Sounds on time - N° d’ordre 232 - 2013 THESE DE L‘UNIVERSITE DE LYON Délivrée par L’UNIVERSITE CLAUDE BERNARD LYON 1 ECOLE DOCTORALE NEUROSCIENCES ET COGNITION (NSCo) DIPLOME DE DOCTORAT EN NEUROSCIENCES COGNITIVES (arrêté du 7 août 2006) Soutenue publiquement le 27 novembre 2013 à Lyon par Floris Tijmen VAN VUGT Directeurs de hèse: Dr. Barbara TILLMANN et Prof. Eckart ALTENMÜLLER Composition du Jury Prof. Eckart ALTENMÜLLER (directeur de thèse) Dr. Barbara TILLMANN (directeur de thèse) Prof. Peter KELLER (rapporteur) Prof. Virginia PENHUNE (rapporteur) Dr. Fabien PERRIN (président du jury) | 3 | Sounds on time - Sounds on time: auditory feedback in motor learning, re-learning and over-learning of timing regularity. Auditory feedback is an auditory signal that contains information about performed movement. Music performance is an excellent candidate to study its inluence on motor actions, since the auditory result is the explicit goal of the movement. Indeed, auditory feedback can guide online motor actions, but its inluence on motor learning has been investigated less. his thesis investigates the inluence of auditory feedback in motor learning, focusing particularly on how we learn temporal control over movements. First, we investigate motor learning in non-musicians, inding that they beneit from temporal information supplied by the auditory signal and are sensitive to distortions of this temporal information. Second, we turn to stroke patients that are re-learning motor actions in a rehabilitation seting. Patients improved their movement capacities but did not depend on the time-locking between movements and the resulting auditory feedback. Surprisingly, they appear to beneit from distortions in feedback. hird, we investigate musical experts, who arguably have established strong links between movement and auditory feedback. We develop a novel analysis framework that allows us to segment timing into systematic and non-systematic variability. Our inding is that these experts have become largely independent of the auditory feedback. he main claim defended in this thesis is that auditory feedback can and does play a role in motor learning of regularity, but the way in which it is used varies qualitatively between diferent populations. hese indings provide new insights into auditory-motor integration and are relevant for developing new perspectives on the role of music in training and rehabilitation setings. Keywords: Sensorimotor Integration, Auditory Feedback, Motor Learning, Motor regularity, Timing, Rehabilitation, Expert musicians, Sequence Production, Sequence Learning | 4 | Sounds on time - Feedback auditif et régularité motrice : apprentissage, rehabilitation et expertise Le feedback auditif se déinit comme un signal auditif qui contient de l'information sur un mouvement. Il a été montré que le feedback auditif peut guider le mouvement en temps réel, mais son inluence sur l'apprentissage moteur est moins clair. Cete thèse a pour but d'examiner l'inluence du feedback auditif sur l'apprentissage moteur, en se focalisant sur le contrôle temporel des mouvements. Premièrement, nous étudions l'apprentissage moteur chez les non-musiciens sains et montrons qu'ils bénéicient de l'information temporelle contenue dans le feedback auditif et qu'ils sont sensibles aux distortions de cete information temporelle. Deuxièmement, nous appliquons ces connaissances à la rehabilitation de patients cérébro-lésés. Nous trouvons que ces patients améliorent leurs capacités de mouvement mais ne dépendent pas de la correspondance temporelle entre le mouvement et le son. Paradoxalement, ces patients ont même benéicié des distortions temporelles dans le feedback. Troisièmement, nous étudions les experts musicaux, car ils ont établi des liens particulièrement forts entre leur mouvement et le son. Nous développons de nouveaux outils d'analyse qui nous permetent de séparer les déviations temporelles en variation systématique et non-systématique. Le résultat principal est que ces experts sont devenu largement indépendents du feedback auditif. La proposition centrale de cete thèse est que le feedback auditif joue un rôle dans l'apprentissage moteur de la regularité, mais la façon dont le cerveau l'utilise dépend de la population étudiée. Ces résultats donnent une nouvelle perspective sur l'intégration audio-motrice et contribuent au développement de nouvelles approches pour l'apprentissage de la musique et la réhabilitation. Mots clés: Intégration sensorimotrice, feedback auditif, apprentissage moteur, regularité motrice, timing, réhabilitation, experts musicaux, production de séquences, apprentissage de séquences. | 5 | Sounds on time - Institutes: Institute of Music Physiology and Musicians’ Medicine, University of Music, Drama and Media, Emmichplatz 