The influence of external and internal motor processes on human auditory rhythm perception

Musical rhythm is composed of organized temporal patterns, and the processes underlying rhythm perception are found to engage both auditory and motor systems. Despite behavioral and neuroscience evidence converging to this audio-motor interaction, relatively little is known about the effect of specific motor processes on auditory rhythm perception. This doctoral thesis was devoted to investigating the influence of both external and internal motor processes on the way we perceive an auditory rhythm. The first half of the thesis intended to establish whether overt body movement had a facilitatory effect on our ability to perceive the auditory rhythmic structure, and whether this effect was modulated by musical training. To this end, musicians and non-musicians performed a pulse-finding task either using natural body movement or through listening only, and produced their identified pulse by finger tapping. The results showed that overt movement benefited rhythm (pulse) perception especially for non-musicians, confirming the facilitatory role of external motor activities in hearing the rhythm, as well as its interaction with musical training. The second half of the thesis tested the idea that indirect, covert motor input, such as that transformed from the visual stimuli, could influence our perceived structure of an auditory rhythm. Three experiments examined the subjectively perceived tempo of an auditory sequence under different visual motion stimulations, while the auditory and visual streams were presented independently of each other. The results revealed that the perceived auditory tempo was accordingly influenced by the concurrent visual motion conditions, and the effect was related to the increment or decrement of visual motion speed. This supported the hypothesis that the internal motor information extracted from the visuomotor stimulation could be incorporated into the percept of an auditory rhythm. Taken together, the present thesis concludes that, rather than as a mere reaction to the given auditory input, our motor system plays an important role in contributing to the perceptual process of the auditory rhythm. This can occur via both external and internal motor activities, and may not only influence how we hear a rhythm but also under some circumstances improve our ability to hear the rhythm.Musikalische Rhythmen bestehen aus zeitlich strukturierten Mustern akustischer Stimuli. Es konnte gezeigt werden, dass die Prozesse, welche der Rhythmuswahrnehmung zugrunde liegen, sowohl motorische als auch auditive Systeme nutzen. Obwohl sich für diese auditiv-motorischen Interaktionen sowohl in den Verhaltenswissenschaften als auch Neurowissenschaften übereinstimmende Belege finden, weiß man bislang relativ wenig über die Auswirkungen spezifischer motorischer Prozesse auf die auditive Rhythmuswahrnehmung. Diese Doktorarbeit untersucht den Einfluss externaler und internaler motorischer Prozesse auf die Art und Weise, wie auditive Rhythmen wahrgenommen werden. Der erste Teil der Arbeit diente dem Ziel herauszufinden, ob körperliche Bewegungen es dem Gehirn erleichtern können, die Struktur von auditiven Rhythmen zu erkennen, und, wenn ja, ob dieser Effekt durch ein musikalisches Training beeinflusst wird. Um dies herauszufinden wurde Musikern und Nichtmusikern die Aufgabe gegeben, innerhalb von präsentierten auditiven Stimuli den Puls zu finden, wobei ein Teil der Probanden währenddessen Körperbewegungen ausführen sollte und der andere Teil nur zuhören sollte. Anschließend sollten die Probanden den gefundenen Puls durch Finger-Tapping ausführen, wobei die Reizgaben sowie die Reaktionen mittels eines computerisierten Systems kontrolliert wurden. Die Ergebnisse zeigen, dass offen ausgeführte Bewegungen die Wahrnehmung des Pulses vor allem bei Nichtmusikern verbesserten. Diese Ergebnisse bestätigen, dass Bewegungen beim Hören von Rhythmen unterstützend wirken. Außerdem zeigte sich, dass hier eine Wechselwirkung mit dem musikalischen Training besteht. Der zweite Teil der Doktorarbeit überprüfte die Idee, dass indirekte, verdeckte Bewegungsinformationen, wie sie z.B. in visuellen Stimuli enthalten sind, die wahrgenommene Struktur von auditiven Rhythmen beeinflussen können. Drei Experimente untersuchten, inwiefern das subjektiv wahrgenommene Tempo einer akustischen Sequenz durch die Präsentation unterschiedlicher visueller Bewegungsreize beeinflusst wird, wobei die akustischen und optischen Stimuli unabhängig voneinander präsentiert wurden. Die Ergebnisse zeigten, dass das wahrgenommene auditive Tempo durch die visuellen Bewegungsinformationen beeinflusst wird, und dass der Effekt in Verbindung mit der Zunahme oder Abnahme der visuellen Geschwindigkeit steht. Dies unterstützt die Hypothese, dass internale Bewegungsinformationen, welche aus visuomotorischen Reizen extrahiert werden, in die Wahrnehmung eines auditiven Rhythmus integriert werden können. Zusammen genommen, 5 zeigt die vorgestellte Arbeit, dass unser motorisches System eine wichtige Rolle im Wahrnehmungsprozess von auditiven Rhythmen spielt. Dies kann sowohl durch äußere als auch durch internale motorische Aktivitäten geschehen, und beeinflusst nicht nur die Art, wie wir Rhythmen hören, sondern verbessert unter bestimmten Bedingungen auch unsere Fähigkeit Rhythmen zu identifizieren

