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Alpha and beta oscillations have been assessed thoroughly during walking due to their potential role as proxies of the corticoreticulospinal tract (CReST) and corticospinal tract (CST), respectively. Given that damage to a descending tract after stroke can cause walking deficits, detailed knowledge of how these oscillations mechanistically contribute to walking could be utilized in strategies for post-stroke locomotor recovery. In this review, the goal was to summarize, synthesize, and discuss the existing evidence on the potential differential role of these oscillations on the motor descending drive, the effect of transcranial alternate current stimulation (tACS) on neurotypical and post-stroke walking, and to discuss remaining gaps in knowledge, future directions, and methodological considerations. Electrophysiological studies of corticomuscular, intermuscular, and intramuscular coherence during walking clearly demonstrate that beta oscillations are predominantly present in the dorsiflexors during the swing phase and may be absent post-stroke. The role of alpha oscillations, however, has not been pinpointed as clearly. We concluded that both animal and human studies should focus on the electrophysiological characterization of alpha oscillations and their potential role to the CReST. Another approach in elucidating the role of these oscillations is to modulate them and then quantify the impact on walking behavior. This is possible through tACS, whose beneficial effect on walking behavior (including boosting of beta oscillations in intramuscular coherence) has been recently demonstrated in both neurotypical adults and stroke patients. However, these studies still do not allow for specific roles of alpha and beta oscillations to be delineated because the tACS frequency used was much lower (i.e., individualized calculated gait frequency was used). Thus, we identify a main gap in the literature, which is tACS studies actually stimulating at alpha and beta frequencies during walking. Overall, we conclude that for beta oscillations there is a clear connection to descending drive in the corticospinal tract. The precise relationship between alpha oscillations and CReST remains elusive due to the gaps in the literature identified here. However, better understanding the role of alpha (and beta) oscillations in the motor control of walking can be used to progress and develop rehabilitation strategies for promoting locomotor recovery.

Brain oscillations emerge during sensory and cognitive processes and have been classified into different frequency bands. Yet, even within the same frequency band and between nearby brain locations, the exact frequencies of brain oscillations can differ. These frequency differences (detuning) have been largely ignored and play little role in current functional theories of brain oscillations. This contrasts with the crucial role that detuning plays in synchronization theory, as originally derived in physical systems. Here, we propose that detuning is equally important to understand synchronization in biological systems. Detuning is a critical control parameter in synchronization, which is not only important in shaping phase-locking, but also in establishing preferred phase relations between oscillators. We review recent evidence that frequency differences between brain locations are ubiquitous and essential in shaping temporal neural coordination. With the rise of powerful experimental techniques to probe brain oscillations, the contributions of exact frequency and detuning across neural circuits will become increasingly clear and will play a key part in developing a new understanding of the role of oscillations in brain function.

It is well-established that both volume conduction and the choice of recording reference (montage) affect the correlation measures obtained from scalp EEG, both in the time and frequency domains. As a result, a number of correlation measures have been proposed aiming to reduce these effects. In our previous work, we have showed that scalp-EEG based functional brain networks in patients with epilepsy exhibit clear periodic patterns at different time scales and that these patterns are strongly correlated to seizure onset, particularly at shorter time scales (around 3 and 5 h), which has important clinical implications. In the present work, we use the same long-duration clinical scalp EEG data (multiple days) to investigate the extent to which the aforementioned results are affected by the choice of reference choice and correlation measure, by considering several widely used montages as well as correlation metrics that are differentially sensitive to the effects of volume conduction. Specifically, we compare two standard and commonly used linear correlation measures, cross-correlation in the time domain, and coherence in the frequency domain, with measures that account for zero-lag correlations: corrected cross-correlation, imaginary coherence, phase lag index, and weighted phase lag index. We show that the graphs constructed with corrected cross-correlation and WPLI are more stable across different choices of reference. Also, we demonstrate that all the examined correlation measures revealed similar periodic patterns in the obtained graph measures when the bipolar and common reference (Cz) montage were used. This includes circadian-related periodicities (e.g., a clear increase in connectivity during sleep periods as compared to awake periods), as well as periodicities at shorter time scales (around 3 and 5 h). On the other hand, these results were affected to a large degree when the average reference montage was used in combination with standard cross-correlation, coherence, imaginary coherence, and PLI, which is likely due to the low number of electrodes and inadequate electrode coverage of the scalp. Finally, we demonstrate that the correlation between seizure onset and the brain network periodicities is preserved when corrected cross-correlation and WPLI were used for all the examined montages. This suggests that, even in the standard clinical setting of EEG recording in epilepsy where only a limited number of scalp EEG measurements are available, graph-theoretic quantification of periodic patterns using appropriate montage, and correlation measures corrected for volume conduction provides useful insights into seizure onset.

