Distinct cortico-muscular coupling between step and stance leg during reactive stepping responses.

Balance recovery often relies on successful stepping responses, which presumably require precise and rapid interactions between the cerebral cortex and the leg muscles. Yet, little is known about how cortico-muscular coupling (CMC) supports the execution of reactive stepping. We conducted an exploratory analysis investigating time-dependent CMC with specific leg muscles in a reactive stepping task. We analyzed high density EEG, EMG, and kinematics of 18 healthy young participants while exposing them to balance perturbations at different intensities, in the forward and backward directions. Participants were instructed to maintain their feet in place, unless stepping was unavoidable. Muscle-specific Granger causality analysis was conducted on single step- and stance-leg muscles over 13 EEG electrodes with a midfrontal scalp distribution. Time-frequency Granger causality analysis was used to identify CMC from cortex to muscles around perturbation onset, foot-off and foot strike events. We hypothesized that CMC would increase compared to baseline. In addition, we expected to observe different CMC between step and stance leg because of their functional role during the step response. In particular, we expected that CMC would be most evident for the agonist muscles while stepping, and that CMC would precede upregulation in EMG activity in these muscles. We observed distinct Granger gain dynamics over theta, alpha, beta, and low/high-gamma frequencies during the reactive balance response for all leg muscles in each step direction. Interestingly, between-leg differences in Granger gain were almost exclusively observed following the divergence of EMG activity. Our results demonstrate cortical involvement in the reactive balance response and provide insights into its temporal and spectral characteristics. Overall, our findings suggest that higher levels of CMC do not facilitate leg-specific EMG activity. Our work is relevant for clinical populations with impaired balance control, where CMC analysis may elucidate the underlying pathophysiological mechanisms.

Midfrontal theta dynamics index the monitoring of postural stability.

Stepping is a common strategy to recover postural stability and maintain upright balance. Postural perturbations have been linked to neuroelectrical markers such as the N1 potential and theta frequency dynamics. Here, we investigated the role of cortical midfrontal theta dynamics of balance monitoring, driven by balance perturbations at different initial standing postures. We recorded electroencephalography, electromyography, and motion tracking of human participants while they stood on a platform that delivered a range of forward and backward whole-body balance perturbations. The participants' postural threat was manipulated prior to the balance perturbation by instructing them to lean forward or backward while keeping their feet-in-place in response to the perturbation. We hypothesized that midfrontal theta dynamics index the engagement of a behavioral monitoring system and, therefore, that perturbation-induced theta power would be modulated by the initial leaning posture and perturbation intensity. Targeted spatial filtering in combination with mixed-effects modeling confirmed our hypothesis and revealed distinct modulations of theta power according to postural threat. Our results provide novel evidence that midfrontal theta dynamics subserve action monitoring of human postural balance. Understanding of cortical mechanisms of balance control is crucial for studying balance impairments related to aging and neurological conditions (e.g. stroke).

Cortical midfrontal theta dynamics following foot strike may index response adaptation during reactive stepping.

Reactive balance recovery often requires stepping responses to regain postural stability following a sudden change in posture. The monitoring of postural stability has been linked to neuroelectrical markers such as the N1 potential and midfrontal theta frequency dynamics. Here, we investigated the role of cortical midfrontal theta dynamics during balance monitoring following foot landing of a reactive stepping response to recover from whole-body balance perturbations. We hypothesized that midfrontal theta dynamics reflect the engagement of a behavioral monitoring system, and therefore that theta would increase time-locked to the moment of foot strike after a stepping response, coinciding with a re-assessment of postural balance to determine if an additional step is necessary. We recorded high-density EEG and kinematic data of 15 healthy young participants while they stood on a platform that delivered multi-directional balance perturbations. Participants were instructed to recover balance with a single step utilizing either their left or right leg (in separate blocks). We used targeted spatial filtering (generalized eigen decomposition) in combination with time-frequency analysis of the EEG data to investigate whether theta dynamics increase following foot strike event. In line with our hypothesis, the results indicate that the foot strike event elicits a midfrontal theta power increase, though only for backward stepping. Counter to our expectations, however, this theta power increase was positively correlated with the margin of stability at foot strike, suggesting a different role of foot strike related theta from monitoring stability. Post-hoc analysis suggests that midfrontal theta dynamics following foot landing may instead facilitate adaptation of stability margins at subsequent stepping responses. We speculate that increase of theta power following foot strikes was not related to stability monitoring but instead may indicate cortical dynamics related to performance monitoring of the balance response.

