Burkhard Maess

Musical syntax is processed in Broca's area: an MEG study

The present experiment was designed to localize the neural substrates that process music-syntactic incongruities, using magnetoencephalography (MEG). Electrically, such processing has been proposed to be indicated by early right-anterior negativity (ERAN), which is elicited by harmonically inappropriate chords occurring within a major-minor tonal context. In the present experiment, such chords elicited an early effect, taken as the magnetic equivalent of the ERAN (termed mERAN). The source of mERAN activity was localized in Brocas area and its right-hemisphere homologue, areas involved in syntactic analysis during auditory language comprehension. We find that these areas are also responsible for an analysis of incoming harmonic sequences, indicating that these regions process syntactic information that is less language-specific than previously believed.

Good practice for conducting and reporting MEG research

Magnetoencephalographic (MEG) recordings are a rich source of information about the neural dynamics underlying cognitive processes in the brain, with excellent temporal and good spatial resolution. In recent years there have been considerable advances in MEG hardware developments and methods. Sophisticated analysis techniques are now routinely applied and continuously improved, leading to fascinating insights into the intricate dynamics of neural processes. However, the rapidly increasing level of complexity of the different steps in a MEG study make it difficult for novices, and sometimes even for experts, to stay aware of possible limitations and caveats. Furthermore, the complexity of MEG data acquisition and data analysis requires special attention when describing MEG studies in publications, in order to facilitate interpretation and reproduction of the results. This manuscript aims at making recommendations for a number of important data acquisition and data analysis steps and suggests details that should be specified in manuscripts reporting MEG studies. These recommendations will hopefully serve as guidelines that help to strengthen the position of the MEG research community within the field of neuroscience, and may foster discussion in order to further enhance the quality and impact of MEG research.

Semantic Category Interference in Overt Picture Naming: Sharpening Current Density Localization by PCA

The study investigated the neuronal basis of the retrieval of words from the mental lexicon. The semantic category interference effect was used to locate lexical retrieval processes in time and space. This effect reflects the finding that, for overt naming, volunteers are slower when naming pictures out of a sequence of items from the same semantic category than from different categories. Participants named pictures blockwise either in the context of same-or mixed-category items while the brain response was registered using magnetoencephalography (MEG). Fifteen out of 20 participants showed longer response latencies in the same-category compared to the mixed-category condition. Event-related MEG signals for the participants demonstrating the interference effect were submitted to a current source density (CSD) analysis. As a new approach, a principal component analysis was applied to decompose the grand average CSD distribution into spatial subcomponents (factors). The spatial factor indicating left temporal activity revealed significantly different activation for the same-category compared to the mixed-category condition in the time window between 150 and 225 msec post picture onset. These findings indicate a major involvement of the left temporal cortex in the semantic interference effect. As this effect has been shown to take place at the level of lexical selection, the data suggest that the left temporal cortex supports processes of lexical retrieval during production.

Memory-matches evoke human gamma-responses

BackgroundHuman brain activity in the gamma frequency range has been shown to be a correlate of numerous cognitive functions like attention, perception and memory access. More specifically, gamma activity has been found to be enhanced when stimuli are stored in or match with short-term memory (STM). We tested the hypothesis that gamma activity is also evoked when stimuli match representations in long-term-memory (LTM). EEG was recorded from 13 subjects performing a choice reaction task. Visual stimuli were either known real-world objects with a memory representation or novel configurations never seen before.ResultsAll stimuli evoked an early gamma response which was maximal over occipital electrodes. This evoked gamma activity was significantly larger for items that matched memory templates.ConclusionsTherefore, we argue that gamma activity results from the feedback from memory into perception systems. This assumption seems to be true for STM as well as LTM.

Localizing pre-attentive auditory memory-based comparison: Magnetic mismatch negativity to pitch change

Changes in the pitch of repetitive sounds elicit the mismatch negativity (MMN) of the event-related brain potential (ERP). There exist two alternative accounts for this index of automatic change detection: (1) A sensorial, non-comparator account according to which ERPs in oddball sequences are affected by differential refractory states of frequency-specific afferent cortical neurons. (2) A cognitive, comparator account stating that MMN reflects the outcome of a memory comparison between a neuronal model of the frequently presented standard sound with the sensory memory representation of the changed sound. Using a condition controlling for refractoriness effects, the two contributions to MMN can be disentangled. The present study used whole-head MEG to further elucidate the sensorial and cognitive contributions to frequency MMN. Results replicated ERP findings that MMN to pitch change is a compound of the activity of a sensorial, non-comparator mechanism and a cognitive, comparator mechanism which could be separated in time. The sensorial part of frequency MMN consisting of spatially dipolar patterns was maximal in the late N1 range (105-125 ms), while the cognitive part peaked in the late MMN-range (170-200 ms). Spatial principal component analyses revealed that the early part of the traditionally measured MMN (deviant minus standard) is mainly due to the sensorial mechanism while the later mainly due to the cognitive mechanism. Inverse modeling revealed sources for both MMN contributions in the gyrus temporales transversus, bilaterally. These MEG results suggest temporally distinct but spatially overlapping activities of non-comparator-based and comparator-based mechanisms of automatic frequency change detection in auditory cortex.

