La présente étude avait pour but d’explorer les modulations fonctionnelles putaminales du signal de spectroscopie par résonance magnétique (SRM) combiné du glutamate et de la glutamine (Glx), ainsi que de l’acide γ-aminobutyrique (GABA) en lien avec l’apprentissage d’une séquence motrice. Nous avons émis l’hypothèse que les concentrations de Glx seraient spécifiquement augmentées pendant et après la pratique d’une telle tâche, et ce comparativement à une condition d’exécution motrice simple conçue pour minimiser l’apprentissage. La tâche d’appuis séquentiels des doigts (« finger taping task ») utilisée est connue pour induire un apprentissage moteur évoluant en phases, avec une progression initialement rapide lors de la première session d’entraînement (phase rapide), puis lente lors de sessions subséquentes (phase lente). Cet apprentissage est également conçu comme dépendant de processus « on-line » (pendant la pratique) d’acquisition et « off-line » (entre les périodes de pratique) de consolidation de la trace mnésique de l’habilité motrice. Une grande quantité de données impliquent le système de neurotransmission glutamatergique, principalement par l’action de ses récepteurs N-Méthyl-D-aspartate (NMDAR) et métabotropiques (mGluR), dans une multitude de domaine de la mémoire. Quelques-unes de ces études suggèrent que cette relation s’applique aussi à des mémoires de type motrice ou dépendante du striatum. De plus, certains travaux chez l’animal montrent qu’une hausse des concentrations de glutamate et de glutamine peut être associée à l’acquisition et/ou consolidation d’une trace mnésique. Nos mesures de SRM à 3.0 Tesla, dont la qualité ne s’est avérée satisfaisante que pour le Glx, démontrent qu’une telle modulation des concentrations de Glx est effectivement détectable dans le putamen après la performance d’une tâche motrice. Elles ne nous permettent toutefois pas de dissocier cet effet putativement attribuable à la plasticité du putamen associée à l’apprentissage moteur de séquence, de celui de la simple activation neuronale causée par l’exécution motrice. L’interprétation de l’interaction non significative, montrant une plus grande modulation par la tâche motrice simple, mène cependant à l’hypothèse alternative que la plasticité glutamatergique détectée est potentiellement plus spécifique à la phase lente de l’apprentissage, suggérant qu’une seconde expérience ainsi orientée et utilisant une méthode de SRM plus sensible au Glx aurait donc de meilleures chances d’offrir des résultats concluants. / The present study explored motor learning-related functional changes in putaminal combined glutamate and glutamine (Glx) and γ-Aminobutyric acid (GABA) magnetic resonance spectroscopy (MRS) signal. It was hypothesized that Glx concentrations would specifically increase during and after learning of a sequential finger tapping task (sFTT), as compared to execution of a simple motor task designed to elicit minimal learning. Learning of sFTT is known to evolve in an initial fast progressing stage during the first practice session (fast learning stage), followed by a slower progression during later sessions (slow learning stage). It is also thought to depend on both on-line (during practice sessions) acquisition and off-line (between practice sessions) consolidation processes to create, transform and assure retention of a motor skill memory trace. A body of data implicates glutamatergic neurotransmission, especially through its N-Methyl-D-aspartate (NMDAR) and metabotropic (mGluR) receptors, in many memory systems, some of which apply to motor learning and striatal-dependant learning. Moreover, some animal studies suggest that Glx concentrations can be upregulated in relation to memory acquisition and/or consolidation. Our MRS acquisitions, of which the quality happened to be sufficient only for Glx quantification, allowed the detection of an augmentation in putaminal Glx occurring after motor task execution. However, our data could not ascribe this modulation specifically to motor learning related plastic changes, at the exclusion of simple neural activation related to motor execution. Nevertheless, the interpretation of the non-significant interaction, showing a larger Glx change in response to the simple motor task compared to sFTT, leads to the possibility that the detected glutamatergic plasticity may be specifically associated to the slow learning phase. We therefore suggest that testing this alternate hypothesis in a second experiment, using an MRS technique with more sensibility to Glx could yield more convincing results.

Le feedback auditif se définit 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 influence sur l'apprentissage moteur est moins clair. Cette thèse a pour but d'examiner l'influence 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 bénéficient de l'information temporelle contenue dans le feedback auditif et qu'ils sont sensibles aux distortions de cette 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 benéficié 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 permettent 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 cette 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.