From locomotion to dance and back : exploring rhythmic sensorimotor synchronization

Le rythme est un aspect important du mouvement et de la perception de l’environnement. Lorsque l’on danse, la pulsation musicale induit une activité neurale oscillatoire qui permet au système nerveux d’anticiper les évènements musicaux à venir. Le système moteur peut alors s’y synchroniser. Cette thèse développe de nouvelles techniques d’investigation des rythmes neuraux non strictement périodiques, tels que ceux qui régulent le tempo naturellement variable de la marche ou la perception rythmes musicaux. Elle étudie des réponses neurales reflétant la discordance entre ce que le système nerveux anticipe et ce qu’il perçoit, et qui sont nécessaire pour adapter la synchronisation de mouvements à un environnement variable. Elle montre aussi comment l’activité neurale évoquée par un rythme musical complexe est renforcée par les mouvements qui y sont synchronisés. Enfin, elle s’intéresse à ces rythmes neuraux chez des patients ayant des troubles de la marche ou de la conscience.Rhythms are central in human behaviours spanning from locomotion to music performance. In dance, self-sustaining and dynamically adapting neural oscillations entrain to the regular auditory inputs that is the musical beat. This entrainment leads to anticipation of forthcoming sensory events, which in turn allows synchronization of movements to the perceived environment. This dissertation develops novel technical approaches to investigate neural rhythms that are not strictly periodic, such as naturally tempo-varying locomotion movements and rhythms of music. It studies neural responses reflecting the discordance between what the nervous system anticipates and the actual timing of events, and that are critical for synchronizing movements to a changing environment. It also shows how the neural activity elicited by a musical rhythm is shaped by how we move. Finally, it investigates such neural rhythms in patient with gait or consciousness disorders

Exploring the neural entrainment to musical rhythms and meter : a steady-state evoked potential approach

Thèse de doctorat réalisé en cotutelle avec l'Université catholique de Louvain, Belgique (Faculté de médecine, Institut de Neuroscience)Percevoir et synchroniser ses mouvements à une pulsation régulière en musique est une capacité largement répandue chez l’Homme, et fondamentale aux comportements musicaux. La pulsation et la métrique en musique désignent généralement une organisation temporelle périodique perçue à partir de stimuli acoustiques complexes, et cette organisation perceptuelle implique souvent une mise en mouvement périodique spontanée du corps. Cependant, les mécanismes neuraux sous-tendant cette perception sont à l’heure actuelle encore méconnus. Le présent travail a donc eu pour objectif de développer une nouvelle approche expérimentale, inspirée par l’approche électrophysiologique des potentiels évoqués stationnaires, afin d’explorer les corrélats neuraux à la base de notre perception de la pulsation et de la métrique induite à l’écoute de rythmes musicaux. L’activité neurale évoquée en relation avec la perception d’une pulsation a été enregistrée par électroencéphalographie (EEG) chez des individus sains, dans divers contextes : (1) dans un contexte d’imagerie mentale d’une métrique appliquée de manière endogène sur un stimulus auditif, (2) dans un contexte d’induction spontanée d’une pulsation à l’écoute de patterns rythmiques musicaux, (3) dans un contexte d’interaction multisensorielle, et (4) dans un contexte de synchronisation sensorimotrice. Pris dans leur ensemble, les résultats de ces études corroborent l’hypothèse selon laquelle la perception de la pulsation en musique est sous-tendue par des processus de synchronisation et de résonance de l’activité neurale dans le cerveau humain. De plus, ces résultats suggèrent que l’approche développée dans le présent travail pourrait apporter un éclairage significatif pour comprendre les mécanismes neuraux de la perception de la pulsation et des rythmes musicaux, et, dans une perspective plus générale, pour explorer les mécanismes de synchronisation neurale.The ability to perceive a regular beat in music and synchronize to it is a widespread human skill. Fundamental to musical behavior, beat and meter refer to the perception of periodicities while listening to musical rhythms, and usually involve spontaneous entrainment to move on these periodicities. However, the neural mechanisms underlying entrainment to beat and meter in Humans remain unclear. The present work tests a novel experimental approach, inspired by the steady-state evoked potential method, to explore the neural dynamics supporting the perception of rhythmic inputs. Using human electroencephalography (EEG), neural responses to beat and meter were recorded in various contexts: (1) mental imagery of meter, (2) spontaneous induction of a beat from rhythmic patterns, (3) multisensory integration, and (4) sensorimotor synchronization. Our results support the view that entrainment and resonance phenomena subtend the processing of musical rhythms in the human brain. Furthermore, our results suggest that this novel approach could help investigating the link between the phenomenology of musical beat and meter and neurophysiological evidence of a bias towards periodicities arising under certain circumstances in the nervous system. Hence, entrainment to music provides an original framework to explore general entrainment phenomena occurring at various levels, from the inter-neural to the inter-individual level

