Abstract

BackgroundConsiderable effort has been devoted to mapping the functional and effective connectivity of the human brain, but these efforts have largely been limited to tasks involving stationary subjects. Recent advances with high-density electroencephalography (EEG) and Independent Components Analysis (ICA) have enabled study of electrocortical activity during human locomotion. The goal of this work was to measure the effective connectivity of cortical activity during human standing and walking.MethodsWe recorded 248-channels of EEG as eight young healthy subjects stood and walked on a treadmill both while performing a visual oddball discrimination task and not performing the task. ICA parsed underlying electrocortical, electromyographic, and artifact sources from the EEG signals. Inverse source modeling methods and clustering algorithms localized posterior, anterior, prefrontal, left sensorimotor, and right sensorimotor clusters of electrocortical sources across subjects. We applied a directional measure of connectivity, conditional Granger causality, to determine the effective connectivity between electrocortical sources.ResultsConnections involving sensorimotor clusters were weaker for walking than standing regardless of whether the subject was performing the simultaneous cognitive task or not. This finding supports the idea that cortical involvement during standing is greater than during walking, possibly because spinal neural networks play a greater role in locomotor control than standing control. Conversely, effective connectivity involving non-sensorimotor areas was stronger for walking than standing when subjects were engaged in the simultaneous cognitive task.ConclusionsOur results suggest that standing results in greater functional connectivity between sensorimotor cortical areas than walking does. Greater cognitive attention to standing posture than to walking control could be one interpretation of that finding. These techniques could be applied to clinical populations during gait to better investigate neural substrates involved in mobility disorders.

Highlights

  • Considerable effort has been devoted to mapping the functional and effective connectivity of the human brain, but these efforts have largely been limited to tasks involving stationary subjects

  • We have recently demonstrated that high-density electroencephalography (EEG) combined with Independent Component Analysis (ICA) enables the study of electrocortical activity related to locomotor control and cognition during walking [7,8,9]

  • Effective connectivities involving nonsensorimotor areas were significantly stronger during walking than standing only when subjects were engaged in the simultaneous cognitive task (p < 0.03) (Figure 2b, red bars)

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Summary

Introduction

Considerable effort has been devoted to mapping the functional and effective connectivity of the human brain, but these efforts have largely been limited to tasks involving stationary subjects. Recent advances with high-density electroencephalography (EEG) and Independent Components Analysis (ICA) have enabled study of electrocortical activity during human locomotion. One way to study brain connectivity during more natural, whole body, behaviors is to combine high-density electroencephalography (EEG), Independent Component Analysis (ICA), and source localization techniques. We have recently demonstrated that high-density electroencephalography (EEG) combined with Independent Component Analysis (ICA) enables the study of electrocortical activity related to locomotor control and cognition during walking [7,8,9]. Clustering electrocortical sources across subjects according to spatial and spectral properties enables calculation of changes in effective connectivity between cortical regions using techniques such as Granger causality [10]

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