Respiration, Heartbeat, and Conscious Tactile Perception.

Previous studies have shown that timing of sensory stimulation during the cardiac cycle interacts with perception. Given the natural coupling of respiration and cardiac activity, we investigated here their joint effects on tactile perception. Forty-one healthy female and male human participants reported conscious perception of finger near-threshold electrical pulses (33% null trials) and decision confidence while electrocardiography, respiratory activity, and finger photoplethysmography were recorded. Participants adapted their respiratory cycle to expected stimulus onsets to preferentially occur during late inspiration/early expiration. This closely matched heart rate variation (sinus arrhythmia) across the respiratory cycle such that most frequent stimulation onsets occurred during the period of highest heart rate probably indicating highest alertness and cortical excitability. Tactile detection rate was highest during the first quadrant after expiration onset. Interindividually, stronger respiratory phase-locking to the task was associated with higher detection rates. Regarding the cardiac cycle, we confirmed previous findings that tactile detection rate was higher during diastole than systole and newly specified its minimum at 250-300 ms after the R-peak corresponding to the pulse wave arrival in the finger. Expectation of stimulation induced a transient heart deceleration which was more pronounced for unconfident decision ratings. Interindividually, stronger poststimulus modulations of heart rate were linked to higher detection rates. In summary, we demonstrate how tuning to the respiratory cycle and integration of respiratory-cardiac signals are used to optimize performance of a tactile detection task.SIGNIFICANCE STATEMENT Mechanistic studies on perception and cognition tend to focus on the brain neglecting contributions of the body. Here, we investigated how respiration and heartbeat influence tactile perception: respiration phase-locking to expected stimulus onsets corresponds to highest heart rate (and presumably alertness/cortical excitability) and correlates with detection performance. Tactile detection varies across the heart cycle with a minimum when the pulse reaches the finger and a maximum in diastole. Taken together with our previous finding of unchanged early event-related potentials across the cardiac cycle, we conclude that these effects are not a peripheral physiological artifact but a result of cognitive processes that model our body's internal state, make predictions to guide behavior, and might also tune respiration to serve the task.

Heart-brain interactions shape somatosensory perception and evoked potentials.

Even though humans are mostly not aware of their heartbeats, several heartbeat-related effects have been reported to influence conscious perception. It is not clear whether these effects are distinct or related phenomena, or whether they are early sensory effects or late decisional processes. Combining electroencephalography and electrocardiography, along with signal detection theory analyses, we identify two distinct heartbeat-related influences on conscious perception differentially related to early vs. late somatosensory processing. First, an effect on early sensory processing was found for the heartbeat-evoked potential (HEP), a marker of cardiac interoception. The amplitude of the prestimulus HEP negatively correlated with localization and detection of somatosensory stimuli, reflecting a more conservative detection bias (criterion). Importantly, higher HEP amplitudes were followed by decreases in early (P50) as well as late (N140, P300) somatosensory-evoked potential (SEP) amplitudes. Second, stimulus timing along the cardiac cycle also affected perception. During systole, stimuli were detected and correctly localized less frequently, relating to a shift in perceptual sensitivity. This perceptual attenuation was accompanied by the suppression of only late SEP components (P300) and was stronger for individuals with a more stable heart rate. Both heart-related effects were independent of alpha oscillations' influence on somatosensory processing. We explain cardiac cycle timing effects in a predictive coding account and suggest that HEP-related effects might reflect spontaneous shifts between interoception and exteroception or modulations of general attentional resources. Thus, our results provide a general conceptual framework to explain how internal signals can be integrated into our conscious perception of the world.

Dissociating Perceptual Awareness and Postperceptual Processing: The P300 Is Not a Reliable Marker of Somatosensory Target Detection.