Synchronization between Keyboard Typing and Neural Oscillations.

Rhythmic neural activity synchronizes with certain rhythmic behaviors, such as breathing, sniffing, saccades, and speech. The extent to which neural oscillations synchronize with higher-level and more complex behaviors is largely unknown. Here, we investigated electrophysiological synchronization with keyboard typing, which is an omnipresent behavior daily engaged by an uncountably large number of people. Keyboard typing is rhythmic, with frequency characteristics roughly the same as neural oscillatory dynamics associated with cognitive control, notably through midfrontal theta (4-7 Hz) oscillations. We tested the hypothesis that synchronization occurs between typing and midfrontal theta and breaks down when errors are committed. Thirty healthy participants typed words and sentences on a keyboard without visual feedback, while EEG was recorded. Typing rhythmicity was investigated by interkeystroke interval analyses and by a kernel density estimation method. We used a multivariate spatial filtering technique to investigate frequency-specific synchronization between typing and neuronal oscillations. Our results demonstrate theta rhythmicity in typing (around 6.5 Hz) through the two different behavioral analyses. Synchronization between typing and neuronal oscillations occurred at frequencies ranging from 4 to 15 Hz, but to a larger extent for lower frequencies. However, peak synchronization frequency was idiosyncratic across participants, therefore not specific to theta nor to midfrontal regions, and correlated somewhat with peak typing frequency. Errors and trials associated with stronger cognitive control were not associated with changes in synchronization at any frequency. As a whole, this study shows that brain-behavior synchronization does occur during keyboard typing but is not specific to midfrontal theta.

Cortical responses to whole-body balance perturbations index perturbation magnitude and predict reactive stepping behavior.

The goal of this study was to determine whether the cortical responses elicited by whole-body balance perturbations were similar to established cortical markers of action monitoring. Postural changes imposed by balance perturbations elicit a robust negative potential (N1) and a brisk increase of theta activity in the electroencephalogram recorded over midfrontal scalp areas. Because action monitoring is a cognitive function proposed to detect errors and initiate corrective adjustments, we hypothesized that the possible cortical markers of action monitoring during balance control (N1 potential and theta rhythm) scale with perturbation intensity and the eventual execution of reactive stepping responses (as opposed to feet-in-place responses). We recorded high-density electroencephalogram from eleven young individuals, who participated in an experimental balance assessment. The participants were asked to recover balance following anteroposterior translations of the support surface at various intensities, while attempting to maintain both feet in place. We estimated source-resolved cortical activity using independent component analysis. Combining time-frequency decomposition and group-level general linear modeling of single-trial responses, we found a significant relation of the interaction between perturbation intensity and stepping responses with multiple cortical features from the midfrontal cortex, including the N1 potential, and theta, alpha, and beta rhythms. Our findings suggest that the cortical responses to balance perturbations index the magnitude of a deviation from a stable postural state to predict the need for reactive stepping responses. We propose that the cortical control of balance may involve cognitive control mechanisms (i.e., action monitoring) that facilitate postural adjustments to maintain postural stability.

The effects of environmental enrichment and age-related differences on inhibitory avoidance in zebrafish (Danio rerio Hamilton).

The inhibitory avoidance paradigm allows the study of mechanisms underlying learning and memory formation in zebrafish (Danio rerio Hamilton). For zebrafish, the physiology and behavior associated with this paradigm are as yet poorly understood. We therefore assessed the effects of environmental enrichment and fish age on inhibitory avoidance learning. Fish raised in an environmentally enriched tank showed decreased anxiety-like behavior and increased exploration. Enrichment greatly reduced inhibitory avoidance in 6-month (6M)- and 12-month (12 M)-old fish. Following inhibitory avoidance, telencephalic mRNA levels of proliferating cell nuclear antigen (pcna), neurogenic differentiation (neurod), cocaine- and amphetamine-regulated transcript 4 (cart4), and cannabinoid receptor 1 (cnr1) were lower in enriched-housed fish, while the ratios of mineralocorticoid receptor (nr3c2)/glucocorticoid receptor α [nr3c1(α)] and glucocorticoid receptor ß [nr3c1(ß)]/glucocorticoid receptor α [nr3c1(α)] were higher. This was observed for 6M-old fish only, not for 24-month (24 M) old fish. Instead, 24 M-old fish showed delayed inhibitory avoidance, no effects of enrichment, and reduced expression of neuroplasticity genes. Overall, our data show strong differences in inhibitory avoidance behavior between zebrafish of different ages and a clear reduction in avoidance behavior following housing under environmental enrichment.