THETA-Rhythm Makes the World Go Round: Dissociative Effects of TMS Theta Versus Alpha Entrainment of Right pTPJ on Embodied Perspective Transformations.

Being able to imagine another person's experience and perspective of the world is a crucial human ability and recent reports suggest that humans "embody" another's viewpoint by mentally rotating their own body representation into the other's orientation. Our recent Magnetoencephalography (MEG) data further confirmed this notion of embodied perspective transformations and pinpointed the right posterior temporo-parietal junction (pTPJ) as the crucial hub in a distributed network oscillating at theta frequency (3-7 Hz). In a subsequent transcranial magnetic stimulation (TMS) experiment we interfered with right pTPJ processing and observed a modulation of the embodied aspects of perspective transformations. While these results corroborated the role of right pTPJ, the notion of theta oscillations being the crucial neural code remained a correlational observation based on our MEG data. In the current study we therefore set out to confirm the importance of theta oscillations directly by means of TMS entrainment. We compared entrainment of right pTPJ at 6 Hz vs. 10 Hz and confirmed that only 6 Hz entrainment facilitated embodied perspective transformations (at 160° angular disparity) while 10 Hz slowed it down. The reverse was true at low angular disparity (60° between egocentric and target perspective) where a perspective transformation was not strictly necessary. Our results further corroborate right pTPJ involvement in embodied perspective transformations and highlight theta oscillations as a crucial neural code.

Irregular Speech Rate Dissociates Auditory Cortical Entrainment, Evoked Responses, and Frontal Alpha.

The entrainment of slow rhythmic auditory cortical activity to the temporal regularities in speech is considered to be a central mechanism underlying auditory perception. Previous work has shown that entrainment is reduced when the quality of the acoustic input is degraded, but has also linked rhythmic activity at similar time scales to the encoding of temporal expectations. To understand these bottom-up and top-down contributions to rhythmic entrainment, we manipulated the temporal predictive structure of speech by parametrically altering the distribution of pauses between syllables or words, thereby rendering the local speech rate irregular while preserving intelligibility and the envelope fluctuations of the acoustic signal. Recording EEG activity in human participants, we found that this manipulation did not alter neural processes reflecting the encoding of individual sound transients, such as evoked potentials. However, the manipulation significantly reduced the fidelity of auditory delta (but not theta) band entrainment to the speech envelope. It also reduced left frontal alpha power and this alpha reduction was predictive of the reduced delta entrainment across participants. Our results show that rhythmic auditory entrainment in delta and theta bands reflect functionally distinct processes. Furthermore, they reveal that delta entrainment is under top-down control and likely reflects prefrontal processes that are sensitive to acoustical regularities rather than the bottom-up encoding of acoustic features. SIGNIFICANCE STATEMENT: The entrainment of rhythmic auditory cortical activity to the speech envelope is considered to be critical for hearing. Previous work has proposed divergent views in which entrainment reflects either early evoked responses related to sound encoding or high-level processes related to expectation or cognitive selection. Using a manipulation of speech rate, we dissociated auditory entrainment at different time scales. Specifically, our results suggest that delta entrainment is controlled by frontal alpha mechanisms and thus support the notion that rhythmic auditory cortical entrainment is shaped by top-down mechanisms.

Getting the beat: entrainment of brain activity by musical rhythm and pleasantness.

Rhythmic entrainment is an important component of emotion induction by music, but brain circuits recruited during spontaneous entrainment of attention by music and the influence of the subjective emotional feelings evoked by music remain still largely unresolved. In this study we used fMRI to test whether the metric structure of music entrains brain activity and how music pleasantness influences such entrainment. Participants listened to piano music while performing a speeded visuomotor detection task in which targets appeared time-locked to either strong or weak beats. Each musical piece was presented in both a consonant/pleasant and dissonant/unpleasant version. Consonant music facilitated target detection and targets presented synchronously with strong beats were detected faster. FMRI showed increased activation of bilateral caudate nucleus when responding on strong beats, whereas consonance enhanced activity in attentional networks. Meter and consonance selectively interacted in the caudate nucleus, with greater meter effects during dissonant than consonant music. These results reveal that the basal ganglia, involved both in emotion and rhythm processing, critically contribute to rhythmic entrainment of subcortical brain circuits by music.