A central challenge in the study of conscious perception lies in dissociating the neural correlates of perceptual awareness from those reflecting its precursors and consequences. No-report paradigms have been instrumental in this endeavor, demonstrating that the event-related potential P300, recorded from the human scalp, reflects reports rather than awareness. However, these paradigms cannot probe the degree to which stimuli are consciously processed from trial to trial and, thus, leave open the possibility that the P300 is a genuine correlate of conscious access enabling reports. Here, instead of removing report requirements, we took the opposite approach and equated postperceptual task demands across conscious and unconscious trials by orthogonalizing target detection and overt reports in a somatosensory detection task. We used Bayesian model selection to track the transformation from physical to perceptual processing stages in the EEG data of 24 male and female participants and show that the early P50 component scaled with physical stimulus intensity, whereas the N140 component was the first correlate of target detection. The late P300 component was elicited for both perceived and unperceived stimuli and was not substantially modulated by target detection. This was in stark contrast to a control experiment using a classical direct report task, which replicated the P50 and N140 effects but additionally showed a strong effect of target detection in the P300 time range. Our results demonstrate the task dependence of the P300 in the somatosensory modality and show that late cortical potentials dissociate from perceptual awareness even when stimuli are always reported.SIGNIFICANCE STATEMENT The time it takes for sensory information to enter our conscious experience can be an indicator of the neural processing stages that lead to perceptual awareness. However, because many cognitive processes routinely correlate with perception, isolating those signals that uniquely reflect perceptual awareness is not a trivial task. Here, we show that late electroencephalography signals cease to correlate with somatosensory awareness when common task confounds are controlled. Importantly, by balancing report requirements instead of abolishing them, we show that the lack of late effects cannot be explained by a lack of conscious access. Instead, we propose that conscious access occurs earlier, at ∼150 ms, supporting the view that early activity in sensory cortices is a neural correlate of conscious perception.

Immediate brain plasticity after one hour of brain-computer interface (BCI).

KEY POINTS: Two groups of inexperienced brain-computer interface users underwent a purely mental EEG-BCI session that rapidly impacted on their brain. Modulations in structural and functional MRI were found after only 1 h of BCI training. Two different types of BCI (based on motor imagery or visually evoked potentials) were employed and analyses showed that the brain plastic changes are spatially specific for the respective neurofeedback. This spatial specificity promises tailored therapeutic interventions (e.g. for stroke patients). ABSTRACT: A brain-computer-interface (BCI) allows humans to control computational devices using only neural signals. However, it is still an open question, whether performing BCI also impacts on the brain itself, i.e. whether brain plasticity is induced. Here, we show rapid and spatially specific signs of brain plasticity measured with functional and structural MRI after only 1 h of purely mental BCI training in BCI-naive subjects. We employed two BCI approaches with neurofeedback based on (i) modulations of EEG rhythms by motor imagery (MI-BCI) or (ii) event-related potentials elicited by visually targeting flashing letters (ERP-BCI). Before and after the BCI session we performed structural and functional MRI. For both BCI approaches we found increased T1-weighted MR signal in the grey matter of the respective target brain regions, such as occipital/parietal areas after ERP-BCI and precuneus and sensorimotor regions after MI-BCI. The latter also showed increased functional connectivity and higher task-evoked BOLD activity in the same areas. Our results demonstrate for the first time that BCI by means of targeted neurofeedback rapidly impacts on MRI measures of brain structure and function. The spatial specificity of BCI-induced brain plasticity promises therapeutic interventions tailored to individual functional deficits, for example in patients after stroke.

Neural correlates of conscious tactile perception: An analysis of BOLD activation patterns and graph metrics.

Theories of human consciousness substantially vary in the proposed spatial extent of brain activity associated with conscious perception as well as in the assumed functional alterations within the involved brain regions. Here, we investigate which local and global changes in brain activity accompany conscious somatosensory perception following electrical finger nerve stimulation, and whether there are whole-brain functional network alterations by means of graph metrics. Thirty-eight healthy participants performed a somatosensory detection task and reported their decision confidence during fMRI. For conscious tactile perception in contrast to undetected near-threshold trials (misses), we observed increased BOLD activity in the precuneus, the intraparietal sulcus, the insula, the nucleus accumbens, the inferior frontal gyrus and the contralateral secondary somatosensory cortex. For misses compared to correct rejections, bilateral secondary somatosensory cortices, supplementary motor cortex and insula showed greater activations. The analysis of whole-brain functional network topology for hits, misses and correct rejections, did not result in any significant differences in modularity, participation, clustering or path length, which was supported by Bayes factor statistics. In conclusion, for conscious somatosensory perception, our results are consistent with an involvement of (probably) domain-general brain areas (precuneus, insula, inferior frontal gyrus) in addition to somatosensory regions; our data do not support the notion of specific changes in graph metrics associated with conscious experience. For the employed somatosensory submodality of fine electrical current stimulation, this speaks for a global broadcasting of sensory content across the brain without substantial reconfiguration of the whole-brain functional network resulting in an integrative conscious experience.