Using Hebb’s (1961) paradigm, it has been shown that older adults (OAs) fail to learn recurrent visuospatial supra-span sequence information (Turcotte, Gagnon, & Poirier, 2005); a deficit which has not been demonstrated on verbal versions of the same task or in younger adults (YAs). Since the Hebb paradigm is thought to rely on working memory and thus attention (Conway & Engle, 1996), one interpretation concerns an OA’s capacity to allocate the necessary attentional resources to carry out the various components of the task. Five studies investigated this proposal. The first three (Article 1) examined attention in a general manner by reducing the amount of attentional resources that a YA could devote to carrying out the visuospatial Hebb supra-span sequence learning task through the implementation of a verbal dual task (DT) procedure. The fourth (Article 2) further investigated the role of attention by using a DT induced at retrieval that overlapped extensively with the requirements (spatial and response features) of the visuospatial Hebb task. The final study (Article 3) aimed to use our previous findings to demonstrate learning among OAs in a visuospatial Hebb learning paradigm in which the motor response was replaced by a verbal response. Our findings confirm that attentional resources employed at the retrieval phase of the task appear to be particularly important for the demonstration of visuospatial sequence learning. The inclusion of a spatial and motor based DT at recall eliminated learning of the repeated sequence in YAs. Interestingly, the learning deficit of OAs was partially eliminated when the motor and spatial requirements at retrieval were reduced. Our findings offer strong support to the contention that supra-span learning of the Hebb type is not altered by the effect of age. However, learning deficits can be observed among OAs when the retrieval component of the task overly taxes attention-related processes. In the case of the visuospatial sequences, the basis of the deficit likely concerns an individual’s capacity to discriminate between responses made to previously presented sequences versus those that need to be made in reaction to the just seen sequence.

Prever eventos ambientais, com base em memórias sobre regularidades passadas, é uma das funções fundamentais de sistemas nervosos complexos. Eventos ordenados serialmente ou sequências estruturadas de estímulos permitem extrair informação passível de descrição formal que define seu padrão serial. Esse padrão inclui informações temporais e espaciais que facultam prever os próximos eventos da sequência, possibilitando a preparação prévia do organismo para lidar com sua ocorrência. Não surpreende que animais, incluindo o ser humano, aprendam, de maneira relativamente rápida, sobre regras e estruturas de padrões sequenciais de estímulos. O uso de tarefas de tempo de reação serial (TRS) é recorrente em estudos envolvendo a formação de associações, antecipação, atenção, as bases da memória e aprendizagem de relações complexas. Resumidamente, voluntários devem responder a estímulos apresentados em sequências repetitivas ou aleatórias. Com o treino, há redução no tempo de reação a cada estímulo, refletindo a aprendizagem de relações percepto-motoras. Essa redução, porém, é maior na sequência repetitiva em relação à sequência aleatória, indicando um aprendizado também sobre a sequência repetitiva, mesmo quando o voluntário não a percebe (conscientemente) e seja incapaz de relatar sua existência. Trata-se, portanto, de uma aquisição (inicialmente) implícita. A complexidade de uma sequência de estímulos pode ser expressa quantitativamente por meio de uma ferramenta matemática proposta por Shannon (1948), a entropia informacional (EI), que considera, entre outras coisas, a probabilidade de ocorrência dos estímulos em diferentes níveis. No presente trabalho, avaliamos em que extensão o conceito de EI permite prever o desempenho de ratos na tarefa de TRS envolvendo sequências com diferentes níveis de complexidade. Ratos foram treinados a reagir (1) a uma sequência repetitiva de estímulos, cuja quantidade de EI no nível 1 (que relaciona os estímulos da sequência 2 a 2) era 2,75. Após atingirem um nível assintótico de desempenho, os animais foram expostos (2) a sequências variáveis de estímulos com a mesma quantidade de EI no nível 1, porém, com maior quantidade de EI no nível 2 (que relaciona os estímulos da sequência 3 a 3). Numa etapa posterior os animais foram expostos (3) a uma nova sequência repetitiva de estímulos, cuja quantidade de EI no nível 1 era 3,00; por fim, os animais foram submetidos (4) a sequências variáveis com a mesma quantidade de EI no nível 1em relação à sequência anterior, porém, com maior quantidade de EI no nível 2 . Os resultados mostraram que os ratos aprenderam sobre os padrões seriais e, mais interessante, que seu desempenho esteve fortemente correlacionado à quantidade de EI no nível 2. Em outras palavras, quanto maior a EI, pior o desempenho dos animais tanto em termos do tempo de reação como em termos da percentagem