Development of full-body rhythmic synchronization in middle childhood.

Rhythmic entrainment is a fundamental aspect of musical behavior, but the skills required to accurately synchronize movement to the beat seem to develop over many years. Motion capture studies of corporeal synchronization have shown immature abilities to lock in to the beat in children before age 5, and reliable synchronization ability in adults without musical training; yet there is a lack of data on full-body synchronization skills between early childhood and adulthood. To document typical rhythmic synchronization during middle childhood, we used a wireless motion capture device to measure period- and phase-locking of full body movement to rhythm and metronome stimuli in 6 to 11 year-old children in comparison with adult data. Results show a gradual improvement with age; however children's performance did not reach adult levels by age 12, suggesting that these skills continue to develop during adolescence. Our results suggest that in the absence of specific music training, full-body rhythmic entrainment skills improve gradually during middle childhood, and provide metrics for examining the continued maturation of these skills during adolescence.

An integrated review of music cognition and rhythmic stimuli in sensorimotor neurocognition and neurorehabilitation.

This work reviews the growing body of interdisciplinary research on music cognition, using biomechanical, kinesiological, clinical, psychosocial, and sociological methods. The review primarily examines the relationship between temporal elements in music and motor responses under varying contexts, with considerable relevance for clinical rehabilitation. After providing an overview of the terminology and approaches pertinent to theories of rhythm and meter from the musical-theoretical and cognitive fields, this review focuses on studies on the effects of rhythmic sensory stimulation on gait, rhythmic cues' effect on the motor system, reactions to rhythmic stimuli attempting to synchronize mobility (i.e., musical embodiment), and the application of rhythm for motor rehabilitation for individuals with Parkinson's disease, stroke, mild cognitive impairment, Alzheimer's disease, and other neurodegenerative or neurotraumatic diseases. This work ultimately bridges the gap between the musical-theoretical and cognitive science fields to facilitate innovative research in which each discipline informs the other.

The Cerebral Cortex and the Songs of Homer: When Neuroscience Meets History and Literature.

In this article we reconsider Homer's poetry in the light of modern achievements in neuroscience. This perspective offers some clues for examining specific patterns of brain functioning. Homer's epics, for instance, painted a synthetic picture of the human body, emphasizing some parts and neglecting others. This led to the formation of a body schema reminiscent of a homunculus, which we call the "Homeric homunculus." Both poems were largely the product of centuries of oral tradition, in which the prodigious memory of courtly rhapsodists was essential to the performance of the epics. The underlying cognitive functions required a close interplay of memory and language skills, supported by the musical and rhythmic cadence of Homeric verse.

Familiarity with music influences stride amplitude and variability during rhythmically-cued walking in individuals with Parkinson's disease.

BACKGROUND: Entrainment of walking with rhythmic auditory cues (e.g., metronome or music) induces gait improvements in people with Parkinson's disease (PD). However, evidence regarding acute impacts of music cues on parkinsonian gait is lacking. Findings in healthy adults imply that familiarity with music cues modulates stride amplitude and stride-to-stride variability; yet the role of familiarity with music on parkinsonian gait remains unknown. RESEARCH QUESTION: To investigate 1) whether familiar and unfamiliar music cueing differentially influences stride and arm swing amplitude and stride-to-stride variability, 2) how stride and arm swing amplitude and stride-to-stride variability are altered by enhanced familiarity with music by repeated listening and walking during rhythmically-cued walking in persons with PD. METHODS: Twenty individuals with idiopathic PD (mean age = 68.9 years, 7 females, H&Y stage 1-3) walked in time with familiar and unfamiliar music cues (Session 1). Participants then repeatedly listened and walked to the same familiar and unfamiliar music cues (Session 2). Spatiotemporal gait parameters in each 2-min trial were recorded with motion capture wearable sensors. RESULTS: In Session 1, gait velocity, stride length, arm swing peak velocity and range of motion, and stride-to-stride variability increased with both music cues compared with baseline; stride length, enjoyment and beat salience were greater in familiar than unfamiliar music cues. In Session 2, repeatedly listening and walking to unfamiliar music, but not familiar music, increased gait velocity, stride length, familiarity, enjoyment, and beat salience, and reduced stride-to-stride time variability. SIGNIFICANCE: Music cues in general induced acute improvements of stride and arm swing amplitude but not stride-to-stride variability, and enhanced familiarity with music improved stride amplitude and variability along with increased enjoyment and reduced cognitive demand in people with PD. Our findings aid in understanding the role of familiarity with music in alleviating gait disturbance and optimizing music-based interventions for PD.

