Philipp Ruhnau

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.

Finding the right control: The mismatch negativity under investigation

OBJECTIVE When investigating auditory perceptual regularity processing, mismatch negativity (MMN) is commonly used. MMN is computed as a difference signal between the event-related potentials (ERPs) elicited by repeated standard tones and rarely occurring deviant tones. This procedure leads to an underestimation of the N1 component elicited by standards compared to the N1 to deviants which might affect the MMN. Consequently, a random control design was previously introduced. This design, however, overestimates the N1 to the deviant. Here, we developed a new paradigm that avoids previous drawbacks. METHODS We designed a regular cascadic sequence as a control to the deviant. ERPs were measured while presenting conventional oddball blocks (standards, deviants), random control blocks and a cascadic control block. RESULTS MMN was observed in each difference signal. Regarding the N1, standards elicited smallest amplitudes. The N1 for the deviant and the cascadic control was comparable. The largest N1 was elicited by the random control. CONCLUSION Standards underestimate N1 refractoriness effects in the responses to deviants, while random control tones overestimate. Cascadic control tones, however, provide a reasonable estimation for the N1. SIGNIFICANCE The new cascadic control design is suitable to investigate auditory perceptual regularity processes while controlling for N1 refractoriness effects.

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.

Stimulus-Driven Brain Oscillations in the Alpha Range: Entrainment of Intrinsic Rhythms or Frequency-Following Response?

Human brain activity is rich in rhythms of various characteristic frequencies. The last few decades have seen an increase in their use as an explanatory means, with a vast literature describing manifold correlations between dynamics of brain rhythms and behavioral performance in perceptual and

The modulation of auditory novelty processing by working memory load in school age children and adults: a combined behavioral and event-related potential study

BackgroundWe investigated the processing of task-irrelevant and unexpected novel sounds and its modulation by working-memory load in children aged 9-10 and in adults. Environmental sounds (novels) were embedded amongst frequently presented standard sounds in an auditory-visual distraction paradigm. Each sound was followed by a visual target. In two conditions, participants evaluated the position of a visual stimulus (0-back, low load) or compared the position of the current stimulus with the one two trials before (2-back, high load). Processing of novel sounds were measured with reaction times, hit rates and the auditory event-related brain potentials (ERPs) Mismatch Negativity (MMN), P3a, Reorienting Negativity (RON) and visual P3b.ResultsIn both memory load conditions novels impaired task performance in adults whereas they improved performance in children. Auditory ERPs reflect age-related differences in the time-window of the MMN as children showed a positive ERP deflection to novels whereas adults lack an MMN. The attention switch towards the task irrelevant novel (reflected by P3a) was comparable between the age groups. Adults showed more efficient reallocation of attention (reflected by RON) under load condition than children. Finally, the P3b elicited by the visual target stimuli was reduced in both age groups when the preceding sound was a novel.ConclusionOur results give new insights in the development of novelty processing as they (1) reveal that task-irrelevant novel sounds can result in contrary effects on the performance in a visual primary task in children and adults, (2) show a positive ERP deflection to novels rather than an MMN in children, and (3) reveal effects of auditory novels on visual target processing.

Maturation of obligatory auditory responses and their neural sources: Evidence from EEG and MEG

Neural auditory responses are known to change from childhood to adulthood. The most prominent components of the event-related potentials (ERPs) found in children are the P1 and N2, while the P1 and N1 are strongest in adults. Previous dipole localizations showed regions of the auditory cortex (AC) underlying these responses. An N1 in children, however, has only been observed in older age or under certain experimental conditions different than commonly applied in adults. The current study aimed to further elucidate on auditory processing and related components in school-aged children. To do this, MEG and EEG was recorded in adults and 9 to 10year old children, while presenting pure tones either repetitively or randomly among tones of different pitch. Furthermore, the current paradigm was explicitly designed to not only investigate the P1 and N2 in children, but moreover to examine N1 modulations based on different refractory states caused by the two conditions. Our results are clear cut. In adults, P1(m) and N1(m) components were localized in AC regions, with the N1(m) largely attenuated for repetitive tones. The P1(m) and N2(m) components observed in children were also localized in AC regions. Most importantly, ERP modulations in the N1 time window (i.e., larger responses for random than repetitive tones) were remarkably similar for adults and children, both in amplitude and latency. This effect indicates that the N1 sub-component reflecting frequency-specific refractoriness is fully developed in 9 to 10year old children. Thus, previous interpretations on the function and maturation of the N1 need reconsideration.

