also applied on the same acupuncture site as control comparisons. Responsive neural substrates were visualized and identified based on both individual and group-level surface activation maps. Discrete regions within the precentral gyrus (area 4) and the fundus of the central sulcus (area 3a) were selectively activated during the real acupuncture stimulation. In SII, the activation was extended in a postero-inferior direction to the fundus of the lateral sulcus. This specific pattern of acupuncture-related activation indicates that deep tissue stimulation (as seen in area 3a activation) and concurrent processing of sensory stimulation (as seen in activation in SII) may mediate neural responses to manual acupuncture.

Differential effects of high and low-frequency TMS on different tactile perceptual tasks and cortical excitability of human somatosensory cortex

Differential effects of high and low-frequency TMS on different tactile perceptual tasks and cortical excitability of human somatosensory cortex

Empathy for Pain and Touch in the Human Somatosensory Cortex

Although feeling pain and touch has long been considered inherently private, recent neuroimaging and neurophysiological studies hint at the social implications of this experience. Here we used somatosensory-evoked potentials (SEPs) to investigate whether mere observation of painful and tactile stimuli delivered to a model would modulate neural activity in the somatic system of an onlooker. Viewing video clips showing pain and tactile stimuli delivered to others, respectively, increased and decreased the amplitude of the P45 SEP component that reflects the activity of the primary somatosensory cortex (S1). These modulations correlated with the intensity but not with the unpleasantness of the pain and touch ascribed to the model or the aversion induced in the onlooker by the video clips. Thus, modulation of S1 activity contingent upon observation of others' pain and touch may reflect the mapping of sensory qualities of observed painful and tactile stimuli. Results indicate that the S1 is not only involved in the actual perception of pain and touch but also plays an important role in extracting somatic features from social interactions.

Transcranial direct current stimulation applied over the somatosensory cortex – Differential effect on low and high frequency SEPs

Objective Transcranial direct current stimulation (tDCS) has an influence on the excitability of the human motor cortex measured by motor evoked potentials (MEPs) after transcranial magnetic stimulation. Low and high frequency (HFOs) components of somatosensory evoked potentials (SEPs) were studied questioning whether a comparable effect can be observed after applying tDCS to the human somatosensory cortex. Methods Multichannel median nerve SEPs were recorded before and after applying tDCS of 1 mA over a period of 9 min with the cathode placed over the somatosensory cortex and the anode over the contralateral forehead and vice versa in a second session. The source activity of the N20, N30 and HFOs was evaluated before and after application of tDCS. Results After cathodal tDCS to the somatosensory cortex we found a significant reduction of the N20 source amplitude while there was no effect after anodal stimulation. For the N30 component and HFOs no change in source activity was observed. Conclusions Corresponding to the results for the motor cortex a sustained reduction of the excitability of the somatosensory cortex after cathodal tDCS was shown. Significance We demonstrated differential effects of tDCS on the high and low frequency components of SEPs confirming the hypothesis of locally and functionally distinct generators of these two components.

Intracortically Distributed Neurovascular Coupling Relationships within and between Human Somatosensory Cortices

The coupling of neuronal cellular activity to its blood supply is of critical importance to the physiology of the human brain and has been under discussion for more than a century. Linearity in this relationship has been demonstrated in some animal studies, but evidence is lacking in humans. In this study, we compared scalp evoked potentials and the functional magnetic resonance imaging (fMRI) blood oxygen level-dependent (BOLD) signal from healthy human volunteers with changes in the intensity of a somatosensory stimulus. By weighting the fMRI images according to the evoked potential amplitude at corresponding intensities, we tested for positive and negative covariation between these 2 data sets and the extent to which these were linear. Hemodynamic changes in primary somatosensory cortex covaried positively with neuronal activity in a predominantly linear manner, with a small quadratic contribution. Simultaneously, other cortical areas corresponding to the nonstimulated limbs were found to covary negatively and linearly in the hemispheres ipsilateral and contralateral to the stimulus. These concurrent and bilateral cortical dynamics, as well as the intraregional features of this neurovascular coupling, are both more complex than had been considered to date, with considerable implications.