Alpha and beta oscillations have been assessed thoroughly during walking due to their potential role as proxies of the corticoreticulospinal tract (CReST) and corticospinal tract (CST), respectively. Given that damage to a descending tract after stroke can cause walking deficits, detailed knowledge of how these oscillations mechanistically contribute to walking could be utilized in strategies for post-stroke locomotor recovery. In this review, the goal was to summarize, synthesize, and discuss the existing evidence on the potential differential role of these oscillations on the motor descending drive, the effect of transcranial alternate current stimulation (tACS) on neurotypical and post-stroke walking, and to discuss remaining gaps in knowledge, future directions, and methodological considerations. Electrophysiological studies of corticomuscular, intermuscular, and intramuscular coherence during walking clearly demonstrate that beta oscillations are predominantly present in the dorsiflexors during the swing phase and may be absent poststroke. The role of alpha oscillations, however, has not been pinpointed as clearly. We concluded that both animal and human studies should focus on the electrophysiological characterization of alpha oscillations and their potential role to the CReST. Another approach in elucidating the role of these oscillations is to modulate them and then quantify the impact on walking behavior. This is possible through tACS, whose beneficial effect on walking behavior (including boosting of beta oscillations in intramuscular coherence) has been recently demonstrated in both neurotypical adults and stroke patients. However, these studies still do not allow for specific roles of alpha and beta oscillations to be delineated because the tACS frequency used was much lower (i.e., individualized calculated gait frequency was used). Thus, we identify a main gap in the literature, which is tACS studies actually stimulating at alpha and beta frequencies during walking. Overall, we conclude that for beta oscillations there is a clear connection to descending drive in the corticospinal tract. The precise relationship between alpha oscillations and CReST remains elusive due to the gaps in the literature identified here. However, better understanding the role of alpha (and beta) oscillations in the motor control of walking can be used to progress and develop rehabilitation strategies for promoting locomotor recovery. [ABSTRACT FROM AUTHOR]

Brain oscillations emerge during sensory and cognitive processes and have been classified into different frequency bands. Yet, even within the same frequency band and between nearby brain locations, the exact frequencies of brain oscillations can differ. These frequency differences (detuning) have been largely ignored and play little role in current functional theories of brain oscillations. This contrasts with the crucial role that detuning plays in synchronization theory, as originally derived in physical systems. Here, we propose that detuning is equally important to understand synchronization in biological systems. Detuning is a critical control parameter in synchronization, which is not only important in shaping phase-locking, but also in establishing preferred phase relations between oscillators. We review recent evidence that frequency differences between brain locations are ubiquitous and essential in shaping temporal neural coordination. With the rise of powerful experimental techniques to probe brain oscillations, the contributions of exact frequency and detuning across neural circuits will become increasingly clear and will play a key part in developing a new understanding of the role of oscillations in brain function. [ABSTRACT FROM AUTHOR]

It is well-established that both volume conduction and the choice of recording reference (montage) affect the correlation measures obtained from scalp EEG, both in the time and frequency domains. As a result, a number of correlation measures have been proposed aiming to reduce these effects. In our previous work, we have showed that scalp-EEG based functional brain networks in patients with epilepsy exhibit clear periodic patterns at different time scales and that these patterns are strongly correlated to seizure onset, particularly at shorter time scales (around 3 and 5 h), which has important clinical implications. In the present work, we use the same long-duration clinical scalp EEG data (multiple days) to investigate the extent to which the aforementioned results are affected by the choice of reference choice and correlation measure, by considering several widely used montages as well as correlation metrics that are differentially sensitive to the effects of volume conduction. Specifically, we compare two standard and commonly used linear correlation measures, cross-correlation in the time domain, and coherence in the frequency domain, with measures that account for zero-lag correlations: corrected cross-correlation, imaginary coherence, phase lag index, and weighted phase lag index. We show that the graphs constructed with corrected cross-correlation and WPLI are more stable across different choices of reference. Also, we demonstrate that all the examined correlation measures revealed similar periodic patterns in the obtained graph measures when the bipolar and common reference (Cz) montage were used. This includes circadian-related periodicities (e.g., a clear increase in connectivity during sleep periods as compared to awake periods), as well as periodicities at shorter time scales (around 3 and 5 h). On the other hand, these results were affected to a large degree when the average reference montage was used in combination with standard cross-correlation, coherence, imaginary coherence, and PLI, which is likely due to the low number of electrodes and inadequate electrode coverage of the scalp. Finally, we demonstrate that the correlation between seizure onset and the brain network periodicities is preserved when corrected cross-correlation and WPLI were used for all the examined montages. This suggests that, even in the standard clinical setting of EEG recording in epilepsy where only a limited number of scalp EEG measurements are available, graph-theoretic quantification of periodic patterns using appropriate montage, and correlation measures corrected for volume conduction provides useful insights into seizure onset. [ABSTRACT FROM AUTHOR]