Emotional content overrides spatial attention.

Spatial attention is our capacity to attend to or ignore particular regions of our spatial environment. However, some classes of stimuli may be able to override our efforts to ignore them. Here we assessed the relationship between involuntary attentional capture with emotional images and spatial attention at early stages of perceptual processing. Multiple scenes of unpleasant and neutral content were displayed in rapid serial visual presentation (RSVP) streams that elicited the steady-state visual evoked potential (SSVEP), a neural marker of selective attention at early visual areas. In a spatial cueing task, participants were cued to covertly attend to RSVP streams presented at 4 and 6 Hz presentation rates in the left and right visual hemifields. The task was to detect square targets occasionally displayed within the image streams, responding only to those appearing on the cued side. The RSVP streams were always neutral pictures in one visual hemifield but would unpredictably switch from neutral to aversive content in the other visual hemifield. We found that SSVEP amplitude was consistently modulated by a change in emotional valence of image streams, regardless of whether the change in content occurred in the attended or unattended spatial location, reflecting an automatic sensory amplification for affective stimuli. The present data provide further evidence in support that emotional images can attract visual processing resources independently of spatial attention allocation, and are consistent with sustained sensory facilitation of early visual areas through re-entrant feedback projections from higher-order cortical areas involved in the extraction of affective information.

Infant Movement Response to Auditory Rhythm.

Rhythmic entrainment occurs when an auditory rhythm drives an internal movement oscillator, thus providing a continuous time reference that improves temporal and spatial movement parameters. Entrainment processes and outcomes are well known for adults, but research is lacking for infants who might benefit from diagnosis and treatment of irregular rhythms within biological, sensorimotor, cognitive, and social domains. The present study used a combination of inertial measurement units and custom-made software to determine the amount, tempo, and regularity of movement in 28 infants aged 6-10 months while they were exposed to silence, an irregular rhythmic cue, or a regular rhythmic cue with tempo changes. We also assessed changes in the infants' movement parameters following a one-week rhythm training protocol. While results revealed no significant effect of auditory condition on amount or tempo of movement, infant movement was significantly more regular when infants were exposed to 120 bpm (beats per minute) than to an irregular rhythmic cue or a 10% faster rhythmic cue (132 bpm). Infants showed no notable changes in movement amount, tempo, or regularity following one week of training involving auditory and physical rhythm. Overall, infants seem to engage in spontaneous movements with or without auditory rhythm but may not show tempo sensitivity through their movements. Increased movement regularity suggests that 120 bpm may be a preferred tempo for infants, at which they are more likely to demonstrate well-timed movements that may reflect interval entrainment. Infants' auditory-motor systems appear not to respond to a 1-week rhythm training protocol.

Imagined Steps: Mental Simulation of Coordinated Rhythmic Movements Effects on Pro-sociality.

Rhythmically coordinating with a partner can increase pro-sociality, but pro-sociality does not appear to change in proportion to coordination success, or particular classes of coordination. Pro-social benefits may have more to do with simply coordinating in a social context than the details of the actual coordination (Cross et al., 2016). This begs the question, how stripped down can a coordination task be and still affect pro-sociality? Would it be sufficient simply to imagine coordinating with others? Imagining a social interaction can lead to many of the same effects as actual interaction (Crisp and Turner, 2009). We report the first experiments to explore whether imagined coordination affects pro-sociality similarly to actual coordination. Across two experiments and over 450 participants, mentally simulated coordination is shown to promote some, but not all, of the pro-social consequences of actual coordination. Imagined coordination significantly increased group cohesion and de-individuation, but did not consistently affect cooperation.

