The Bereitschaftspotential (BP) is an event related potential believed to reflect motor planning and preparedness. Although the relationship between the BP and volitional movements of the distal limbs is well established, studies of the BP and speech have produced inconclusive findings. The most heavily debated of these findings were reports of left lateralized hemispheric asymmetry in the BP topography, shortly before speech onset. Several researchers argued that these shifts were artifacts produced by movements of the articulatory muscles. However, methodological differences between the studies could also explain why the asymmetry was not always found. In the present study it was proposed that articulatory complexity and semantic processing each contribute to observed variations in the speech BP topography. Eighteen healthy volunteers performed 3 speech tasks, designed to distinguish semantic and articulatory contributions to the BP topography. The findings suggested that articulatory complexity and semantic processing each uniquely contribute to the frontolateral and medial BP topographic distribution. The present study also introduced the use of Doppler imaging of the tongue as a means of eliminating potential artifactual tongue movements from the speech BP. / Ph. D.

Submitted by Daniella Sodre (daniella.sodre@ufpe.br) on 2015-03-13T16:07:58Z No. of bitstreams: 2 DISSERTAÇÃO Isabella Araújo Mota.pdf: 2205369 bytes, checksum: c6172d160e112abf88146c5aec509245 (MD5) license_rdf: 1232 bytes, checksum: 66e71c371cc565284e70f40736c94386 (MD5) / Made available in DSpace on 2015-03-13T16:07:58Z (GMT). No. of bitstreams: 2 DISSERTAÇÃO Isabella Araújo Mota.pdf: 2205369 bytes, checksum: c6172d160e112abf88146c5aec509245 (MD5) license_rdf: 1232 bytes, checksum: 66e71c371cc565284e70f40736c94386 (MD5) Previous issue date: 2014-04-11 / O Bereitschaftspotential (BP) ou potencial de prontidão é uma onda negativa observada retrogradamente no eletroencefalograma (EEG) precedendo um ato motor. O BP tem sido usado na investigação dos mecanismos corticais de geração do movimento voluntário. A lentificação dos movimentos (bradicinesia) é um dos sinais cardinais da doença de Parkinson (DP). Objetivos: Estudar o BP precedendo o piscamento ocular em sujeitos normais e em pacientes com DP durante as fases Off (sem levodopa) e On (com levodopa). Métodos: Foram avaliados 18 sujeitos saudáveis (oito homens) entre 17 e 60 anos (média 35 anos) e 10 pacientes com DP (nove homens) entre 43 e 86 anos (média 58 anos) nos estágios 1 e 2 da escala de Hoehn & Yahr (H&Y). O EEG foi registrado em 11 posições do sistema 10-20 de colocação de eletrodos, em referência aos mastoides interligados. O eletro-oculograma vertical (VEOG) foi registrado por dois eletrodos posicionados acima e abaixo do olho direito. O eletrodo terra foi colocado na clavícula direita. Nos sujeitos normais os registros foram obtidos durante o piscamento palpebral espontâneo e voluntário. Nos pacientes com DP apenas o piscamento voluntário foi analisado e os registros obtidos durante as fases Off e On. Segmentos do EEG entre três segundos antes e dois segundos após o início do potencial do piscamento (identificado visualmente no canal de VEOG), livres de artefatos, foram promediados off-line e analisados em programa dedicado escrito em Matlab. A análise de significância estatística foi feita com modelos de Análise de Variância ANOVA. Resultados: Nos sujeitos normais a média do BP iniciou em torno de 1700 ms antes do início do piscamento voluntário, com amplitude média no pico da negatividade de 3.3 μV. Não se observou BP precedendo o piscamento espontâneo. Nos pacientes com DP, a média do BP iniciou em torno de 1500 ms antes do piscamento voluntário, com amplitude média no pico de negatividade de 0.6 μV. A média de amplitude do BP não foi significativamente diferente durante as fases Off e On. Conclusão: Este estudo demonstrou a presença de um BP precedendo o piscamento palpebral 11 voluntário, porém não precedendo ao piscamento palpebral espontâneo. Além disso, demontrou a presença de um BP com amplitude acentuadamente reduzida precedendo o piscamento voluntário de pacientes com DP nas fases 1 e 2 da escala H&Y. O uso da levodopa não aumentou a amplitude do BP. Estes dados indicam que o BP precedendo o piscamento palpebral é significativamente acometido já nas fases iniciais da DP, independentemente do uso da levodopa.