Inter-individual and age-dependent variability in simulated electric fields induced by conventional transcranial electrical stimulation.

Variations in head and brain anatomy determine the strength and distribution of electrical fields in humans and may account for inconsistent behavioral and neurophysiological results in transcranial electrical stimulation (tES) studies. However, it is insufficiently understood which anatomical features contribute to the variability of the modelled electric fields, and if their impact varies across age groups. In the present study, we tested the associations of global head anatomy, indexed by extra- and intra-cranial volumes, with electric field measures, comparing young and older adults. We modelled six "conventional" electrode montages typically used in tES studies using SimNIBS software in 40 individuals (20 young, 20 older adults; 20-35, 64-79 years). We extracted individual electric field strengths and focality values for each montage to identify tissue volumes that account for variability of the induced electric fields in both groups. Linear mixed models explained most of the inter-individual variability of the overall induced field strength in the brain, but not of field focality. Higher absolute head volume and relative volume of skin, skull and cerebrospinal fluid (CSF) were associated with lower overall electric field strengths. Additionally, we found interactions of age group with head volume and CSF, indicating that this relationship was mitigated in the older group. Our results demonstrate the importance to adjust brain stimulation not only according to brain atrophy, but also to additional parameters of head anatomy. Future studies need to elucidate the mechanisms underlying individual variability of tES effects in young and older adults, and verify the usefulness of the proposed models in terms of neurophysiology and behavior in empirical studies.

A novel approach for assessing hypoperfusion in stroke using spatial independent component analysis of resting-state fMRI.

Individualized treatment of acute stroke depends on the timely detection of ischemia and potentially salvageable tissue in the brain. Using functional MRI (fMRI), it is possible to characterize cerebral blood flow from blood-oxygen-level-dependent (BOLD) signals without the administration of exogenous contrast agents. In this study, we applied spatial independent component analysis to resting-state fMRI data of 37 stroke patients scanned within 24 hr of symptom onset, 17 of whom received follow-up scans the next day. Our analysis revealed "Hypoperfusion spatially-Independent Components" (HICs) whose spatial patterns of BOLD signal resembled regions of delayed perfusion depicted by dynamic susceptibility contrast MRI. These HICs were detected even in the presence of excessive patient motion, and disappeared following successful tissue reperfusion. The unique spatial and temporal features of HICs allowed them to be distinguished with high accuracy from other components in a user-independent manner (area under the curve = 0.93, balanced accuracy = 0.90, sensitivity = 1.00, and specificity = 0.85). Our study therefore presents a new, noninvasive method for assessing blood flow in acute stroke that minimizes interpretative subjectivity and is robust to severe patient motion.

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

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 opposite effects on neural signaling and perception. We suggest that the common neural basis is preferential activation of cortical inhibitory interneurons. While the inhibitory impact of a subthreshold single pulse (reflected by mu synchronization) is not psychophysically detectable-rather perception may be facilitated-repetition of the same subthreshold pulse shifts the excitation-inhibition balance toward an inhibitory cortical state (reflected by perceptual impediment) accompanied by mu desynchronization. These differential findings provide novel insights on the notion of alpha activity mediating functional inhibition.

Dissociable neural correlates of stimulation intensity and detection in somatosensation.