DRUM-PD: The use of a drum circle to improve the symptoms and signs of Parkinson's disease (PD).

BACKGROUND: Physical therapy can improve motor function in patients with PD. Music performance may be used to improve motor skills by rhythmic entrainment. Drumming has long been a part of traditional healing rituals worldwide, and is increasingly being utilized as a therapeutic strategy. METHODS: This pilot controlled prospective cohort trial assessed feasibility and effects of twice-weekly group West African drum circle classes for 6 weeks on PD patients' quality of life, symptoms, motor findings, cognition, and mood. Ten patients with PD were recruited into the drum circle group. Ten patients with PD were matched pairwise to each of the drum circle participants, and enrolled in a no-intervention control group. Both groups completed the PD-specific Parkinson Disease Questionnaire (PDQ)-39 quality of life assessment and the Geriatric Depression Scale (GDS), and underwent motor and cognitive assessments by a rater blinded to group at baseline, 6 weeks, and 12 weeks. RESULTS: Drummers had significantly improved PDQ-39 scores from baseline to 6 weeks (-5.8, p=0.042), whereas the control group's scores were unchanged. Walking performance was significantly faster at baseline for controls; after 6 weeks of drumming this difference was no longer significant, and remained non-significant at 12 weeks. The drummers trended (p=0.069) toward improvement in walking from baseline to 12 weeks. Other outcomes did not significantly change from baseline to 6 or 12 weeks. CONCLUSIONS: Drum circle classes significantly and reversibly improved quality of life in patients with PD. This pilot trial's findings merit larger controlled investigations comparing drumming classes to established interventions in PD, such as physical therapy.

Tracing the neural basis of auditory entrainment.

Neurons in the auditory cortex synchronize their responses to temporal regularities in sound input. This coupling or "entrainment" is thought to facilitate beat extraction and rhythm perception in temporally structured sounds, such as music. As a consequence of such entrainment, the auditory cortex responds to an omitted (silent) sound in a regular sequence. Although previous studies suggest that the auditory brainstem frequency-following response (FFR) exhibits some of the beat-related effects found in the cortex, it is unknown whether omissions of sounds evoke a brainstem response. We simultaneously recorded cortical and brainstem responses to isochronous and irregular sequences of consonant-vowel syllable /da/ that contained sporadic omissions. The auditory cortex responded strongly to omissions, but we found no evidence of evoked responses to omitted stimuli from the auditory brainstem. However, auditory brainstem responses in the isochronous sound sequence were more consistent across trials than in the irregular sequence. These results indicate that the auditory brainstem faithfully encodes short-term acoustic properties of a stimulus and is sensitive to sequence regularity, but does not entrain to isochronous sequences sufficiently to generate overt omission responses, even for sequences that evoke such responses in the cortex. These findings add to our understanding of the processing of sound regularities, which is an important aspect of human cognitive abilities like rhythm, music and speech perception.

Beat Keeping in a Sea Lion As Coupled Oscillation: Implications for Comparative Understanding of Human Rhythm.

Human capacity for entraining movement to external rhythms-i.e., beat keeping-is ubiquitous, but its evolutionary history and neural underpinnings remain a mystery. Recent findings of entrainment to simple and complex rhythms in non-human animals pave the way for a novel comparative approach to assess the origins and mechanisms of rhythmic behavior. The most reliable non-human beat keeper to date is a California sea lion, Ronan, who was trained to match head movements to isochronous repeating stimuli and showed spontaneous generalization of this ability to novel tempos and to the complex rhythms of music. Does Ronan's performance rely on the same neural mechanisms as human rhythmic behavior? In the current study, we presented Ronan with simple rhythmic stimuli at novel tempos. On some trials, we introduced "perturbations," altering either tempo or phase in the middle of a presentation. Ronan quickly adjusted her behavior following all perturbations, recovering her consistent phase and tempo relationships to the stimulus within a few beats. Ronan's performance was consistent with predictions of mathematical models describing coupled oscillation: a model relying solely on phase coupling strongly matched her behavior, and the model was further improved with the addition of period coupling. These findings are the clearest evidence yet for parity in human and non-human beat keeping and support the view that the human ability to perceive and move in time to rhythm may be rooted in broadly conserved neural mechanisms.

