Differential motor system entrainment to auditory and visual rhythms.

Perception of, and synchronization to, auditory rhythms is known to be more accurate than with flashing visual rhythms. The motor system is known to play a role in the processing of timing information for auditory rhythm perception, but it is unclear if the motor system plays the same role for visual rhythm perception. One demonstrated component of auditory rhythm perception is neural entrainment at the frequency of the auditory rhythm. In this study, we use EEG to measure the entrainment of both auditory and visual rhythms from the motor cortex while subjects either tapped in synchrony with or passively attended to the presented rhythms. To isolate activity from motor cortex, we used independent component analysis to first separate out neural sources, then selected components using a combination of component topography, dipole location, mu activation, and beta modulation. This process took advantage of the fact that tapping activity results in reduced mu power, and characteristic beta modulation, which helped select motor components. Our findings suggest neural entrainment in motor components was stronger for visual rhythms than auditory rhythms and strongest during the tapping conditions for both modalities. We also find mu power increased in response to both auditory and visual rhythms. These findings indicate that the generally greater rhythm perception capabilities of the auditory system over the visual system may not depend entirely on neural entrainment in the motor system, but rather how the motor system is able to use the timing information made available to it. NEW & NOTEWORTHY We investigated neural entrainment in the motor system for both auditory and visual isochronous rhythms using electroencephalogram. Counter to expectations, our findings suggest stronger entrainment for visual rhythms than for auditory rhythms. Motor system activity was isolated with a novel procedure using independent component analysis as a means of blind source separation, along with known markers of mu activity from the motor system to identify motor components.

A systematic review and meta-analysis of the efficacy of intermittent theta burst stimulation (iTBS) on cognitive enhancement.

Intermittent theta-burst stimulation (iTBS) has been used to focally regulate excitability of neural cortex over the past decade - however there is little consensus on the generalizability of effects reported in individual studies. Many studies use small sample sizes (N < 30), and there is a considerable amount of methodological heterogeneity in application of the stimulation itself. This systematic meta-analysis aims to consolidate the extant literature and determine if up-regulatory theta-burst stimulation reliably enhances cognition through measurable behavior. Results show that iTBS - when compared to suitable control conditions - may enhance cognition when outlier studies are removed, but also that there is a significant amount of heterogeneity across studies. Significant contributors to between-study heterogeneity include location of stimulation and method of navigation to the stimulation site. Surprisingly, the type of cognitive domain investigated was not a significant contributor of heterogeneity. The findings of this meta-analysis demonstrate that standardization of iTBS is urgent and necessary to determine if neuroenhancement of particular cognitive faculties are reliable and robust, and measurable through observable behavior.

Cortical mu rhythms during action and passive music listening.

Brain systems supporting body movement are active during music listening in the absence of overt movement. This covert motor activity is not well understood, but some theories propose a role in auditory timing prediction facilitated by motor simulation. One question is how music-related covert motor activity relates to motor activity during overt movement. We address this question using scalp electroencephalogram by measuring mu rhythms-cortical field phenomena associated with the somatomotor system that appear over sensorimotor cortex. Lateralized mu enhancement over hand sensorimotor cortex during/just before foot movement in foot versus hand movement paradigms is thought to reflect hand movement inhibition during current/prospective movement of another effector. Behavior of mu during music listening with movement suppressed has yet to be determined. We recorded 32-channel EEG (n = 17) during silence without movement, overt movement (foot/hand), and music listening without movement. Using an independent component analysis-based source equivalent dipole clustering technique, we identified three mu-related clusters, localized to left primary motor and right and midline premotor cortices. Right foot tapping was accompanied by mu enhancement in the left lateral source cluster, replicating previous work. Music listening was accompanied by similar mu enhancement in the left, as well as midline, clusters. We are the first, to our knowledge, to report, and also to source-resolve, music-related mu modulation in the absence of overt movements. Covert music-related motor activity has been shown to play a role in beat perception (Ross JM, Iversen JR, Balasubramaniam R. Neurocase 22: 558-565, 2016). Our current results show enhancement in somatotopically organized mu, supporting overt motor inhibition during beat perception.NEW & NOTEWORTHY We are the first to report music-related mu enhancement in the absence of overt movements and the first to source-resolve mu activity during music listening. We suggest that music-related mu modulation reflects overt motor inhibition during passive music listening. This work is relevant for the development of theories relating to the involvement of covert motor system activity for predictive beat perception.

Modality-specific frequency band activity during neural entrainment to auditory and visual rhythms.