Somatosensory stimulation intensity and behavioral detection are positively related, and both correlate with neural responses. However, it is still controversial as to what extent stimulus intensity and early somatosensory evoked potentials (SEP) predict detection and how these parameters interact with pre-stimulus brain oscillatory states, which also influence sensory processing. Here we investigated how early SEP components encode stimulation intensity, how pre-stimulus alpha- and beta-band amplitudes interact with SEPs, and which neural markers predict stimulus detection. To this end, we randomly presented electrical finger nerve stimulation with various intensities distributed along the individual psychometric response function (including catch trials) while recording the EEG. Participants reported stimulus presence on a trial-by-trial basis (one-alternative-forced-choice). For the lowest (imperceptible) intensities, participants showed zero (behavioral) sensitivity despite measurable early cortical processing reflected by the P50 component. The P50 amplitude scaled with increasing stimulation intensities but was not predictive of stimulus detection. Instead, detection was associated with the later negative N150 component, as well as with pre-stimulus lowered somatosensory alpha- and increased frontal beta-band amplitudes. Our results give evidence for a serial representation of stimulus intensity and detection, as reflected by the P50 and N150 amplitude, respectively. Furthermore, stimulus detection seems to depend on the current brain state, rendering upcoming stimulation being reportable or not.

Reduction of somatosensory functional connectivity by transcranial alternating current stimulation at endogenous mu-frequency.

Alpha, the most prominent human brain rhythm, might reflect a mechanism of functional inhibition for gating neural processing. This concept has been derived predominantly from local measures of inhibition, while large-scale network mechanisms to guide information flow are largely unknown. Here, we investigated functional connectivity changes on a whole-brain level by concurrent transcranial alternating current stimulation (tACS) and resting-state functional MRI in humans. We specifically focused on somatosensory alpha-band oscillations by adjusting the tACS frequency to each individual´s somatosensory (mu-) alpha peak frequency (mu-tACS). Potential differences of Eigenvector Centrality of primary somatosensory cortex (S1) as well as on a whole brain level between mu-tACS and sham were analyzed. Our results demonstrate that mu-tACS induces a locally-specific decrease in whole-brain functional connectivity of left S1. An additional exploratory analysis revealed that this effect primarily depends on a decrease in functional connectivity between S1 and a network of regions that are crucially involved in somatosensory processing. Furthermore, the decrease in functional centrality was specific to mu-tACS and was not observed when tACS was applied in the gamma-range in an independent study. Our findings provide evidence that modulated somatosensory (mu-) alpha-activity may affect whole-brain network level activity by decoupling primary sensory areas from other hubs involved in sensory processing.

Alpha-Band Brain Oscillations Shape the Processing of Perceptible as well as Imperceptible Somatosensory Stimuli during Selective Attention.

Attention filters and weights sensory information according to behavioral demands. Stimulus-related neural responses are increased for the attended stimulus. Does alpha-band activity mediate this effect and is it restricted to conscious sensory events (suprathreshold), or does it also extend to unconscious stimuli (subthreshold)? To address these questions, we recorded EEG in healthy male and female volunteers undergoing subthreshold and suprathreshold somatosensory electrical stimulation to the left or right index finger. The task was to detect stimulation at the randomly alternated cued index finger. Under attention, amplitudes of somatosensory evoked potentials increased 50-60 ms after stimulation (P1) for both suprathreshold and subthreshold events. Prestimulus amplitude of peri-Rolandic alpha, that is mu, showed an inverse relationship to P1 amplitude during attention compared to when the finger was unattended. Interestingly, intermediate and high amplitudes of mu rhythm were associated with the highest P1 amplitudes during attention and smallest P1 during lack of attention, that is, these levels of alpha rhythm seemed to optimally support the behavioral goal ("detect" stimuli at the cued finger while ignoring the other finger). Our results show that attention enhances neural processing for both suprathreshold and subthreshold stimuli and they highlight a rather complex interaction between attention, Rolandic alpha activity, and their effects on stimulus processing.SIGNIFICANCE STATEMENT Attention is crucial in prioritizing processing of relevant perceptible (suprathreshold) stimuli: it filters and weights sensory input. The present study investigates the controversially discussed question whether this attention effect extends to imperceptible (subthreshold) stimuli as well. We found noninvasive EEG signatures for attentional modulation of neural events following perceptible and imperceptible somatosensory stimulation in human participants. Specifically, stimulus processing for both kinds of stimulation, subthreshold and suprathreshold, is enhanced by attention. Interestingly, Rolandic alpha rhythm strength and its influence on stimulus processing are strikingly altered by attention most likely to optimally achieve the behavioral goal.