How Moving Together Brings Us Together: When Coordinated Rhythmic Movement Affects Cooperation.

Although it is well established that rhythmically coordinating with a social partner can increase cooperation, it is as yet unclear when and why intentional coordination has such effects. We distinguish three dimensions along which explanations might vary. First, pro-social effects might require in-phase synchrony or simply coordination. Second, the effects of rhythmic movements on cooperation might be direct or mediated by an intervening variable. Third, the pro-social effects might occur in proportion to the quality of the coordination, or occur once some threshold amount of coordination has occurred. We report an experiment and two follow-ups which sought to identify which classes of models are required to account for the positive effects of coordinated rhythmic movement on cooperation. Across the studies, we found evidence (1) that coordination, and not just synchrony, can have pro-social consequences (so long as the social nature of the task is perceived), (2) that the effects of intentional coordination are direct, not mediated, and (3) that the degree of the coordination did not predict the degree of cooperation. The fact of inter-personal coordination (moving together in time and in a social context) is all that's required for pro-social effects. We suggest that future research should use the kind of carefully controllable experimental task used here to continue to develop explanations for when and why coordination affects pro-social behaviors.

Unpredicted Pitch Modulates Beta Oscillatory Power during Rhythmic Entrainment to a Tone Sequence.

Extracting temporal regularities in external stimuli in order to predict upcoming events is an essential aspect of perception. Fluctuations in induced power of beta band (15-25 Hz) oscillations in auditory cortex are involved in predictive timing during rhythmic entrainment, but whether such fluctuations are affected by prediction in the spectral (frequency/pitch) domain remains unclear. We tested whether unpredicted (i.e., unexpected) pitches in a rhythmic tone sequence modulate beta band activity by recording EEG while participants passively listened to isochronous auditory oddball sequences with occasional unpredicted deviant pitches at two different presentation rates. The results showed that the power in low-beta (15-20 Hz) was larger around 200-300 ms following deviant tones compared to standard tones, and this effect was larger when the deviant tones were less predicted. Our results suggest that the induced beta power activities in auditory cortex are consistent with a role in sensory prediction of both "when" (timing) upcoming sounds will occur as well as the prediction precision error of "what" (spectral content in this case). We suggest, further, that both timing and content predictions may co-modulate beta oscillations via attention. These findings extend earlier work on neural oscillations by investigating the functional significance of beta oscillations for sensory prediction. The findings help elucidate the functional significance of beta oscillations in perception.

Motor system evolution and the emergence of high cognitive functions.

In human and nonhuman primates, the cortical motor system comprises a collection of brain areas primarily related to motor control. Existing evidence suggests that no other mammalian group has the number, extension, and complexity of motor-related areas observed in the frontal lobe of primates. Such diversity is probably related to the wide behavioral flexibility that primates display. Indeed, recent comparative anatomical, psychophysical, and neurophysiological studies suggest that the evolution of the motor cortical areas closely correlates with the emergence of high cognitive abilities. Advances in understanding the cortical motor system have shown that these areas are also related to functions previously linked to higher-order associative areas. In addition, experimental observations have shown that the classical distinction between perceptual and motor functions is not strictly followed across cortical areas. In this paper, we review evidence suggesting that evolution of the motor system had a role in the shaping of different cognitive functions in primates. We argue that the increase in the complexity of the motor system has contributed to the emergence of new abilities observed in human and nonhuman primates, including the recognition and imitation of the actions of others, speech perception and production, and the execution and appreciation of the rhythmic structure of music.

Recovery after aerobic exercise is manipulated by tempo change in a rhythmic sound pattern, as indicated by autonomic reaction on heart functioning.

