Imperceptible somatosensory stimulation alters sensorimotor background rhythm and connectivity (original) (raw)

Early and late activity in somatosensory cortex reflects changes in bodily self-consciousness: An evoked potential study

Neuroscience, 2012

How can we investigate the brain mechanisms underlying self-consciousness? Recent behavioural studies on multisensory bodily perception have shown that multisensory conflicts can alter bodily self-consciousness such as in the ''full body illusion'' (FBI) in which changes in self-identification with a virtual body and tactile perception are induced. Here we investigated whether experimental changes in self-identification during the FBI are accompanied by activity changes in somatosensory cortex by recording somatosensory-evoked potentials (SEPs). To modulate self-identification, participants were filmed by a video camera from behind while their backs were stroked, either synchronously (illusion condition) or asynchronously (control condition) with respect to the stroking seen on their virtual body. Tibial nerve SEPs were recorded during the FBI and analysed using evoked potential (EP) mapping. Tactile mislocalisation was measured using the crossmodal congruency task. SEP mapping revealed five sequential periods of brain activation during the FBI, of which two differed between the illusion condition and the control condition. Activation at 30-50 ms (corresponding to the P40 component) in primary somatosensory cortex was stronger in the illusion condition. A later activation at $110-200 ms, likely originating in higher-tier somatosensory regions in parietal cortex, was stronger and lasted longer in the control condition. These data show that changes in bodily self-consciousness modulate activity in primary and higher-tier somatosensory cortex at two distinct processing steps. We argue that early modulations of primary somatosensory cortex may be a consequence of (1) multisensory integration of synchronous vs. asynchronous visuo-tactile stimuli and/or (2) differences in spatial attention (to near or far space) between the conditions. The later activation in higher-tier parietal cortex (and potentially other regions in temporo-parietal and frontal cortex) likely reflects the detection of visuo-tactile conflicts in the asynchronous condition.

Early Neural Correlates of Conscious Somatosensory Perception

Journal of Neuroscience, 2005

The cortical processing of consciously perceived and unperceived somatosensory stimuli is thought to be identical during the first 100 -120 ms after stimulus onset. Thereafter, the electrophysiological correlates of conscious perception have been shown to be reflected in the N1 component of the evoked response as well as in later (Ͼ200 ms) nonstimulus-locked ␥-band (28 -50 Hz) oscillatory activity. To evaluate more specifically the time course and correlation of neuronal oscillations with conscious perception, we recorded neuromagnetic responses to threshold-intensity somatosensory stimuli. We show here that cortical broadband activities phase locked to the subsequently perceived stimuli in somatosensory, frontal, and parietal regions as early as 30 -70 ms from stimulus onset, whereas the phase locking to the unperceived stimuli was weak and primarily restricted to somatosensory regions. Such stimulus locking also preceded the perceived stimuli, indicating that the phase of ongoing cortical activities biases subsequent perception. Furthermore, the data show that the stimulus locking was present in the -(4 -8 Hz), ␣-(8 -14 Hz), ␤-(14 -28 Hz), and ␥-(28 -40 Hz) frequency bands, of which the widespread ␣-band component was dominant for the consciously perceived stimuli but virtually unobservable for the unperceived stimuli. Our results show that the neural correlates of conscious perception are already found during the earliest stages of cortical processing from 30 to 150 ms after stimulus onset and suggest that ␣-frequency-band oscillations have a role in the neural mechanisms of sensory awareness.

The Tactile Window to Consciousness is Characterized by Frequency-Specific Integration and Segregation of the Primary Somatosensory Cortex