A execução de movimentos voluntários é precedida por processos preparatórios que manifestam-se eletrofisiologicamente como um potencial elétrico lento denominado Bereitschaftspotential. As investigações sobre o significado fisiológico desses potenciais mostram-se muitas vezes inconclusivas devido à dificuldade de isolamento dos fatores ambientais e endógenos que os influenciam. Dentre essas variáveis, a expressão consciente do ato motor tem sido negligenciada por grande parte dos pesquisadores. O presente trabalho consiste em uma série de estudos eletrofisiológicos que visam demonstrar a influência do controle consciente sobre os processos preparatórios motores. Parte-se do desenvolvimento de protocolos comportamentais que modulam o envolvimento consciente sobre processos motores. Desse modo, verificamos que grande parte do Bereitschaftspotential comumente observado durante a geração de movimentos espontâneos estão associados com a escolha deliberada de agir. Em seguida, são investigadas como a variável em questão pode afetar os processos relacionados com a livre escolha de movimentos. Nossos resultados indicam que processos relacionados ao controle consciente afetam nitidamente a expressão de livre escolha sobre os potenciais preparatórios motores. Por fim, nós demonstramos como a influência do controle consciente nos processos motores pode estar afetada em pacientes com transtorno obsessivo-compulsivo / The execution of voluntary movements is preceded by preparatory processes that electrophysiologically are manifested as a slow electric potential called Bereitschaftspotential. Investigations on the physiological significance of these potentials are often inconclusive due to the difficulty of isolating the environmental or endogenous factors that influence them. Among these factors, the conscious expression of the motor act has been neglected by most researchers. The present work consists of a series of electrophysiological studies aimed at demonstrating the influence of conscious control on preparatory motor processes. Firstly, behavioral protocols that modulate the conscious involvement in motor control are proposed. Using these methods, we found that great part of the Bereitschaftspotential commonly observed during the generation of spontaneous movements is associated with the deliberate choice to act. Next, we investigated how this factor variable affects the processes related to the free mode of movement selection. Our results indicate that processes related to conscious control clearly affect the manifestations of free choice in preparatory motor potentials. Finally, we here demonstrated how the influence of conscious control on the motor processes can be affected in the obsessive-compulsive disorder

A execução de movimentos voluntários é precedida por processos preparatórios que manifestam-se eletrofisiologicamente como um potencial elétrico lento denominado Bereitschaftspotential. As investigações sobre o significado fisiológico desses potenciais mostram-se muitas vezes inconclusivas devido à dificuldade de isolamento dos fatores ambientais e endógenos que os influenciam. Dentre essas variáveis, a expressão consciente do ato motor tem sido negligenciada por grande parte dos pesquisadores. O presente trabalho consiste em uma série de estudos eletrofisiológicos que visam demonstrar a influência do controle consciente sobre os processos preparatórios motores. Parte-se do desenvolvimento de protocolos comportamentais que modulam o envolvimento consciente sobre processos motores. Desse modo, verificamos que grande parte do Bereitschaftspotential comumente observado durante a geração de movimentos espontâneos estão associados com a escolha deliberada de agir. Em seguida, são investigadas como a variável em questão pode afetar os processos relacionados com a livre escolha de movimentos. Nossos resultados indicam que processos relacionados ao controle consciente afetam nitidamente a expressão de livre escolha sobre os potenciais preparatórios motores. Por fim, nós demonstramos como a influência do controle consciente nos processos motores pode estar afetada em pacientes com transtorno obsessivo-compulsivo / The execution of voluntary movements is preceded by preparatory processes that electrophysiologically are manifested as a slow electric potential called Bereitschaftspotential. Investigations on the physiological significance of these potentials are often inconclusive due to the difficulty of isolating the environmental or endogenous factors that influence them. Among these factors, the conscious expression of the motor act has been neglected by most researchers. The present work consists of a series of electrophysiological studies aimed at demonstrating the influence of conscious control on preparatory motor processes. Firstly, behavioral protocols that modulate the conscious involvement in motor control are proposed. Using these methods, we found that great part of the Bereitschaftspotential commonly observed during the generation of spontaneous movements is associated with the deliberate choice to act. Next, we investigated how this factor variable affects the processes related to the free mode of movement selection. Our results indicate that processes related to conscious control clearly affect the manifestations of free choice in preparatory motor potentials. Finally, we here demonstrated how the influence of conscious control on the motor processes can be affected in the obsessive-compulsive disorder