Rhythm perception depends on the ability to predict the onset of rhythmic events. Previous studies indicate beta band modulation is involved in predicting the onset of auditory rhythmic events (Fujioka et al., 2009, 2012; Snyder & Large, 2005). We sought to determine if similar processes are recruited for prediction of visual rhythms by investigating whether beta band activity plays a role in a modality-dependent manner for rhythm perception. We looked at electroencephalography time-frequency neural correlates of prediction using an omission paradigm with auditory and visual rhythms. By using omissions, we can separate out predictive timing activity from stimulus-driven activity. We hypothesized that there would be modality-independent markers of rhythm prediction in induced beta band oscillatory activity, and our results support this hypothesis. We find induced and evoked predictive timing in both auditory and visual modalities. Additionally, we performed an exploratory-independent components-based spatial clustering analysis, and describe all resulting clusters. This analysis reveals that there may be overlapping networks of predictive beta activity based on common activation in the parietal and right frontal regions, auditory-specific predictive beta in bilateral sensorimotor regions, and visually specific predictive beta in midline central, and bilateral temporal/parietal regions. This analysis also shows evoked predictive beta activity in the left sensorimotor region specific to auditory rhythms and implicates modality-dependent networks for auditory and visual rhythm perception.

Motor and Predictive Processes in Auditory Beat and Rhythm Perception.

In this article, we review recent advances in research on rhythm and musical beat perception, focusing on the role of predictive processes in auditory motor interactions. We suggest that experimental evidence of the motor system's role in beat perception, including in passive listening, may be explained by the generation and maintenance of internal predictive models, concordant with the Active Inference framework of sensory processing. We highlight two complementary hypotheses for the neural underpinnings of rhythm perception: The Action Simulation for Auditory Prediction hypothesis (Patel and Iversen, 2014) and the Gradual Audiomotor Evolution hypothesis (Merchant and Honing, 2014) and review recent experimental progress supporting each of these hypotheses. While initial formulations of ASAP and GAE explain different aspects of beat-based timing-the involvement of motor structures in the absence of movement, and physical entrainment to an auditory beat respectively-we suggest that work under both hypotheses provide converging evidence toward understanding the predictive role of the motor system in the perception of rhythm, and the specific neural mechanisms involved. We discuss future experimental work necessary to further evaluate the causal neural mechanisms underlying beat and rhythm perception.

Sensorimotor Synchronization With Auditory and Visual Modalities: Behavioral and Neural Differences.

It has long been known that the auditory system is better suited to guide temporally precise behaviors like sensorimotor synchronization (SMS) than the visual system. Although this phenomenon has been studied for many years, the underlying neural and computational mechanisms remain unclear. Growing consensus suggests the existence of multiple, interacting, context-dependent systems, and that reduced precision in visuo-motor timing might be due to the way experimental tasks have been conceived. Indeed, the appropriateness of the stimulus for a given task greatly influences timing performance. In this review, we examine timing differences for sensorimotor synchronization and error correction with auditory and visual sequences, to inspect the underlying neural mechanisms that contribute to modality differences in timing. The disparity between auditory and visual timing likely relates to differences in the processing specialization between auditory and visual modalities (temporal vs. spatial). We propose this difference could offer potential explanation for the differing temporal abilities between modalities. We also offer suggestions as to how these sensory systems interface with motor and timing systems.

Neural responses to perturbations in visual and auditory metronomes during sensorimotor synchronization.

Tapping in synchrony to an isochronous rhythm involves several key functions of the sensorimotor system including timing, prediction and error correction. While auditory sensorimotor synchronization (SMS) has been well studied, much less is known about mechanisms involved in visual SMS. By comparing error correction in auditory and visual SMS, it can be determined if the neural mechanisms for detection and correction of synchronization errors are generalized or domain specific. To study this problem, we measured EEG while subjects tapped in synchrony to separate visual and auditory metronomes that both contained small temporal perturbations to induce errors. The metronomes had inter-onset intervals of 600 ms and the perturbations where of 4 kinds: ± 66 ms to induce period corrections, and ± 16 ms to induce phase corrections. We hypothesize that given the less precise nature of visual SMS, error correction to perturbed visual flashing rhythms will be more gradual than with the equivalent auditory perturbations. Additionally, we expect this more gradual error correction will be reflected in the visual evoked potentials. Our findings indicate that the visual system is only capable of more gradual phase corrections to even the larger induced errors. This is opposed to the swifter period correction of the auditory system to large induced errors. EEG data found the peak N1 auditory evoked potential is modulated by the size and direction of an induced error in line with previous research, while the P1 visual evoked potential was only effected by the large late-coming perturbations resulting in reduced peak latency. Looking at the error response EEG data, an Error Related Negativity (ERN) and related Error Positivity (pE) was found only in the auditory + 66 condition, while no ERN or pE were found in any of the visual perturbation conditions. In addition to the ERPs, we performed a dipole source localization and clustering analysis indicating that the anterior cingulate was active in the error detection of the perturbed stimulus for both auditory and visual conditions in addition to being involved in producing the ERN and pE induced by the auditory + 66 perturbation. Taken together, these results confirm that the visual system is less developed for synchronizing and error correction with flashing rhythms by its more gradual error correction. The reduced latency of the P1 to the visual + 66 suggests that the visual system can detect these errors, but that detection does not translate into any meaningful improvement in error correction. This indicates that the visual system is not as tightly coupled to the motor system as the auditory system is for SMS, suggesting the mechanisms of SMS are not completely domain general.