Investigating ongoing brain oscillations and their influence on conscious perception - network states and the window to consciousness.

In cognitive neuroscience, prerequisites of consciousness are of high interest. Within recent years it has become more commonly understood that ongoing brain activity, mainly measured with electrophysiology, can predict whether an upcoming stimulus is consciously perceived. One approach to investigate the relationship between ongoing brain activity and conscious perception is to conduct near-threshold (NT) experiments and focus on the pre-stimulus period. The current review will, in the first part, summarize main findings of pre-stimulus research from NT experiments, mainly focusing on the alpha band (8–14 Hz). It is probable that the most prominent finding is that local (mostly sensory) areas show enhanced excitatory states prior to detection of upcoming NT stimuli, as putatively reflected by decreased alpha band power. However, the view of a solely local excitability change seems to be too narrow. In a recent paper, using a somatosensory NT task, Weisz et al. (2014) replicated the common alpha finding and, furthermore, conceptually embedded this finding into a more global framework called “Windows to Consciousness” (Win2Con). In this review, we want to further elaborate on the crucial assumption of “open windows” to conscious perception, determined by pre-established pathways connecting sensory and higher order areas. Methodologically, connectivity and graph theoretical analyses are applied to source-imaging magnetoencephalographic data to uncover brain regions with strong network integration as well as their connection patterns. Sensory regions with stronger network integration will more likely distribute information when confronted with weak NT stimuli, favoring its subsequent conscious perception. First experimental evidence confirms our aforementioned “open window” hypothesis. We therefore emphasize that future research on prerequisites of consciousness needs to move on from investigating solely local excitability to a more global view of network connectivity.

Processing of complex distracting sounds in school-aged children and adults: evidence from EEG and MEG data.

When a perceiver performs a task, rarely occurring sounds often have a distracting effect on task performance. The neural mismatch responses in event-related potentials to such distracting stimuli depend on age. Adults commonly show a negative response, whereas in children a positive as well as a negative mismatch response has been reported. Using electro- and magnetoencephalography (EEG/MEG), here we investigated the developmental changes of distraction processing in school-aged children (9–10 years) and adults. Participants took part in an auditory-visual distraction paradigm comprising a visuo-spatial primary task and task-irrelevant environmental sounds distracting from this task. Behaviorally, distractors delayed reaction times (RTs) in the primary task in both age groups, and this delay was of similar magnitude in both groups. The neurophysiological data revealed an early as well as a late mismatch response elicited by distracting stimuli in both age groups. Together with previous research, this indicates that deviance detection is accomplished in a hierarchical manner in the auditory system. Both mismatch responses were localized to auditory cortex areas. All mismatch responses were generally delayed in children, suggesting that not all neurophysiological aspects of deviance processing are mature in school-aged children. Furthermore, the P3a, reflecting involuntary attention capture, was present in both age groups in the EEG with comparable amplitudes and at similar latencies, but with a different topographical distribution. This suggests that involuntary attention shifts toward complex distractors operate comparably in school-aged children and adults, yet undergoing generator maturation.

Prestimulus Network Integration of Auditory Cortex Predisposes Near-Threshold Perception Independently of Local Excitability

An ever-increasing number of studies are pointing to the importance of network properties of the brain for understanding behavior such as conscious perception. However, with regards to the influence of prestimulus brain states on perception, this network perspective has rarely been taken. Our recent framework predicts that brain regions crucial for a conscious percept are coupled prior to stimulus arrival, forming pre-established pathways of information flow and influencing perceptual awareness. Using magnetoencephalography (MEG) and graph theoretical measures, we investigated auditory conscious perception in a near-threshold (NT) task and found strong support for this framework. Relevant auditory regions showed an increased prestimulus interhemispheric connectivity. The left auditory cortex was characterized by a hub-like behavior and an enhanced integration into the brain functional network prior to perceptual awareness. Right auditory regions were decoupled from non-auditory regions, presumably forming an integrated information processing unit with the left auditory cortex. In addition, we show for the first time for the auditory modality that local excitability, measured by decreased alpha power in the auditory cortex, increases prior to conscious percepts. Importantly, we were able to show that connectivity states seem to be largely independent from local excitability states in the context of a NT paradigm.

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.