Gianpaolo Demarchi

Friends, not foes: Magnetoencephalography as a tool to uncover brain dynamics during transcranial alternating current stimulation

Brain oscillations are supposedly crucial for normal cognitive functioning and alterations are associated with cognitive dysfunctions. To demonstrate their causal role on behavior, entrainment approaches in particular aim at driving endogenous oscillations via rhythmic stimulation. Within this context, transcranial electrical stimulation, especially transcranial alternating current stimulation (tACS), has received renewed attention. This is likely due to the possibility of defining oscillatory stimulation properties precisely. Also, measurements comparing pre-tACS with post-tACS electroencephalography (EEG) have shown impressive modulations. However, the period during tACS has remained a blackbox until now, due to the enormous stimulation artifact. By means of application of beamforming to magnetoencephalography (MEG) data, we successfully recovered modulations of the amplitude of brain oscillations during weak and strong tACS. Additionally, we demonstrate that also evoked responses to visual and auditory stimuli can be recovered during tACS. The main contribution of the present study is to provide critical evidence that during ongoing tACS, subtle modulations of oscillatory brain activity can be reconstructed even at the stimulation frequency. Future tACS experiments will be able to deliver direct physiological insights in order to further the understanding of the contribution of brain oscillations to cognition and behavior.

Eyes wide shut: Transcranial alternating current stimulation drives alpha rhythm in a state dependent manner

Transcranial alternating current stimulation (tACS) is used to modulate brain oscillations to measure changes in cognitive function. It is only since recently that brain activity in human subjects during tACS can be investigated. The present study aims to investigate the phase relationship between the external tACS signal and concurrent brain activity. Subjects were stimulated with tACS at individual alpha frequency during eyes open and eyes closed resting states. Electrodes were placed at Cz and Oz, which should affect parieto-occipital areas most strongly. Source space magnetoencephalography (MEG) data were used to estimate phase coherence between tACS and brain activity. Phase coherence was significantly increased in areas in the occipital pole in eyes open resting state only. The lag between tACS and brain responses showed considerable inter-individual variability. In conclusion, tACS at individual alpha frequency entrains brain activity in visual cortices. Interestingly, this effect is state dependent and is clearly observed with eyes open but only to a lesser extent with eyes closed.

The contribution of primary and secondary somatosensory cortices to the representation of body parts and body sides: An fmri adaptation study

Although the somatosensory homunculus is a classically used description of the way somatosensory inputs are processed in the brain, the actual contributions of primary (SI) and secondary (SII) somatosensory cortices to the spatial coding of touch remain poorly understood. We studied adaptation of the fMRI BOLD response in the somatosensory cortex by delivering pairs of vibrotactile stimuli to the finger tips of the index and middle fingers. The first stimulus (adaptor) was delivered either to the index or to the middle finger of the right or left hand, and the second stimulus (test) was always administered to the left index finger. The overall BOLD response evoked by the stimulation was primarily contralateral in SI and was more bilateral in SII. However, our fMRI adaptation approach also revealed that both somatosensory cortices were sensitive to ipsilateral as well as to contralateral inputs. SI and SII adapted more after subsequent stimulation of homologous as compared with nonhomologous fingers, showing a distinction between different fingers. Most importantly, for both somatosensory cortices, this finger-specific adaptation occurred irrespective of whether the tactile stimulus was delivered to the same or to different hands. This result implies integration of contralateral and ipsilateral somatosensory inputs in SI as well as in SII. Our findings suggest that SI is more than a simple relay for sensory information and that both SI and SII contribute to the spatial coding of touch by discriminating between body parts (fingers) and by integrating the somatosensory input from the two sides of the body (hands).

Waves of regret: A MEG study of emotion and decision-making

Recent fMRI studies have investigated brain activity involved in the feeling of regret and disappointment by manipulating the feedback participants saw after making a decision to play certain gambles: full-feedback (regret: participant sees the outcomes from both the chosen and unchosen gamble) vs. partial-feedback (disappointment: participant only sees the outcome from chosen gamble). However, regret and disappointment are also characterized by differential agency attribution: personal agency for regret, external agency for disappointment. In this study, we investigate the neural correlates of these two characterizations of regret and disappointment using magnetoencephalography (MEG). To do this, we experimentally induced each emotion by manipulating feedback (chosen gamble vs. unchosen gamble), agency (human vs. computer choice) and outcomes (win vs. loss) in a fully randomized design. At the behavioral level the emotional experience of regret and disappointment were indeed affected by both feedback and agency manipulations. These emotions also differentially affect subsequent choices, with regret leading to riskier behavior. At the neural level both feedback and agency affected the brain responses associated with regret and disappointment, demonstrating differential localization in the brain for each. Notably, feedback regret showed greater brain activity in the right anterior and posterior regions, with agency regret producing greater activity in the left anterior region. These findings extend the evidence for neural activity in processing both regret and disappointment by highlighting for the first time the respective importance of feedback and agency, as well as outlining the temporal dynamics of these emotions.