de respostas corretas. Portanto, o conceito de EI permite não apenas quantificar a complexidade de sequências empregadas em estudos envolvendo aprendizagem serial, mas também prever o desempenho dos animais. / Prediction of environmental events, relying on memories of past regularities, is one of the fundamental functions of complex nervous systems. Sequences of serially ordered stimuli allow extracting information that defines its serial pattern. These patterns allow prediction of the next item in a sequence of events, facultating previous preparation to deal with its occurrence. Not surprisingly, animals, including humans, can identify rules present in serial structures of stimuli. Serial reaction time tasks (SRTT) have been extensively used in studies involving association, anticipation, attention, and learning and memory. Typically, subjects have to react to stimuli presented either in random or in repetitive sequences. As training proceeds, reaction time to each stimulus decreases, reflecting acquisition of this perceptual-motor skill. However, reaction time reduction is greater for repetitive sequences relative to the random sequences, indicating acquisition about the repetitive structure of the sequence. In human beings, this may occur even when the subject in uncapable of reporting the existence of a sequence, indicating that the acquisition was (at least initially) implicit rather than explicit. The complexity of a sequence of stimuli, at different levels, may be quantifyed by means of a mathematical tool proposed by Shannon (1948), the information entropy (IE). In this study we evaluated to which extent IE can predict performance of rats in SRTT involving sequences of stimuli organized at different levels of complexity. Rats were trained to react (1) a repeated sequence of stimuli which IE at the level \"1\" (i.e., expressing to which extent a given item allow prediction of the next) was 2.75. After reaching an asymptotic level of performance, the animals were exposed (2) a variable sequence of stimuli with the same amount of IE in the level \"1\", but with more IE in the level \"2\" (i.e., expressing to which extent two given items allow prediction of the next). Later the animals were exposed to (3) a new repeated sequence of stimuli, which IE at the level \"1\" was 3.00. Finally, the animals were submitted to (4) a random sequence of stimuli with the same amount of IE at the level \"1\", i.e., 3.00, but with greater IE in level 2. Results showed that rats learned about the serial patterns and, more interestingly, their performance strongly correlated to the amount of IE at the level \"2 \", both in terms of reaction times and in terms of percentage of correct responses. Therefore, IE allows not only to quantify complexity of sequences in studies involving serial learning, but also to predict performance of the subjects.

Using Hebb’s (1961) paradigm, it has been shown that older adults (OAs) fail to learn recurrent visuospatial supra-span sequence information (Turcotte, Gagnon, & Poirier, 2005); a deficit which has not been demonstrated on verbal versions of the same task or in younger adults (YAs). Since the Hebb paradigm is thought to rely on working memory and thus attention (Conway & Engle, 1996), one interpretation concerns an OA’s capacity to allocate the necessary attentional resources to carry out the various components of the task. Five studies investigated this proposal. The first three (Article 1) examined attention in a general manner by reducing the amount of attentional resources that a YA could devote to carrying out the visuospatial Hebb supra-span sequence learning task through the implementation of a verbal dual task (DT) procedure. The fourth (Article 2) further investigated the role of attention by using a DT induced at retrieval that overlapped extensively with the requirements (spatial and response features) of the visuospatial Hebb task. The final study (Article 3) aimed to use our previous findings to demonstrate learning among OAs in a visuospatial Hebb learning paradigm in which the motor response was replaced by a verbal response. Our findings confirm that attentional resources employed at the retrieval phase of the task appear to be particularly important for the demonstration of visuospatial sequence learning. The inclusion of a spatial and motor based DT at recall eliminated learning of the repeated sequence in YAs. Interestingly, the learning deficit of OAs was partially eliminated when the motor and spatial requirements at retrieval were reduced. Our findings offer strong support to the contention that supra-span learning of the Hebb type is not altered by the effect of age. However, learning deficits can be observed among OAs when the retrieval component of the task overly taxes attention-related processes. In the case of the visuospatial sequences, the basis of the deficit likely concerns an individual’s capacity to discriminate between responses made to previously presented sequences versus those that need to be made in reaction to the just seen sequence.