Cortico-muscular coherence in sensorimotor synchronisation

This thesis sets out to investigate the neuro-muscular control mechanisms underlying the ubiquitous phenomenon of sensorimotor synchronisation (SMS). SMS is the coordination of movement to external rhythms, and is commonly observed in everyday life. A large body of research addresses the processes underlying SMS at the levels of behaviour and brain. Comparatively, little is known about the coupling between neural and behavioural processes, i.e. neuro-muscular processes. Here, the neuro-muscular processes underlying SMS were investigated in the form of cortico-muscular coherence measured based on Electroencephalography (EEG) and Electromyography (EMG) recorded in human healthy participants. These neuro-muscular processes were investigated at three levels of engagement: passive listening and observation of rhythms in the environment, imagined SMS, and executed SMS, which resulted in the testing of three hypotheses: (i) Rhythms in the environment, such as music, spontaneously modulate cortico-muscular coupling, (ii) Movement intention modulates cortico-muscular coupling, and (iii) Cortico-muscular coupling is dynamically modulated during SMS time-locked to the stimulus rhythm. These three hypotheses were tested through two studies that used Electroencephalography (EEG) and Electromyography (EMG) recordings to measure Cortico-muscular coherence (CMC). First, CMC was tested during passive music listening, to test whether temporal and spectral properties of music stimuli known to induce groove, i.e., the subjective experience of wanting to move, can spontaneously modulate the overall strength of the communication between the brain and the muscles. Second, imagined and executed movement synchronisation was used to investigate the role of movement intention and dynamics on CMC. The two studies indicate that both top-down, and somatosensory and/or proprioceptive processes modulate CMC during SMS tasks. Although CMC dynamics might be linked to movement dynamics, no direct correlation between movement performance and CMC was found. Furthermore, purely passive auditory or visual rhythmic stimulation did not affect CMC. Together, these findings thus indicate that movement intention and active engagement with rhythms in the environment might be critical in modulating CMC. Further investigations of the mechanisms and function of CMC are necessary, as they could have important implications for clinical and elderly populations, as well as athletes, where optimisation of motor control is necessary to compensate for impaired movement or to achieve elite performance

Frontal alpha oscillations distinguish leaders from followers: Multivariate decoding of mutually interacting brains

Successful social interactions rely upon the abilities of two or more people to mutually exchange information in real-time, while simultaneously adapting to one another. The neural basis of social cognition has mostly been investigated in isolated individuals, and more recently using two-person paradigms to quantify the neuronal dynamics underlying social interaction. While several studies have shown the relevance of understanding complementary and mutually adaptive processes, the neural mechanisms underlying such coordinative behavioral patterns during joint action remain largely unknown. Here, we employed a synchronized finger-tapping task while measuring dual-EEG from pairs of human participants who either mutually adjusted to each other in an interactive task or followed a computer metronome. Neurophysiologically, the interactive condition was characterized by a stronger suppression of alpha and low-beta oscillations over motor and frontal areas in contrast to the non-interactive computer condition. A multivariate analysis of two-brain activity to classify interactive versus non-interactive trials revealed asymmetric patterns of the frontal alpha-suppression in each pair, during both task anticipation and execution, such that only one member showed the frontal component. Analysis of the behavioral data showed that this distinction coincided with the leader–follower relationship in 8/9 pairs, with the leaders characterized by the stronger frontal alpha-suppression. This suggests that leaders invest more resources in prospective planning and control. Hence our results show that the spontaneous emergence of leader–follower relationships in dyadic interactions can be predicted from EEG recordings of brain activity prior to and during interaction. Furthermore, this emphasizes the importance of investigating complementarity in joint action