Physical prowess is associated with rapid recovery from exhaustion. Here we examined whether recovery from aerobic exercise could be manipulated with a rhythmic sound pattern that either decreased or increased in tempo. Six men and six women exercised repeatedly for six minutes on a cycle ergometer at 60 percent of their individual maximal oxygen consumption, and then relaxed for six minutes while listening to one of two sound pattern conditions, which seemed to infinitely either decrease or increase in tempo, during which heart and breathing activity was measured. Participants exhibited more high-frequent heart rate variability when listening to decreasing tempo than when listening to increasing tempo, accompanied by a non-significant trend towards lower heart rate. The results show that neuropsychological entrainment to a sound pattern may directly affect the autonomic nervous system, which in turn may facilitate physiological recovery after exercise. Applications using rhythmic entrainment to aid physical recovery are discussed.

Situational influences on rhythmicity in speech, music, and their interaction.

Brain processes underlying the production and perception of rhythm indicate considerable flexibility in how physical signals are interpreted. This paper explores how that flexibility might play out in rhythmicity in speech and music. There is much in common across the two domains, but there are also significant differences. Interpretations are explored that reconcile some of the differences, particularly with respect to how functional properties modify the rhythmicity of speech, within limits imposed by its structural constraints. Functional and structural differences mean that music is typically more rhythmic than speech, and that speech will be more rhythmic when the emotions are more strongly engaged, or intended to be engaged. The influence of rhythmicity on attention is acknowledged, and it is suggested that local increases in rhythmicity occur at times when attention is required to coordinate joint action, whether in talking or music-making. Evidence is presented which suggests that while these short phases of heightened rhythmical behaviour are crucial to the success of transitions in communicative interaction, their modality is immaterial: they all function to enhance precise temporal prediction and hence tightly coordinated joint action.

Are non-human primates capable of rhythmic entrainment? Evidence for the gradual audiomotor evolution hypothesis.

We propose a decomposition of the neurocognitive mechanisms that might underlie interval-based timing and rhythmic entrainment. Next to reviewing the concepts central to the definition of rhythmic entrainment, we discuss recent studies that suggest rhythmic entrainment to be specific to humans and a selected group of bird species, but, surprisingly, is not obvious in non-human primates. On the basis of these studies we propose the gradual audiomotor evolution hypothesis that suggests that humans fully share interval-based timing with other primates, but only partially share the ability of rhythmic entrainment (or beat-based timing). This hypothesis accommodates the fact that non-human primates (i.e., macaques) performance is comparable to humans in single interval tasks (such as interval reproduction, categorization, and interception), but show differences in multiple interval tasks (such as rhythmic entrainment, synchronization, and continuation). Furthermore, it is in line with the observation that macaques can, apparently, synchronize in the visual domain, but show less sensitivity in the auditory domain. And finally, while macaques are sensitive to interval-based timing and rhythmic grouping, the absence of a strong coupling between the auditory and motor system of non-human primates might be the reason why macaques cannot rhythmically entrain in the way humans do.

How musical training affects cognitive development: rhythm, reward and other modulating variables.

Musical training has recently gained additional interest in education as increasing neuroscientific research demonstrates its positive effects on brain development. Neuroimaging revealed plastic changes in the brains of adult musicians but it is still unclear to what extent they are the product of intensive music training rather than of other factors, such as preexisting biological markers of musicality. In this review, we synthesize a large body of studies demonstrating that benefits of musical training extend beyond the skills it directly aims to train and last well into adulthood. For example, children who undergo musical training have better verbal memory, second language pronunciation accuracy, reading ability and executive functions. Learning to play an instrument as a child may even predict academic performance and IQ in young adulthood. The degree of observed structural and functional adaptation in the brain correlates with intensity and duration of practice. Importantly, the effects on cognitive development depend on the timing of musical initiation due to sensitive periods during development, as well as on several other modulating variables. Notably, we point to motivation, reward and social context of musical education, which are important yet neglected factors affecting the long-term benefits of musical training. Further, we introduce the notion of rhythmic entrainment and suggest that it may represent a mechanism supporting learning and development of executive functions. It also hones temporal processing and orienting of attention in time that may underlie enhancements observed in reading and verbal memory. We conclude that musical training uniquely engenders near and far transfer effects, preparing a foundation for a range of skills, and thus fostering cognitive development.