Differential effects of high and low-frequency TMS on different tactile perceptual tasks and cortical excitability of human somatosensory cortex

Differential effects of high and low-frequency TMS on different tactile perceptual tasks and cortical excitability of human somatosensory cortex

Bodily Illusions Modulate Tactile Perception

Touch differs from other exteroceptive senses in that the body itself forms part of the tactile percept. Interactions between proprioception and touch provide a powerful way to investigate the implicit body representation underlying touch. Here, we demonstrate that an intrinsic primary quality of a tactile object, for example its size, is directly affected by the perceived size of the body part touching it. We elicited proprioceptive illusions that the left index finger was either elongating or shrinking by vibrating the biceps or triceps tendon of the right arm while subjects grasped the tip of their left index finger. Subjects estimated the distance between two simultaneous tactile contacts on the left finger during tendon vibration. We found that tactile distances feel bigger when the touched body part feels elongated. Control tests showed that the modulation of touch was linked to the perceived index-finger size induced by tendon vibration. Vibrations that did not produce proprioceptive illusion had no effect on touch. Our results show that the perception of tactile objects is referenced to an implicit body representation and that proprioception contributes to this body representation. We also provide, for the first time, a quantitative, implicit measure of distortions of body size.

Timing‐dependent plasticity in human primary somatosensory cortex

Animal experiments suggest that cortical sensory representations may be remodelled as a consequence of changing synaptic efficacy by timing-dependent associative neuronal activity. Here we describe a timing-based associative form of plasticity in human somatosensory cortex. Paired associative stimulation (PAS) was performed by combining repetitive median nerve stimulation with transcranial magnetic stimulation (TMS) over the contralateral postcentral region. PAS increased exclusively the amplitude of the P25 component of the median nerve-evoked somatosensory-evoked potential (MN-SSEP), which is probably generated in the superficial cortical layers of area 3b. SSEP components reflecting neuronal activity in deeper cortical layers (N20 component) or subcortical regions (P14 component) remained constant. PAS-induced enhancement of P25 amplitude displayed topographical specificity both for the recording (MN-SSEP versus tibial nerve-SSEP) and the stimulation (magnetic stimulation targeting somatosensory versus motor cortex) arrangements. Modulation of P25 amplitude was confined to a narrow range of interstimulus intervals (ISIs) between the MN pulse and the TMS pulse, and the sign of the modulation changed with ISIs differing by only 15 ms. The function describing the ISI dependence of PAS effects on somatosensory cortex resembled one previously observed in motor cortex, shifted by approximately 7 ms. The findings suggest a simple model of modulation of excitability in human primary somatosensory cortex, possibly by mechanisms related to the spike-timing-dependent plasticity of neuronal synapses located in upper cortical layers.

Brain Response to Visceral Aversive Conditioning: A Functional Magnetic Resonance Imaging Study

Brain-imaging studies to date have confounded visceral pain perception with anticipation. We used functional magnetic resonance imaging of the human brain to study the neuroanatomic network involved in aversive conditioning of visceral pain and, thus, anticipation. Eight healthy volunteers (5 male) participated in the study. We used a classic conditioning paradigm in which 3 neutral stimuli (differently colored circles) that acted as conditioned stimuli were paired with painful esophageal distention, air puff to the wrist, or nothing, which acted as unconditioned stimuli. Neural activity was measured during learning, anticipation (pairing only 50% of conditioned stimuli with their unconditioned stimuli), and extinction (unpaired conditioned stimuli) phases. For magnetic resonance imaging, axial slices depicting blood oxygen level-dependent contrast were acquired with a 1.5-T system. Neural responses during the learning phase included areas commonly associated with visceral pain (anterior cingulate cortex, insula, and primary and secondary somatosensory cortices) and innocuous somatosensory perception (primary and secondary somatosensory cortices and insula). During the anticipation and extinction phases of aversive stimulation, brain activity resembled that seen during actual painful esophageal stimulation. In contrast, anticipation and extinction of the innocuous somatic stimulus failed to show that effect. We have shown that actual and anticipated visceral pain elicit similar cortical responses. These results have implications for the design and interpretation of brain-imaging studies of visceral pain. They not only contribute to our understanding of the processing of visceral pain, but also have clinical implications for the management of chronic pain states.

Functional representation of the finger and face in the human somatosensory cortex: intraoperative intrinsic optical imaging

We applied the intrinsic optical imaging technique to the human primary somatosensory cortex during brain tumor/epilepsy surgery for nine patients. The cortical surface was illuminated with a Xenon light through an operating microscope, and the reflected light, which passed through a 605 nm bandpass filter, was detected by a CCD camera-based optical imaging system. Individual electrical stimulation of five digits induced changes in the reflected light intensities. Visualizing the intrinsic optical responses, we constructed maps of finger representation in Brodmann's area 1. In the maps, response areas of Digits I to V were sequentially aligned along the central sulcus in the crown of the postcentral gyrus from the latero-inferior region (Digit I) to the medio-superior region (Digit V). The neighboring response areas partially overlapped each other, as previously described in the monkey somatosensory cortex. Similar results were obtained in the face region with stimulation of the three branches of the trigeminal nerve. These results suggest that the overlap of the response areas is a common feature in the somatosensory cortex not only in monkeys, but also in humans.