Emotiv EPOC is a low-cost consumer headset capable of acquiring a raw EEG signal. The thesis evaluates its usage for an acquisition of research event-related potentials. A controlled laboratory experiment was performed with an objective of isolating the Bereitschaftspotential (Readiness Potential) and other movement-related potentials and comparing the results with the results of the previous research. The possibility of measuring the potential with Emotiv EPOC was not confirmed, most probably due to procedural issues during the experiment, however, further analysis of the data suggests its presence. Other outcomes of the research include qualitative findings about the headset, mainly its hardware construction, and testing feedback of EEGLab, an open source toolbox for EEG processing and visualization.

京都大学 / 0048 / 新制・論文博士 / 博士(医学) / 乙第12817号 / 論医博第2079号 / 新制||医||1004(附属図書館) / 31304 / 京都大学大学院医学研究科脳統御医科学系専攻 / (主査)教授 河野 憲二, 教授 金子 武嗣, 教授 大森 治紀 / 学位規則第4条第2項該当 / Doctor of Medical Science / Kyoto University / DFAM

This study examined the brain electrical activity of normal speakers in a non-delayed auditory feedback (DAF) condition and when experiencing DAF to determine the effect DAF would have on the Bereitschaftspotential (BP). The BP reflects the preparatory state of a person prior to motor execution of an act and can be observed 1500 to 500 ms prior to voluntary movement. The participants in the study included 10 adults with normal speech. Each read a series of 30 sentences, both without DAF and with DAF, while the BP was measured. Results indicate that the BP is present across the scalp in both the control condition and the DAF condition; however, the BP is reduced in the DAF condition. The scalp distribution maps indicate an increased negativity in the left frontal lobe in the DAF condition. These findings suggest that while the brain is engaged in processing current information that has already been initiated, the motor system may not be able to be primed for the next sequential motor event. There is still a need for more research to explore the motor control of speech and the ways altered feedback may disrupt the speech motor control.

