Walking reduces sensorimotor network connectivity compared to standing.

BACKGROUND: 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. The goal of this work was to measure the effective connectivity of cortical activity during human standing and walking. METHODS: We 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. RESULTS: Connections 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. CONCLUSIONS: Our 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.

Weighted phase lag index stability as an artifact resistant measure to detect cognitive EEG activity during locomotion.

BACKGROUND: High-density electroencephalography (EEG) with active electrodes allows for monitoring of electrocortical dynamics during human walking but movement artifacts have the potential to dominate the signal. One potential method for recovering cognitive brain dynamics in the presence of gait-related artifact is the Weighted Phase Lag Index. METHODS: We tested the ability of Weighted Phase Lag Index to recover event-related potentials during locomotion. Weighted Phase Lag Index is a functional connectivity measure that quantified how consistently 90° (or 270°) phase 'lagging' one EEG signal was compared to another. 248-channel EEG was recorded as eight subjects performed a visual oddball discrimination and response task during standing and walking (0.8 or 1.2 m/s) on a treadmill. RESULTS: Applying Weighted Phase Lag Index across channels we were able to recover a p300-like cognitive response during walking. This response was similar to the classic amplitude-based p300 we also recovered during standing. We also showed that the Weighted Phase Lag Index detects more complex and variable activity patterns than traditional voltage-amplitude measures. This variability makes it challenging to compare brain activity over time and across subjects. In contrast, a statistical metric of the index's variability, calculated over a moving time window, provided a more generalized measure of behavior. Weighted Phase Lag Index Stability returned a peak change of 1.8% + -0.5% from baseline for the walking case and 3.9% + -1.3% for the standing case. CONCLUSIONS: These findings suggest that both Weighted Phase Lag Index and Weighted Phase Lag Index Stability have potential for the on-line analysis of cognitive dynamics within EEG during human movement. The latter may be more useful from extracting general principles of neural behavior across subjects and conditions.

Local dynamics of gap-junction-coupled interneuron networks.

Interneurons coupled by both electrical gap-junctions (GJs) and chemical GABAergic synapses are major components of forebrain networks. However, their contributions to the generation of specific activity patterns, and their overall contributions to network function, remain poorly understood. Here we demonstrate, using computational methods, that the topological properties of interneuron networks can elicit a wide range of activity dynamics, and either prevent or permit local pattern formation. We systematically varied the topology of GJ and inhibitory chemical synapses within simulated networks, by changing connection types from local to random, and changing the total number of connections. As previously observed we found that randomly coupled GJs lead to globally synchronous activity. In contrast, we found that local GJ connectivity may govern the formation of highly spatially heterogeneous activity states. These states are inherently temporally unstable when the input is uniformly random, but can rapidly stabilize when the network detects correlations or asymmetries in the inputs. We show a correspondence between this feature of network activity and experimental observations of transient stabilization of striatal fast-spiking interneurons (FSIs), in electrophysiological recordings from rats performing a simple decision-making task. We suggest that local GJ coupling enables an active search-and-select function of striatal FSIs, which contributes to the overall role of cortical-basal ganglia circuits in decision-making.

The resonance frequency shift, pattern formation, and dynamical network reorganization via sub-threshold input.

We describe a novel mechanism that mediates the rapid and selective pattern formation of neuronal network activity in response to changing correlations of sub-threshold level input. The mechanism is based on the classical resonance and experimentally observed phenomena that the resonance frequency of a neuron shifts as a function of membrane depolarization. As the neurons receive varying sub-threshold input, their natural frequency is shifted in and out of its resonance range. In response, the neuron fires a sequence of action potentials, corresponding to the specific values of signal currents, in a highly organized manner. We show that this mechanism provides for the selective activation and phase locking of the cells in the network, underlying input-correlated spatio-temporal pattern formation, and could be the basis for reliable spike-timing dependent plasticity. We compare the selectivity and efficiency of this pattern formation to a supra-threshold network activation and a non-resonating network/neuron model to demonstrate that the resonance mechanism is the most effective. Finally we show that this process might be the basis of the phase precession phenomenon observed during firing of hippocampal place cells, and that it may underlie the active switching of neuronal networks to locking at various frequencies.

Interaction between connectivity and oscillatory currents in a heterogeneous neuronal network.

Intrinsic oscillations are thought to play important and distinct roles in cognitive processes across nearly all regions of the brain. Their specific roles are highly dependent on their properties: low-frequency θ is thought to be important in the gating of cognitive processes, while high-frequency γ is believed to be essential for binding and spike-timing-dependent plasticity. We investigated the role of an oscillatory drive for pattern formation of heterogeneous networks. Network heterogeneities were implemented as network regions having increased connectivity as compared to the rest of the network. We varied the properties of the oscillatory drive as well as network connectivity. We observed that the disparity in spatiotemporal patterning of activity between the structurally enhanced region and rest of the network was highly dependent on the frequency and amplitude of the oscillatory drive as well as network connectivity, generally favoring bigger enhancement of activity for high-frequency oscillations and phase locking with moderate enhancement of activity for lower-frequency oscillations. Thus, these results indicate that the specific role of the observed oscillations may depend on their dynamical interactions with the heterogeneous network.