Scientific Reports, 2016

We recently proposed that besides levels of local cortical excitability, also distinct pre-stimulus network states (windows to consciousness) determine whether a near-threshold stimulus will be consciously perceived. In the present magnetoencephalography study, we scrutinised these pre-stimulus network states with a focus on the primary somatosensory cortex. For this purpose participants performed a simple near-threshold tactile detection task. Confirming previous studies, we found reduced alpha and beta power in the somatosensory region contralateral to stimulation prior to correct stimulus detection as compared to undetected stimuli, and stronger event-related responses following successful stimulus detection. As expected, using graph theoretical measures, we also observed modulated pre-stimulus network level integration. Specifically, the right primary somatosensory cortex contralateral to stimulation showed an increased integration in the theta band, and additionally, a decreased integration in the beta band. Overall, these results underline the importance of network states for enabling conscious perception. Moreover, they indicate that also a reduction of irrelevant functional connections contributes to the window to consciousness by tuning pre-stimulus pathways of information flow. Studies investigating pre-stimulus effects in near-threshold (NT) paradigms observed that correctly perceived stimuli are preceded by low alpha power in task-relevant areas. This was shown for the visual 1-4 and the somatosensory cortex 5-9 depending on the task. These observations are usually interpreted according to the notion that alpha activity reflects the cortical excitability with strong alpha reflecting functional inhibition 10,11. The straightforward rationale, thus, states that an upcoming NT stimulus will become conscious when pre-stimulus local excitability (e.g., in the visual cortex) is high such that a weak input causes ignition of relevant neural assemblies. Hence, alpha power effects are interpreted in local terms. Despite providing an intuitive explanation of pre-stimulus determinants of conscious perception that are well linked to a strong conceptual framework, this interpretation has a major shortcoming. As this viewpoint emphasizes local pre-stimulus cortical excitability, it predicts that successful stimulus detection depends on a bottom-up input sweep. In this case, effects in sensory regions should become evident immediately, as soon as an ignition threshold is crossed. Interestingly, evidence for this implicit prediction is rather scarce. In contrast, effects in sensory regions are reported to appear relatively late, probably due to recurrent activation from downstream areas 12,13. Apart from this empirical discrepancy of what should be expected if pre-stimulus effects were interpreted along the functional inhibition hypothesis, major neuroscientific frameworks of conscious perception stress a network perspective. For example the global neuronal workspace model (GNW) 14 suggests that sensory stimuli

Baseline brain activity fluctuations predict somatosensory perception in humans

Proceedings of the National Academy of Sciences, 2007

In perceptual experiments, within-individual fluctuations in perception are observed across multiple presentations of the same stimuli, a phenomenon that remains only partially understood. Here, by means of thulium-yttrium/aluminum-garnet laser and event-related functional MRI, we tested whether variability in perception of identical stimuli relates to differences in prestimulus, baseline brain activity. Results indicate a positive relationship between conscious perception of low-intensity somatosensory stimuli and immediately preceding levels of baseline activity in medial thalamus and the lateral frontoparietal network, respectively, which are thought to relate to vigilance and ''external monitoring.'' Conversely, there was a negative correlation between subsequent reporting of conscious perception and baseline activity in a set of regions encompassing posterior cingulate/ precuneus and temporoparietal cortices, possibly relating to introspection and self-oriented processes. At nociceptive levels of stimulation, pain-intensity ratings positively correlated with baseline fluctuations in anterior cingulate cortex in an area known to be involved in the affective dimension of pain. These results suggest that baseline brain-activity fluctuations may profoundly modify our conscious perception of the external world.

Imperceptible Somatosensory Single Pulse and Pulse Train Stimulation Oppositely Modulate Mu Rhythm Activity and Perceptual Performance

Cerebral Cortex

Subliminal stimulation alters conscious perception – a potential mechanism is the modulation of cortical background rhythms especially in the alpha range. Here, in the human somatosensory domain, we assessed effects of subthreshold (imperceptible) electrical finger nerve stimulation – either presented as single pulses or as brief (1 s) 7 Hz pulse trains—on mu-alpha rhythm and perceptual performance. In electroencephalography, subthreshold single pulses transiently (~150–350 ms poststimulus) increased mu activity (event-related synchronization), while, interestingly, subthreshold trains led to prolonged (>1 s) mu desynchronization. In psychophysics, detection of near-threshold target stimuli was consistently reduced when presented together with subthreshold trains (at three delays), whereas for targets paired with subthreshold single pulses detection remained unaffected (30 and 180 ms) or was even elevated (60 ms). Though both imperceptible, single pulses and pulse trains exerted ...