Age-dependent effects of brain stimulation on network centrality.

Functional magnetic resonance imaging (fMRI) studies have suggested that advanced age may mediate the effects of transcranial direct current stimulation (tDCS) on brain function. However, studies directly comparing neural tDCS effects between young and older adults are scarce and limited to task-related imaging paradigms. Resting-state (rs-) fMRI, that is independent of age-related differences in performance, is well suited to investigate age-associated differential neural tDCS effects. Three "online" tDCS conditions (anodal, cathodal, sham) were compared in a cross-over, within-subject design, in 30 young and 30 older adults. Active stimulation targeted the left sensorimotor network (active electrode over left sensorimotor cortex with right supraorbital reference electrode). A graph-based rs-fMRI data analysis approach (eigenvector centrality mapping) and complementary seed-based analyses characterized neural tDCS effects. An interaction between anodal tDCS and age group was observed. Specifically, centrality in bilateral paracentral and posterior regions (precuneus, superior parietal cortex) was increased in young, but decreased in older adults. Seed-based analyses revealed that these opposing patterns of tDCS-induced centrality modulation originated from differential effects of tDCS on functional coupling of the stimulated left paracentral lobule. Cathodal tDCS did not show significant effects. Our study provides first evidence for differential tDCS effects on neural network organization in young and older adults. Anodal stimulation mainly affected coupling of sensorimotor with ventromedial prefrontal areas in young and decoupling with posteromedial areas in older adults.

Relationship Between Changes in the Temporal Dynamics of the Blood-Oxygen-Level-Dependent Signal and Hypoperfusion in Acute Ischemic Stroke.

BACKGROUND AND PURPOSE: Changes in the blood-oxygen-level-dependent (BOLD) signal provide a noninvasive measure of blood flow, but a detailed comparison with established perfusion parameters in acute stroke is lacking. We investigated the relationship between BOLD signal temporal delay and dynamic susceptibility contrast magnetic resonance imaging (DSC-MRI) in stroke patients. METHODS: In 30 patients with acute (<24 hours) ischemic stroke, we performed Pearson correlation and multiple linear regression between DSC-MRI parameters (time to maximum [Tmax], mean transit time, cerebral blood flow, and cerebral blood volume) and BOLD-based parameters (BOLD delay and coefficient of BOLD variation). Prediction of severe hypoperfusion (Tmax >6 seconds) was assessed using receiver-operator characteristic (ROC) analysis. RESULTS: Correlation was highest between Tmax and BOLD delay (venous sinus reference; time shift range 7; median r=0.60; interquartile range=0.49-0.71). Coefficient of BOLD variation correlated with cerebral blood volume (median r= 0.37; interquartile range=0.24-0.51). Mean R2 for predicting BOLD delay by DSC-MRI was 0.54 (SD=0.2) and for predicting coefficient of BOLD variation was 0.37 (SD=0.17). BOLD delay (whole-brain reference, time shift range 3) had an area under the curve of 0.76 for predicting severe hypoperfusion (sensitivity=69.2%; specificity=80%), whereas BOLD delay (venous sinus reference, time shift range 3) had an area under the curve of 0.76 (sensitivity=67.3%; specificity=83.5%). CONCLUSIONS: BOLD delay is related to macrovascular delay and microvascular hypoperfusion, can identify severely hypoperfused tissue in acute stroke, and is a promising alternative to gadolinium contrast agent-based perfusion assessment in acute stroke. CLINICAL TRIAL REGISTRATION: URL: http://www.clinicaltrials.gov. Unique identifier: NCT00715533 and NCT02077582.

Brain Network Reconfiguration and Perceptual Decoupling During an Absorptive State of Consciousness.