1, 30175 Hanover Germany Lyon Neuroscience Research Center CNRS-UMR 5292, INSERM U1028, University Lyon-1, 50 av Tony Garnier, 69007 Lyon, France his research was supported by the EBRAMUS (European Brain and Music) grant, ITN MC FP7, GA 238157. | 6 | Sounds on time - he musician may sing to you of the rhythm which is in all space, but he cannot give you the ear which arrests the rhythm nor the voice that echoes it. – Kahlil Gibran, he Prophet | 7 | Sounds on time - | 8 | Sounds on time - Table of Contents I. Sensorimotor integration: a conceptual map.......................19 I.1. Introduction.......................................20 1.1 Aim.................................................20 1.2 Disclaimer: inding a balance between unity and diversity...............21 I.2. Preliminaries......................................22 2.1 he human motor system – a brief overview........................23 2.1.a Motor control areas – a brief outline..............................23 2.1.b Control Strategies..........................................26 I.3. Towards a deinition of sensorimotor integration................28 3.1 Sensorimotor integration....................................28 3.2 Aim.................................................29 I.4. Motor theories of cognition..............................31 I.5. Motor involvement in perception...........................32 5.1 Movement making perception possible..........................34 5.1.a Microsaccades (vision)......................................35 5.1.b Motor-induced suppression....................................36 5.2 Action perception.......................................39 5.2.a Mirror neurons...........................................39 5.2.b Mirror networks...........................................43 5.2.c Direct matching hypothesis and motor resonance......................46 5.2.d Plasticity of mirror networks: arbitrariness of the associations..............47 5.3 Speech perception and comprehension...........................50 5.3.a Perception of speech sounds...................................50 5.3.b Syntactic processing........................................54 | 9 | Sounds on time - 5.3.c Semantic processing........................................55 5.3.d Conversational interaction....................................56 5.3.e Anatomical substrates.......................................57 5.3.f Necessity and role of motor involvement............................57 5.4 Music perception.........................................58 5.4.a Why music?.............................................58 5.4.b Motor systems supporting perceptual memory formation.................59 5.4.c Music perception in novices: a focus on rhythm perception................61 5.4.d Musical experts..........................................62 5.4.e Creation of auditory-motor associations: music learning in novices...........64 I.6. Sensory involvement in action............................66 6.1 Sensory feedback in action...................................70 6.1.a Representing action by its sensory consequences (action selection)............70 6.1.b Sensory guidance of action (action execution).........................72 6.1.c Motor learning through sensory feedback (action learning)................73 6.2 Sensory stimulation in action................................75 6.2.a Sensorimotor corticospinal facilitation............................75 6.2.b Sensation as a reference for movement.............................77 6.3 Speech production........................................79 6.3.a Sensory co-activation........................................79 6.3.b Reliance on feedback........................................79 6.4 Music production........................................80 6.4.a Sensory co-activation during silent performance......................81 6.4.b Reliance on feedback........................................82 6.4.c Soniication feedback in musical novices...........................82 I.7. Synthesis.........................................84 I.8. Outline of the thesis...................................85 II. Learning in musical novices...............................87 II.1. Introduction......................................88 1.1 Motor regularity........................................88 | 10 | Sounds on time - 1.2 Serial Reaction Time......................................89 1.3 Aims and Hypotheses......................................91 II.2. hresholds of auditory-motor coupling measured with a simple task in musicians and non-musicians: Was the sound simultaneous to the keystroke?.92 2.1 Introduction............................................94 2.2 Materials and Methods....................................96 2.2.a Participants..............................................96