Digital filter design for electrophysiological data – A practical approach

BACKGROUND Filtering is a ubiquitous step in the preprocessing of electroencephalographic (EEG) and magnetoencephalographic (MEG) data. Besides the intended effect of the attenuation of signal components considered as noise, filtering can also result in various unintended adverse filter effects (distortions such as smoothing) and filter artifacts. METHOD We give some practical guidelines for the evaluation of filter responses (impulse and frequency response) and the selection of filter types (high-pass/low-pass/band-pass/band-stop; finite/infinite impulse response, FIR/IIR) and filter parameters (cutoff frequencies, filter order and roll-off, ripple, delay and causality) to optimize signal-to-noise ratio and avoid or reduce signal distortions for selected electrophysiological applications. RESULTS Various filter implementations in common electrophysiology software packages are introduced and discussed. Resulting filter responses are compared and evaluated. CONCLUSION We present strategies for recognizing common adverse filter effects and filter artifacts and demonstrate them in practical examples. Best practices and recommendations for the selection and reporting of filter parameters, limitations, and alternatives to filtering are discussed.

Perception of phrase structure in music

Neither music nor spoken language form uniform auditory streams, rather, they are structured into phrases. For the perception of such structures, the detection of phrase boundaries is crucial. We discovered electroencephalography (EEG) and magnetoencephalography (MEG) correlates for the perception of phrase boundaries in music. In EEG, this process was marked by a positive wave approximately between 500 and 600 ms after the offset of a phrase boundary with a centroparietal maximum. In MEG, we found major activity in an even broader time window (400–700 ms). Source localization revealed that likely candidates for the generation of the observed effects are structures in the limbic system, including anterior and posterior cingulate as well as posterior mediotemporal cortex. The timing and topography of the EEG effect bear some resemblance to a positive shift (closure positive shift, CPS) found for prosodic phrase boundaries during speech perception in an earlier study, suggesting that the underlying processes might be related. Because the brain structures, which possibly underlie the observed effects, are known to be involved in memory and attention processes, we suggest that the CPS may not reflect the detection of the phrase boundary as such, but those memory and attention related processes that are necessary to guide the attention focus from one phrase to the next, thereby closing the former and opening up the next phrase. Hum Brain Mapp 24:259–273, 2005.

Adverse Listening Conditions and Memory Load Drive a Common Alpha Oscillatory Network

How does acoustic degradation affect the neural mechanisms of working memory? Enhanced alpha oscillations (8–13 Hz) during retention of items in working memory are often interpreted to reflect increased demands on storage and inhibition. We hypothesized that auditory signal degradation poses an additional challenge to human listeners partly because it draws on the same neural mechanisms. In an adapted Sternberg paradigm, auditory memory load and acoustic degradation were parametrically varied and the magnetoencephalographic response was analyzed in the time–frequency domain. Notably, during the stimulus-free delay interval, alpha power monotonically increased at central–parietal sensors as functions of memory load (higher alpha power with more memory load) and of acoustic degradation (also higher alpha power with more severe acoustic degradation). This alpha effect was superadditive when highest load was combined with most severe degradation. Moreover, alpha oscillatory dynamics during stimulus-free delay were predictive of response times to the probe item. Source localization of alpha power during stimulus-free delay indicated that alpha generators in right parietal, cingulate, supramarginal, and superior temporal cortex were sensitive to combined memory load and acoustic degradation. In summary, both challenges of memory load and acoustic degradation increase activity in a common alpha-frequency network. The results set the stage for future studies on how chronic or acute degradations of sensory input affect mechanisms of executive control.

Localizing the distributed language network responsible for the N400 measured by MEG during auditory sentence processing

We studied auditory sentence comprehension using magnetoencephalography while subjects listened to sentences whose correctness they had to judge subsequently. The localization and the time course of brain electrical activity during processing of correct and semantically incorrect sentences were estimated by computing a brain surface current density within a cortical layer for both conditions. Finally, a region of interest (ROI) analysis was conducted to determine the time course of specific locations. A magnetic N400 was present in six spatially different ROIs. Semantic anomalies caused an exclusive involvement of the ventral portion of the left inferior frontal gyrus (BA 47) and left pars triangularis (BA 45). The anterior parts of the superior (BA 22) and inferior (BA 20/21) temporal gyri bilaterally were activated by both conditions. The activation for the correct condition, however, peaked earlier in both left temporal regions (approximately 32 ms). In general, activation due to semantic violations was more pronounced, started later, and lasted longer as compared to correct sentences. The findings reveal a clear left-hemispheric dominance during language processing indicated firstly by the mere number of activated regions (four in the left vs. two in the right hemisphere) and secondly by the observed specificity of the left inferior frontal ROIs to semantic violations. The temporal advantage observed for the correct condition in the left temporal regions is supporting the notion that the established context eases the processing of the final word. Semantically incorrect words that do not fit into the context result in longer integration times.

A MEG analysis of the P300 in visual discrimination tasks

Based on recent research that indicated that P300 scalp topography varies as a function of task and/or information to be processed, this study examined scalp-recorded magnetic fields correlated with the P300 by means of whole-head magnetoencephalography. Subjects performed two discrimination tasks, in which targets, defined on either object or spatial characteristics of the same visual stimuli, had to be discriminated. Based on the across-subject root mean square (RMS) functions a sequence of 4 components could be identified in both tasks, N1m, P3m, and two later components, which, based on their estimated neuronal sources, were classified as representing motor processes during and following the manual responses to target stimuli. Reliable between-task differences in source localization were obtained for the P3m component, but not for the other components. Inferior-medial sources were found for the P3m evoked by both spatial and object targets, with these sources being located about 3.5 cm more anterior for object targets. These results suggest that different neuronal sources, possibly located in subcortical regions in the vicinity of the thalamus, contribute to the P3m evoked by target stimuli defined by either object or spatial stimulus characteristics.