Trance is an absorptive state of consciousness characterized by narrowed awareness of external surroundings and has long been used-for example, by shamans-to gain insight. Shamans across cultures often induce trance by listening to rhythmic drumming. Using functional magnetic resonance imaging (fMRI), we examined the brain-network configuration associated with trance. Experienced shamanic practitioners (n = 15) listened to rhythmic drumming, and either entered a trance state or remained in a nontrance state during 8-min scans. We analyzed changes in network connectivity. Trance was associated with higher eigenvector centrality (i.e., stronger hubs) in 3 regions: posterior cingulate cortex (PCC), dorsal anterior cingulate cortex (dACC), and left insula/operculum. Seed-based analysis revealed increased coactivation of the PCC (a default network hub involved in internally oriented cognitive states) with the dACC and insula (control-network regions involved in maintaining relevant neural streams). This coactivation suggests that an internally oriented neural stream was amplified by the modulatory control network. Additionally, during trance, seeds within the auditory pathway were less connected, possibly indicating perceptual decoupling and suppression of the repetitive auditory stimuli. In sum, trance involved coactive default and control networks, and decoupled sensory processing. This network reconfiguration may promote an extended internal train of thought wherein integration and insight can occur.

Connections between intraparietal sulcus and a sensorimotor network underpin sustained tactile attention.

Previous studies on sustained tactile attention draw conclusions about underlying cortical networks by averaging over experimental conditions without considering attentional variance in single trials. This may have formed an imprecise picture of brain processes underpinning sustained tactile attention. In the present study, we simultaneously recorded EEG-fMRI and used modulations of steady-state somatosensory evoked potentials (SSSEPs) as a measure of attentional trial-by-trial variability. Therefore, frequency-tagged streams of vibrotactile stimulations were simultaneously presented to both index fingers. Human participants were cued to sustain attention to either the left or right finger stimulation and to press a button whenever they perceived a target pulse embedded in the to-be-attended stream. In-line with previous studies, a classical general linear model (GLM) analysis based on cued attention conditions revealed increased activity mainly in somatosensory and cerebellar regions. Yet, parametric modeling of the BOLD response using simultaneously recorded SSSEPs as a marker of attentional trial-by-trial variability quarried the intraparietal sulcus (IPS). The IPS in turn showed enhanced functional connectivity to a modality-unspecific attention network. However, this was only revealed on the basis of cued attention conditions in the classical GLM. By considering attentional variability as captured by SSSEPs, the IPS showed increased connectivity to a sensorimotor network, underpinning attentional selection processes between competing tactile stimuli and action choices (press a button or not). Thus, the current findings highlight the potential value by considering attentional variations in single trials and extend previous knowledge on the role of the IPS in tactile attention.

Imperceptible somatosensory stimulation alters sensorimotor background rhythm and connectivity.

Most sensory input to our body is not consciously perceived. Nevertheless, it may reach the cortex and influence our behavior. In this study, we investigated noninvasive neural signatures of unconscious cortical stimulus processing to understand mechanisms, which (1) prevent low-intensity somatosensory stimuli from getting access to conscious experience and which (2) can explain the associated impediment of conscious perception for additional stimuli. Stimulation of digit 2 in humans far below the detection threshold elicited a cortical evoked potential (P1) at 60 ms, but no further somatosensory evoked potential components. No event-related desynchronization was detected; rather, there was a transient synchronization in the alpha frequency range. Using the same stimulation during fMRI, a reduced centrality of contralateral primary somatosensory cortex (SI) was found, which appeared to be mainly driven by reduced functional connectivity to frontoparietal areas. We conclude that after subthreshold stimulation the (excitatory) feedforward sweep of bottom-up processing terminates in SI preventing access to conscious experience. We speculate that this interruption is due to a predominance of inhibitory processing in SI. The increase in alpha activity and the disconnection of SI from frontoparietal areas are likely correlates of an elevated perception threshold and may thus serve as a gating mechanism for the access to conscious experience.

Electrical noise modulates perception of electrical pulses in humans: sensation enhancement via stochastic resonance.

