Alpha power indexes task-related networks on large and small scales: A multimodal ECoG study in humans and a non-human primate.

Performing different tasks, such as generating motor movements or processing sensory input, requires the recruitment of specific networks of neuronal populations. Previous studies suggested that power variations in the alpha band (8-12Hz) may implement such recruitment of task-specific populations by increasing cortical excitability in task-related areas while inhibiting population-level cortical activity in task-unrelated areas (Klimesch et al., 2007; Jensen and Mazaheri, 2010). However, the precise temporal and spatial relationships between the modulatory function implemented by alpha oscillations and population-level cortical activity remained undefined. Furthermore, while several studies suggested that alpha power indexes task-related populations across large and spatially separated cortical areas, it was largely unclear whether alpha power also differentially indexes smaller networks of task-related neuronal populations. Here we addressed these questions by investigating the temporal and spatial relationships of electrocorticographic (ECoG) power modulations in the alpha band and in the broadband gamma range (70-170Hz, indexing population-level activity) during auditory and motor tasks in five human subjects and one macaque monkey. In line with previous research, our results confirm that broadband gamma power accurately tracks task-related behavior and that alpha power decreases in task-related areas. More importantly, they demonstrate that alpha power suppression lags population-level activity in auditory areas during the auditory task, but precedes it in motor areas during the motor task. This suppression of alpha power in task-related areas was accompanied by an increase in areas not related to the task. In addition, we show for the first time that these differential modulations of alpha power could be observed not only across widely distributed systems (e.g., motor vs. auditory system), but also within the auditory system. Specifically, alpha power was suppressed in the locations within the auditory system that most robustly responded to particular sound stimuli. Altogether, our results provide experimental evidence for a mechanism that preferentially recruits task-related neuronal populations by increasing cortical excitability in task-related cortical areas and decreasing cortical excitability in task-unrelated areas. This mechanism is implemented by variations in alpha power and is common to humans and the non-human primate under study. These results contribute to an increasingly refined understanding of the mechanisms underlying the selection of the specific neuronal populations required for task execution.

The effects of spatial filtering and artifacts on electrocorticographic signals.

OBJECTIVE: Electrocorticographic (ECoG) signals contain noise that is common to all channels and noise that is specific to individual channels. Most published ECoG studies use common average reference (CAR) spatial filters to remove common noise, but CAR filters may introduce channel-specific noise into other channels. To address this concern, scientists often remove artifactual channels prior to data analysis. However, removing these channels depends on expert-based labeling and may also discard useful data. Thus, the effects of spatial filtering and artifacts on ECoG signals have been largely unknown. This study aims to quantify these effects and thereby address this gap in knowledge. APPROACH: In this study, we address these issues by exploring the effects of application of two types of unsupervised spatial filters and three methods of detecting signal artifacts using a large ECoG data set (20 subjects, four task conditions in each subject). MAIN RESULTS: Our results confirm that spatial filtering improves performance, i.e., it reduces ECoG signal variance that is not related to the task. They also show that removing artifactual channels automatically (using quantitatively defined rejection criteria) or manually (using expert opinion) does not increase the total amount of task-related information, but does avoid potential contamination from one or more noisy channels. Finally, applying a novel 'median average reference' filter does not require the elimination of artifactual channels prior to spatial filtering and still mitigates the influence of channels with channel-specific noise. Thus, it allows the investigator to retain more potentially useful task-related data. SIGNIFICANCE: In summary, our results show that appropriately designed spatial filters that account for both common noise and channel-specific noise greatly improve the quality of ECoG signal analyses, and that artifacts in only a single channel can result in profound and undesired effects on all other channels.