Modelling single-person and multi-person event-based synchronisation

A linear phase correction model has been shown to accurately reflect the corrective processes involved in synchronising motor actions to an external rhythmic cue. The model originated from studies of finger tapping to an isochronous metronome beat and is based on the time series of asynchronies between the metronome and corresponding finger tap onsets, along with their associated intervals. Over recent years the model has evolved and been applied to more complex scenarios, including phase perturbed cues, tempo variations and, most recently, timing within groups. Here, we review the studies that have contributed to the development of the linear phase correction model and the associated findings related to human timing performance. The review provides a background to the studies examining single-person timing to simple metronome cues. We then further expand on the more complex analyses of motor timing to phase and tempo shifted cues. Finally, recent studies investigating inter-personal synchronisation between groups of two or more individuals are discussed, along with a brief overview on the implications of these studies for social interactions. We conclude with a discussion on future areas of research that will be important for understanding corrective timing processes between people

When Coordinating Finger Tapping to a Variable Beat the Variability Scaling Structure of the Movement and the Cortical BOLD Signal are Both Entrained to the Auditory Stimuli

Rhythmic actions are characterizable as a repeating invariant pattern of movement together with variability taking the form of cycle-to-cycle fluctuations. Variability in behavioral measures is atypically random, and often exhibits serial temporal dependencies and statistical self-similarity in the scaling of variability magnitudes across timescales. Self-similar (i.e. fractal) variability scaling is evident in measures of both brain and behavior. Variability scaling structure can be quantified via the scaling exponent (α) from detrended fluctuation analysis (DFA). Here we study the task of coordinating thumb-finger tapping to the beats of constructed auditory stimuli. We test the hypothesis that variability scaling evident in tap-to-tap intervals as well as in the fluctuations of cortical hemodynamics will become entrained to (i.e. drawn toward) manipulated changes in the variability scaling of a stimulus’s beat-to-beat intervals. Consistent with this hypothesis, manipulated changes of the exponent α of the experimental stimuli produced corresponding changes in the exponent α of both tap-to-tap intervals and cortical hemodynamics. The changes in hemodynamics were observed in both motor and sensorimotor cortical areas in the contralateral hemisphere. These results were observed only for the longer timescales of the detrended fluctuation analysis used to measure the exponent α. These findings suggest that complex auditory stimuli engage both brain and behavior at the level of variability scaling structures

Neural alpha oscillations index the balance between self-other integration and segregation in real-time joint action

Shared knowledge and interpersonal coordination are prerequisites for most forms of social behavior. Influential approaches to joint action have conceptualized these capacities in relation to the separate constructs of co-representation (knowledge) and self-other entrainment (coordination). Here we investigated how brain mechanisms involved in co-representation and entrainment interact to support joint action. To do so, we used a musical joint action paradigm to show that the neural mechanisms underlying co-representation and self-other entrainment are linked via a process – indexed by EEG alpha oscillations – regulating the balance between self-other integration and segregation in real time. Pairs of pianists performed short musical items while action familiarity and interpersonal (behavioral) synchronization accuracy were manipulated in a factorial design. Action familiarity referred to whether or not pianists had rehearsed the musical material performed by the other beforehand. Interpersonal synchronization was manipulated via congruent or incongruent tempo change instructions that biased performance timing towards the impending, new tempo. It was observed that, when pianists were familiar with each other's parts, millisecond variations in interpersonal synchronized behavior were associated with a modulation of alpha power over right centro-parietal scalp regions. Specifically, high behavioral entrainment was associated with self-other integration, as indexed by alpha suppression. Conversely, low behavioral entrainment encouraged reliance on internal knowledge and thus led to self-other segregation, indexed by alpha enhancement. These findings suggest that alpha oscillations index the processing of information about self and other depending on the compatibility of internal knowledge and external (environmental) events at finely resolved timescales