Synchronization of Neural Activity across Cortical Areas Correlates with Conscious Perception

Journal of Neuroscience, 2007

Subliminal stimuli can be deeply processed and activate similar brain areas as consciously perceived stimuli. This raises the question which signatures of neural activity critically differentiate conscious from unconscious processing. Transient synchronization of neural activity has been proposed as a neural correlate of conscious perception. Here we test this proposal by comparing the electrophysiological responses related to the processing of visible and invisible words in a delayed matching to sample task. Both perceived and nonperceived words caused a similar increase of local (gamma) oscillations in the EEG, but only perceived words induced a transient long-distance synchronization of gamma oscillations across widely separated regions of the brain. After this transient period of temporal coordination, the electrographic signatures of conscious and unconscious processes continue to diverge. Only words reported as perceived induced (1) enhanced theta oscillations over frontal regions during the maintenance interval, (2) an increase of the P300 component of the eventrelated potential, and (3) an increase in power and phase synchrony of gamma oscillations before the anticipated presentation of the test word. We propose that the critical process mediating the access to conscious perception is the early transient global increase of phase synchrony of oscillatory activity in the gamma frequency range.

Direct electrical stimulation of human cortex evokes high gamma activity that predicts conscious somatosensory perception

Journal of neural engineering, 2017

Direct electrical stimulation (DES) is a clinical gold standard for human brain mapping and readily evokes conscious percepts, yet the neurophysiological changes underlying these percepts are not well understood. To determine the neural correlates of DES, we stimulated the somatosensory cortex of ten human participants at frequency-amplitude combinations that both elicited and failed to elicit conscious percepts, meanwhile recording neural activity directly surrounding the stimulation site. We then compared the neural activity of perceived trials to that of non-perceived trials. We found that stimulation evokes distributed high gamma activity, which correlates with conscious perception better than stimulation parameters themselves. Our findings suggest that high gamma activity is a reliable biomarker for perception evoked by both natural and electrical stimuli.

Functional deactivations: Multiple ipsilateral brain areas engaged in the processing of somatosensory information

Human Brain Mapping, 2011

Somatosensory signals modulate activity throughout a widespread network in both of the brain hemispheres: the contralateral as well as the ipsilateral side of the brain relative to the stimulated limb. To analyze the ipsilateral somatosensory brain areas that are engaged during limb stimulation, we performed functional magnetic resonance imaging (fMRI) in 12 healthy subjects during electrical median nerve stimulation using both a block-and an event-related fMRI design. Data were analyzed through the use of model-dependent (SPM) and model-independent (ICA) approaches. Beyond the well-known positive blood oxygenation level-dependent (BOLD) responses, negative deflections of the BOLD response were found consistently in several ipsilateral brain areas, including the primary somatosensory cortex, the supplementary motor area, the insula, the dorsal part of the posterior cingulate cortex, and the contralateral cerebellum. Compared to their positive counterparts, the negative hemodynamic responses showed a different time course, with an onset time delay of 2.4 s and a peak delay of 0.7 s. This characteristic delay was observed in all investigated areas and verified by a second (purely tactile) event-related paradigm, suggesting a systematic difference for brain areas involved in the processing of somatosensory information. These findings may indicate that the physiological basis of these deactivations differs from that of the positive BOLD responses. Therefore, an altered model for the negative BOLD response may be beneficial to further model-dependent fMRI analyses. Hum Brain Mapp 32:127-140, in Wiley Online Library (wileyonlinelibrary.com). V C 2010 Wiley-Liss, Inc. r Brain Engaged in the Processing of Somatosensory Information r r 129 r r Brain Engaged in the Processing of Somatosensory Information r r 139 r

Re-wiring the brain: Increased functional connectivity within primary somatosensory cortex following synchronous co-activation

NeuroImage, 2014

The primary somatosensory cortex shows precise topographical organisation, but can be quickly modified by alterations to sensory inputs. Temporally correlated sensory inputs to the digits can result in the merging of digit representations on the cortical surface. Underlying mechanisms driving these changes are unclear but the strengthening of intra-cortical synaptic connections via Hebbian mechanisms has been suggested. We use fMRI measures of temporal coherence to infer alterations in the relative strength of neuronal connections between digit regions 2 and 4 following 3 hours of synchronous and asynchronous co-activation. Following synchronous co-activation we find a 20% increase in temporal coherence of the fMRI signal (p = 0.0004). No significant change is seen following asynchronous co-activation suggesting that temporal coincidence between the two digit inputs during co-activation is driving this coherence change. In line with previous work we also find a trend towards reduced separation of the digit representations following synchronous co-activation and significantly increased separation for the asynchronous case. Increased coherence is significantly correlated with reduced digit separation for the synchronous case. This study shows that passive synchronous stimulation to the digits strengthens the underlying cortical connections between the digit regions in only a few hours, and that this mechanism may be related to topographical re-organisation.