Although noise is usually considered to be harmful for signal detection and information transmission, stochastic resonance (SR) describes the counterintuitive phenomenon of noise enhancing the detection and transmission of weak input signals. In mammalian sensory systems, SR-related phenomena may arise both in the peripheral and the central nervous system. Here, we investigate behavioral SR effects of subliminal electrical noise stimulation on the perception of somatosensory stimuli in humans. We compare the likelihood to detect near-threshold pulses of different intensities applied on the left index finger during presence vs. absence of subliminal noise on the same or an adjacent finger. We show that (low-pass) noise can enhance signal detection when applied on the same finger. This enhancement is strong for near-threshold pulses below the 50% detection threshold and becomes stronger when near-threshold pulses are applied as brief trains. The effect reverses at pulse intensities above threshold, especially when noise is replaced by subliminal sinusoidal stimulation, arguing for a peripheral direct current addition. Unfiltered noise applied on longer pulses enhances detection of all pulse intensities. Noise applied to an adjacent finger has two opposing effects: an inhibiting effect (presumably due to lateral inhibition) and an enhancing effect (most likely due to SR in the central nervous system). In summary, we demonstrate that subliminal noise can significantly modulate detection performance of near-threshold stimuli. Our results indicate SR effects in the peripheral and central nervous system.

Functional connectivity-based parcellation of the human sensorimotor cortex.

Task-based functional magnetic resonance imaging (fMRI) has been successfully employed to obtain somatotopic maps of the human sensorimotor cortex. Here, we showed through direct comparison that a similar functional map can be obtained, independently of a task, by performing a connectivity-based parcellation of the sensorimotor cortex based on resting-state fMRI. Cortex corresponding to two adjacent Brodmann areas (BA 3 and BA 4) was selected as the sensorimotor area. Parcellation was obtained along a medial-lateral axis, which was confirmed to be somatotopic (corresponding roughly to an upper, middle and lower limb, respectively) by comparing it with maps obtained using motoric task-based fMRI in the same participants. Interestingly, the resting-state parcellation map demonstrated higher correspondence to the task-based divisions after individuals performed the motor task. Using the resting-state fMRI data, we also observed higher functional correlations between the centrally located hand region and the other two regions, than between the foot and tongue. The functional relevance of these somatosensory parcellation results indicates the feasibility of a wide range of potential applications to brain mapping.

Internal ventilation system of MR scanners induces specific EEG artifact during simultaneous EEG-fMRI.

During simultaneous EEG-fMRI acquisition, the EEG signal suffers from tremendous artifacts caused by the scanner "environment". Particularly, gradient artifacts and the ballistocardiogram have been well characterized, along with methods to eliminate them. Here, we describe another systematic artifact in the EEG signal, which is induced by the internal ventilation system of Siemens TRIO and VERIO MR scanners. A ventilation-level dependent vibration induces specific peaks in the frequency spectrum of the EEG. These frequency peaks are in the range of physiologically relevant brain rhythms (gamma frequency range), and thus interfere with their reliable acquisition. This ventilation dependent artifact was most prominent on the electrodes placed directly on the subject's head, so it is not sufficient to simply place the EEG's amplifier outside the scanner tube. Instead, the ventilator must be switched off to fully eliminate the ventilator's artificial manipulation of EEG recordings. Without the internal ventilator system being on, the temperature within the scanner tube may rise, thus requiring shorter scanning sessions or an additional external ventilation system.

Content Representation of Tactile Mental Imagery in Primary Somatosensory Cortex.

The imagination of tactile stimulation has been shown to activate primary somatosensory cortex (S1) with a somatotopic specificity akin to that seen during the perception of tactile stimuli. Using fMRI and multivariate pattern analysis, we investigate whether this recruitment of sensory regions also reflects content-specific activation (i.e., whether the activation in S1 is specific to the mental content participants imagined). To this end, healthy volunteers (n = 21) either perceived or imagined three types of vibrotactile stimuli (mental content) while fMRI data were acquired. Independent of the content, during tactile mental imagery we found activation of frontoparietal regions, supplemented with activation in the contralateral BA2 subregion of S1, replicating previous reports. While the imagery of the three different stimuli did not reveal univariate activation differences, using multivariate pattern classification, we were able to decode the imagined stimulus type from BA2. Moreover, cross-classification revealed that tactile imagery elicits activation patterns similar to those evoked by the perception of the respective stimuli. These findings promote the idea that mental tactile imagery involves the recruitment of content-specific activation patterns in sensory cortices, namely in S1.