La navigation spatiale est une fonction complexe qui nécessite de combiner des informations sur l’environnement et notre mouvement propre pour construire une représentation du monde et trouver le chemin le plus direct vers notre but. Cette intégration multimodale suggère qu’un large réseau de structures corticales et sous-corticales interagit avec l’hippocampe, structure clé de la navigation. Je me suis concentrée chez la souris sur la navigation de type séquence (ou stratégie égocentrique séquentielle) qui repose sur l’organisation temporelle de mouvements associés à des points de choix spatialement distincts. Après avoir montré que l’apprentissage de cette navigation de type séquence nécessitait l’hippocampe et le striatum dorso-médian, nous avons caractérisé le réseau fonctionnel la sous-tendant en combinant de l’imagerie Fos, de l’analyse de connectivité fonctionnelle et une approche computationnelle. Les réseaux fonctionnels changent au cours de l’apprentissage. Lors de la phase précoce, le réseau impliqué comprend un ensemble de régions cortico-striatales fortement corrélées. L’hippocampe était activé ainsi que des structures impliquées dans le traitement d’informations de mouvement propre (cervelet), dans la manipulation de représentations mentales de l’espace (cortex rétrosplénial, pariétal, entorhinal) et dans la planification de trajectoires dirigées vers un but (boucle cortex préfrontal-ganglions de la base). Le réseau de la phase tardive est caractérisé par l’apparition d’activations coordonnées de l’hippocampe et du cervelet avec le reste du réseau. Parallèlement, nous avons testé si l’intégration de chemin, de l’apprentissage par renforcement basé modèle ou non-basé modèle pouvaient reproduire le comportement des souris. Seul un apprentissage par renforcement non-basé modèle auquel une mémoire rétrospective était ajoutée pouvait reproduire les dynamiques d’apprentissage à l’échelle du groupe ainsi que la variabilité individuelle. Ces résultats suggèrent qu’un modèle d’apprentissage par renforcement suffit à l’apprentissage de la navigation de type séquence et que l’ensemble des structures que cet apprentissage requiert adaptent leurs interactions fonctionnelles au cours de l’apprentissage. / Spatial navigation is a complex function requiring the combination of external and self-motion cues to build a coherent representation of the external world and drive optimal behaviour directed towards a goal. This multimodal integration suggests that a large network of cortical and subcortical structures interacts with the hippocampus, a key structure in navigation. I have studied navigation in mice through this global approach and have focused on one particular type of navigation, which consists in remembering a sequence of turns, named sequence-based navigation or sequential egocentric strategy. This navigation specifically relies on the temporal organization of movements at spatially distinct choice points. We first showed that sequence-based navigation learning required the hippocampus and the dorsomedial striatum. Our aim was to identify the functional network underlying sequence-based navigation using Fos imaging and computational approaches. The functional networks dynamically changed across early and late learning stages. The early stage network was dominated by a highly inter-connected cortico-striatal cluster. The hippocampus was activated alongside structures known to be involved in self-motion processing (cerebellar cortices), in mental representation of space manipulations (retrosplenial, parietal, entorhinal cortices) and in goal-directed path planning (prefrontal-basal ganglia loop). The late stage was characterized by the emergence of correlated activity between the hippocampus, the cerebellum and the cortico-striatal structures. Conjointly, we explored whether path integration, model-based or model-free reinforcement learning algorithms could explain mice’s learning dynamics. Only the model-free system, as long as a retrospective memory component was added to it, was able to reproduce both the group learning dynamics and the individual variability observed in the mice. These results suggest that a unique model-free reinforcement learning algorithm was sufficient to learn sequence-based navigation and that the multiple structures this learning required adapted their functional interactions across learning.