In the course of development organisms learn to associate their actions with the effects these actions have in the environment. Recent studies have shown that perceiving or anticipating action-effects automatically activates actions, which formerly have been experienced to cause these effects (Elsner & Hommel, 2001). Using subliminal priming paradigms and electrophysiological measures I investigated whether subliminally (i.e., not consciously perceivable) presented action-effects can automatically activate associated actions and if so, whether this response priming by action-effects can bias free-choice actions. Secondly I investigated whether action-effects with different emotional valences influence response selection differently. To address the first question three experiments were performed. Each experiment consisted of two experimental phases. The first phase, the acquisition-phase, was a learning phase were simple key-press actions were associated with simple visual stimuli (i.e., action-effects; diamond or square) that were contingent on the actions. Immediately after the acquisition-phase the test-phase followed, in which participants performed free-choice actions after the presentation of a Go-signal. In Experiments 2 and 3 a NoGo-signal indicating that responses had to be withheld could appear with the same likelihood as the Go-signal. Unknown to the participants, one of the former action-effects (diamond or square) was presented subliminally prior to each Go- and NoGo-signal to investigate the influence of unconscious action-effects on response selection. Taken together, the results of the test-phases provided strong evidence that even subliminally presented (i.e., unconscious) action-effects can automatically activate associated responses. The response priming by action-effects became evident in the lateralized readiness potential (LRP), an electrophysiological indicator of specific response activation processes. Under certain circumstanced this automatic response activation can bias free-choice actions although participants experienced the actions as freely chosen. In the test-phase of the first experiment more acquisition-phase-consistent than –inconsistent responses were chosen. If, for instance, a left key-press had been associated with a square during the acquisition-phase, the left key was chosen significantly more often after the subliminal presentation of a square in the test-phase. At least three factors seemed to influence which responses were chosen and executed: The strength of the priming effect, the complexity of the task (i.e., pure Go-blocks or intermixed Go/NoGo-blocks), and the elapsed time between the prime stimulus and the Go-signal. To address the second question simple key-press actions were linked to action-effects with different emotional valences (positive vs. negative pictures accompanied by high or low tones) during the acquisition-phase. In the subsequent test-phase, the effects-tones that had been associated with negative or positive pictures were presented and followed by a Go-signal, after which participants had to freely choose to press one of the two response keys. Results indicated that the anticipation of the emotional valence of an action-effect influenced free-choice action. Whereas the effect-tones induced a clear response bias (i.e., more acquisition-consistent than –inconsistent key-choices) if they had been associated with a positive emotional valence, this response bias was not reliable for action-effects associated with negative emotional features. In summary, the present results provide further proof for ideomotor theories of action control (James, 1890; Elsner & Hommel, 2001) which state that actions are automatically activated by anticipating their consequences.

In the course of development organisms learn to associate their actions with the effects these actions have in the environment. Recent studies have shown that perceiving or anticipating action-effects automatically activates actions, which formerly have been experienced to cause these effects (Elsner & Hommel, 2001). Using subliminal priming paradigms and electrophysiological measures I investigated whether subliminally (i.e., not consciously perceivable) presented action-effects can automatically activate associated actions and if so, whether this response priming by action-effects can bias free-choice actions. Secondly I investigated whether action-effects with different emotional valences influence response selection differently. To address the first question three experiments were performed. Each experiment consisted of two experimental phases. The first phase, the acquisition-phase, was a learning phase were simple key-press actions were associated with simple visual stimuli (i.e., action-effects; diamond or square) that were contingent on the actions. Immediately after the acquisition-phase the test-phase followed, in which participants performed free-choice actions after the presentation of a Go-signal. In Experiments 2 and 3 a NoGo-signal indicating that responses had to be withheld could appear with the same likelihood as the Go-signal. Unknown to the participants, one of the former action-effects (diamond or square) was presented subliminally prior to each Go- and NoGo-signal to investigate the influence of unconscious action-effects on response selection. Taken together, the results of the test-phases provided strong evidence that even subliminally presented (i.e., unconscious) action-effects can automatically activate associated responses. The response priming by action-effects became evident in the lateralized readiness potential (LRP), an electrophysiological indicator of specific response activation processes. Under certain circumstanced this automatic response activation can bias free-choice actions although participants experienced the actions as freely chosen. In the test-phase of the first experiment more acquisition-phase-consistent than –inconsistent responses were chosen. If, for instance, a left key-press had been associated with a square during the acquisition-phase, the left key was chosen significantly more often after the subliminal presentation of a square in the test-phase. At least three factors seemed to influence which responses were chosen and executed: The strength of the priming effect, the complexity of the task (i.e., pure Go-blocks or intermixed Go/NoGo-blocks), and the elapsed time between the prime stimulus and the Go-signal. To address the second question simple key-press actions were linked to action-effects with different emotional valences (positive vs. negative pictures accompanied by high or low tones) during the acquisition-phase. In the subsequent test-phase, the effects-tones that had been associated with negative or positive pictures were presented and followed by a Go-signal, after which participants had to freely choose to press one of the two response keys. Results indicated that the anticipation of the emotional valence of an action-effect influenced free-choice action. Whereas the effect-tones induced a clear response bias (i.e., more acquisition-consistent than –inconsistent key-choices) if they had been associated with a positive emotional valence, this response bias was not reliable for action-effects associated with negative emotional features. In summary, the present results provide further proof for ideomotor theories of action control (James, 1890; Elsner & Hommel, 2001) which state that actions are automatically activated by anticipating their consequences.