Faith and oscillations recovered: On analyzing EEG/MEG signals during tACS

ABSTRACT Despite recent success in analyzing brain oscillations recorded during transcranial alternating current stimulation (tACS), the field still requires further research to establish standards in artifact removal methods. This includes taking a step back from the removal of the tACS artifact and thoroughly characterizing the to‐be‐removed artifact. A recent study by Noury et al. (2016) contributed importantly to this endeavour by showing the existence of nonlinear artefacts in the tACS signal as seen by MEG and EEG. Unfortunately however this paper conveys the message that current artifact removal attempts have failed altogether and that—based on these available tools—brain oscillations recorded during tACS cannot be analyzed using MEG and EEG. Here we want to balance this overly pessimistic conclusion: In‐depth reanalyses of our own data and phantom‐head measurements indicate that nonlinearities can occur, but only when technical limits of the stimulator are reached. As such they are part of the “real” stimulation and not a specific MEG analysis problem. Future tACS studies should consider these technical limits to avoid any nonlinear modulations of the tACS artifact. We conclude that even with current approaches, brain oscillations recorded during tACS can be meaningfully studied in many practical cases.

Neural Correlates of Finger Gnosis

Neuropsychological studies have described patients with a selective impairment of finger identification in association with posterior parietal lesions. However, evidence of the role of these areas in finger gnosis from studies of the healthy human brain is still scarce. Here we used functional magnetic resonance imaging to identify the brain network engaged in a novel finger gnosis task, the intermanual in-between task (IIBT), in healthy participants. Several brain regions exhibited a stronger blood oxygenation level-dependent (BOLD) response in IIBT than in a control task that did not explicitly rely on finger gnosis but used identical stimuli and motor responses as the IIBT. The IIBT involved stronger signal in the left inferior parietal lobule (IPL), bilateral precuneus (PCN), bilateral premotor cortex, and left inferior frontal gyrus. In all regions, stimulation of nonhomologous fingers of the two hands elicited higher BOLD signal than stimulation of homologous fingers. Only in the left anteromedial IPL (a-mIPL) and left PCN did signal strength decrease parametrically from nonhomology, through partial homology, to total homology with stimulation delivered synchronously to the two hands. With asynchronous stimulation, the signal was stronger in the left a-mIPL than in any other region, possibly indicating retention of task-relevant information. We suggest that the left PCN may contribute a supporting visuospatial representation via its functional connection to the right PCN. The a-mIPL may instead provide the core substrate of an explicit bilateral body structure representation for the fingers that when disrupted can produce the typical symptoms of finger agnosia.

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.

Cortical Reorganization after Damage to the Central Nervous System

Findings regarding cortical reorganization have raised hope for the development of new rehabilitation therapies in brain damaged patients. Powerful therapeutic concepts have been developed. However, as the understanding of cortical reorganization has evolved, limits of cortical reorganization have become also more evident. In humans, the knowledge about cortical reorganization relies mainly on functional imaging methods that infer changes in activation patterns of the brain. Based on studies that have been performed by the authors, it is discussed whether altered activity patterns in brain damaged patients are indeed related to cortical reorganization and to the recovery of brain functions.

Effects of looming and static sounds on somatosensory processing: A MEG study

The present study aims to assess the mechanisms involved in the processing of potentially threatening stimuli presented within the peri-head space of humans. Magnetic fields evoked by air-puffs presented at the peri-oral area of fifteen participants were recorded by using magnetoencephalography (MEG). Crucially, each air puff was preceded by a sound, which could be either perceived as looming, stationary and close to the body (i.e., within the peri-head space) or stationary and far from the body (i.e., extrapersonal space). The comparison of the time courses of the global field power (GFP) indicated a significant difference in the time window ranging from 70 to 170 ms between the conditions. When the air puff was preceded by a stationary sound located far from the head stronger somatosensory activity was evoked as compared to the conditions where the sounds were located close to the head. No difference could be shown for the looming and the stationary prime stimulus close to the head. Source localization was performed assuming a pair of symmetric dipoles in a spherical head model that was fitted to the MRI images of the individual participants. Results showed sources in primary and secondary somatosensory cortex. Source activities in secondary somatosensory cortex differed between the three conditions, with larger effects evoked by the looming sounds and smaller effects evoked by the far stationary sounds, and the close stationary sounds evoking intermediate effects. Overall, these findings suggest the existence of a system involved in the detection of approaching objects and protecting the body from collisions in humans.