Task-oriented chatbots can be used to automate a specific task, such as finding a restaurant and making a reservation. Implementing such a conversational system can be difficult, requiring domain knowledge and handcrafted rules. The focus of this thesis was to evaluate the possibility of using a neural network-based model to create an end-to-end trainable chatbot that can automate a restaurant reservation service. For this purpose, a sequence-to-sequence model was implemented and trained on dialog data. The strengths and limitations of the system were evaluated and the prediction accuracy of the system was compared against several baselines. With our relatively simple model, we were able to achieve results comparable to the most advanced baseline model. The evaluation has shown some promising strengths of the system but also significant flaws that cannot be overlooked. The current model cannot be used as a standalone system to successfully conduct full conversations with the goal of making a restaurant reservation. The review has, however, contributed with a thorough examination of the current system, and shown where future work ought to be focused. / Chatbotar kan användas för att automatisera enkla uppgifter, som att hitta en restaurang och boka ett bord. Att skapa ett sådant konversationssystem kan dock vara svårt, tidskrävande, och kräva mycket domänkunskap. I denna uppsats undersöks om det är möjligt att använda ett neuralt nätverk för att skapa en chatbot som kan lära sig att automatisera en tjänst som hjälper användaren hitta en restaurang och boka ett bord. För att undersöka detta implementerades en så kallad ``sequence-to-sequence''-modell som sedan tränades på domänspecifik dialogdata. Systemets styrkor och svagheter utvärderades och dess förmåga att generera korrekta svar jämfördes med flera andra modeller. Vår relativt enkla modell uppnådde liknande resultat som den mest avancerade av de andra modellerna. Resultaten visar modellens styrkor, men påvisar även signifikanta brister. Dessa brister gör att systemet, i sig självt, inte kan användas för att skapa en chatbot som kan hjälpa en användare att hitta en passande restaurang. Utvärderingen har dock bidragit med en grundlig undersökning av vilka fel som görs, vilket kan underlätta framtida arbete inom området.

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Dyslexi är en form av läs- och skrivsvårighet som inte kan förklaras av t.ex. bristfällig undervisning, låg begåvning eller synproblem. Orsaken till dyslexi anses vanligen vara nedsatt förmåga av fonologiskt processande. Flera studier har dock visat att dyslexi även är förknippat med språkliga, kognitiva och motoriska svårigheter som inte kan förklaras av nedsatt förmåga av fonologiskt processande. Det procedurella minnessystemet engageras vid inlärning, processande och konsolidering av kognitiva och motoriska färdigheter. Enligt the procedural deficit hypothesis (PDH) orsakas de svårigheter förekommande vid dyslexi av en nedsättning i detta minnessystem. Forskningen inom området är inte entydig då det har rapporterats om både intakt och nedsatt procedurellt minne hos personer med dyslexi. Föreliggande studie var den första att undersöka procedurellt minne hos svenska universitetsstuderande med dyslexi. Syftet var att testa PDH med hjälp av alternating serial reaction time task (ASRT), samt att undersöka relationen mellan procedurellt minne, närmare bestämt implicit sekvensinlärning, och läsförmåga samt aspekter av språklig/fonologisk förmåga. I studien deltog totalt 37 universitetsstuderande, 11 med dyslexi och 26 utan dyslexi. Resultatet indikerade att förmågan till inlärning och konsolidering i det procedurella minnet inte skilde sig åt mellan grupperna. Resultatet ger således inte stöd åt PDH. Vidare kunde inga signifikanta korrelationer påvisas, vilket tyder på att det inte föreligger något samband mellan procedurellt minne och respektive läsförmåga, fonologisk medvetenhet och snabb benämning (rapid automatized naming, RAN). / Developmental dyslexia is characterized by reading and writing difficulties that do not derive from poor education, low intelligence or visual impairment. The disorder is commonly explained as a result of an underlying phonological deficit. However, studies have shown that dyslexia is also associated with linguistic, cognitive, and motor impairments that can not be explained by a specific phonological deficit. The procedural memory underlies learning, processing and consolidation of motor and cognitive skills. The procedural deficit hypothesis (PDH) suggests that an impairment of procedural memory is the underlying cause of developmental dyslexia. Previous research has yielded inconsistent results as studies have reported intact as well as impaired procedural memory in dyslexics. This study was the first to examine procedural memory in Swedish university students. The aim was to test the PDH, by using an alternating serial reaction time task (ASRT), and to examine the correlation between procedural memory and reading ability as well as aspects of linguistic/phonological ability. In this study there was a total of 37 university students, 11 dyslexics and 26 nondyslexics. The result showed learning and consolidation of the procedural memory in both groups and there was no significant difference between the groups. Thus, the results do not support the PDH. No significant correlations were found, which indicates that there is no association between procedural memory and reading ability, phonological awareness or rapid automatized naming (RAN).