Die Studie hat zum Ziel, im Rahmen des Äußerungsproduktionsmodells die Annahme thinking for speaking (Slobin, 1996) experimentell zu überprüfen. Ansatzpunkt ist der Unterschied in der Stellung der Markierung einer Entscheidungsfrage zwischen drei Sprachen. Während der Fragemodus im Deutschen/Polnischen vor dem Frageinhalt markiert wird, geschieht die Modusmarkierung im Chinesischen nach dem Inhalt. Um die entsprechende Satzstruktur aufzubauen, sollte der Fragemodus beim syntaktischen Kodierungsprozess (speaking) im Deutschen/Polnischen vor, im Chinesischen nach dem Inhalt verarbeitet werden. Unter Zugrundelegung der inkrementellen Äußerungsproduktion gehen wir davon aus, dass die Verarbeitungsreihenfolge beim vorangehenden Konzeptualisierungsprozess (thinking) mit der syntaktischen Kodierungsabfolge übereinstimmt: Der Fragemodus wird im Deutschen/ Polnischen vor, im Chinesischen hingegen nach dem Inhalt konzeptualisiert. Um den zeitlichen Ablauf der zwei Konzeptualisierungsprozess zu ermitteln, bedienten wir uns des lateralisierten Bereitschaftspotenzials (lateralized readiness potential, LRP) im binären Wahlreaktions-Go/Nogo-Paradigma. Im Versuch reagierten deutsche, chinesische und polnische Muttersprachler auf dargebotene Bilder mit Tastendrücken und Sprechen. Zu beobachten war das Auftreten eines LRP bei Nogo, das signalisierte, in welcher Reihenfolge sich die Handwahl und die Nogo-Entscheidung realisierten Ein Nogo-LRP trat bei allen drei Sprechergruppen auf. Zudem wurde festgestellt, dass die Nogo-Entscheidung, die erwartungsgemäß mit der sprachlichen Verarbeitung des Fragemodus verbunden wurde, bei den drei Sprechergruppen ungefähr zeitgleich getroffen wurde. Die Befunde legen nahe, dass der Fragemodus in den drei Sprachen zeitlich nicht unterschiedlich, sondern einheitlich geplant wurde. Die Schlussfolgerung wird im Rahmen von thinking for speaking sowie dem Äußerungsproduktionsmodell diskutiert. / Languages differ in the marking of the sentence mood of a polar interrogative (yes/no question). For instance, the interrogative mood is marked at the beginning of the surface structure in Polish, whereas the marker appears at the end in Chinese. In order to generate the corresponding sentence frame, the syntactic specification of the interrogative mood is early in Polish and late in Chinese. In this respect, German belongs to an interesting intermediate class. The yes/no-question is expressed by a shift of the finite verb from its final position in the underlying structure into the utterance initial position, a move affecting, hence, both the sentence final and the sentence initial constituents. The present study aimed to investigate whether during generation of the semantic structure of a polar interrogative, i.e. the processing preceding the grammatical formulation, the interrogative mood is encoded according to its position in the syntactic structure at distinctive time points in Chinese, German, and Polish. In a two-choice go/nogo experimental design, native speakers of the three languages responded to pictures by pressing buttons and producing utterances in their native language while their brain potentials were recorded. The emergence and latency of lateralized readiness potentials (LRP) in nogo conditions, in which speakers asked a yes/no question, should indicate the time point of processing the interrogative mood. The results revealed that Chinese, German, and Polish native speakers did not differ from each other in the electrophysiological indicator. The findings suggest that the semantic encoding of the interrogative mood is temporally consistent across languages despite its disparate syntactic specification. The consistent encoding may be ascribed to economic processing of interrogative moods at various sentential positions of the syntactic structures in languages or, more generally, to the overarching status of sentence mood in the semantic structure.