Les ganglions de la base forment un ensemble de structures sous-corticales connues pour leur implication dans les processus sensori-moteurs, cognitifs et motivationnels. L’objectif de ce travail était d’approfondir le rôle des neurones du noyau subthalamique (NST), des neurones de projections et interneurones cholinergiques du striatum et des neurones du globus pallidus externe (GPe) dans la mise en place et l’exécution d’un comportement dans différents contextes motivationnels. Nous nous sommes intéressés à l’influence de l’effort et de la récompense sur l’activité de ces neurones grâce à une approche comportementale associée à des enregistrements extracellulaires unitaires chez le singe éveillé. L’influence de ces facteurs a été appréhendée dans une tâche visuo-motrice dans laquelle différents niveaux d’effort et de récompense étaient imposés à l’animal. Nos résultats comportementaux ont montré une prise en compte de la valeur des stimuli par les animaux. Les résultats électrophysiologiques obtenus montrent une implication de chacune des populations étudiées dans le traitement des informations relatives à l’effort et à la récompense. Ils suggèrent un rôle des neurones du NST, du striatum et du GPe respectivement dans la mise en place, l’exécution et l’évaluation de l’action sur la base de la valeur subjective de la récompense. Nos résultats apportent des informations nouvelles sur les substrats neurophysiologiques qui sous-tendent les processus motivationnels dans la circuiterie des ganglions de la base. / The basal ganglia form a set of subcortical structures known to be involved in sensorimotor, cognitive and motivational processes. The aim of this work was to study the role of the subthalamic nucleus (STN) neurons, the cholinergic interneurons and the projection neurons of the striatum and the neurons of the external part of the globus pallidus (GPe) in the establishment and the execution of a behavior under different motivational contexts. We examined the influence of effort and reward on the activity of these neurons with a behavioral approach combined with extracellular recordings in awake monkeys. The influence of these factors has been investigated in a visuo-motor task in which different levels of effort and reward were imposed on the animal. Our behavioral results showed a consideration of the value of the visual stimuli by the animals. Electrophysiological results showed an implication of each of the neuronal populations studied in the encoding of force and reward related information. These data suggest a role of STN, striatum and GPe in the establishment, the execution and the update of the benefit of the action based on subjective reward value. Our results bring out new features on the neurophysiological substrates underlying motivational processes in basal ganglia circuitry.

L'attention est une fonction au cœur de la cognition qui, à tout moment, nous permet de sélectionner les informations pertinentes à traiter, tout en ignorant les autres. Cette sélection de l’information qui s’opère à la fois dans l'espace et dans le temps résulte de l’intégration des informations sensorielles et d’un contrôle de "haut niveau" en fonction de nos buts. Cette fonction dépend d’un réseau cérébral incluant le système fronto-pariétal et est sous l’influence de différents neuromodulateurs, en particulier la noradrénaline, dont l’action reste encore mal connue. Mon travail de thèse consistait à comprendre le rôle de la noradrénaline sur les processus attentionnels. Mes objectifs étaient d’une part de vérifier notre hypothèse selon laquelle la noradrénaline modulerait les différentes facettes de l’attention (attention spatiale et attention soutenue) et d’autre part d’élucider les mécanismes d’action par lesquelles la noradrénaline exercerait ces effets. Pour répondre à ces questions, nous avons testé l’impact d’une augmentation de la transmission noradrénergique (administration intramusculaire d'atomoxétine) chez le singe, dans des tâches comportementales nécessitant une sélection de l’information visuelle soit dans l’espace (tâche d'attention avec indice et exploration spontanée d'images) soit au cours du temps (tâche de discrimination go/nogo). Nos résultats démontrent que l’atomoxétine facilite les processus attentionnels à la fois dans l’espace et au cours du temps. Dans l’espace, l’atomoxétine module l’orientation de l’attention visuo-spatiale en fonction du contexte, en ajustant le taux d’accumulation sensorielle ou l’impact de la saillance des images sur l’orientation de l’attention. Au cours du temps, l’atomoxétine ajuste la relation entre la sensibilité à discriminer la cible parmi des distracteurs et le biais de réponse des animaux. En résumé, mes résultats démontrent que la noradrénaline influence les deux facettes, spatiale et temporelle de l’attention et suggèrent une action via un ajustement des processus de traitement de l’information sensorielle et un ajustement du contrôle de l’attention au contexte / Attention is a function at the heart of cognition that, at any given moment, enables us to select some information for further processing, while setting aside others. This selection of information that operates both in space and time, results from the integration of sensory information and higher-level control according to our goals. This function depends on a cerebral network including the fronto-parietal system. It is also under the influence of different neuromodulators, in particular norepinephrine, the action of which is still poorly understood.The aim of my PhD work was to understand the role of norepinephrine on attentional processes. My objectives were, on the one hand, to test our hypothesis that norepinephrine is capable of acting on the different facets of attention (spatial attention and sustained attention) and, on the other hand, to elucidate the mechanisms of action by which noradrenaline exerts its action. To answer these questions, we tested the impact of an increase in noradrenergic transmission (intramuscular administration of atomoxetine) in monkeys, using behavioral tasks requiring a selection of visual information in space (cued attentional task and spontaneous image exploration) or over time (go/nogo discrimination task). Our results demonstrate that atomoxetine facilitates attentional processes both in space and over time. In space, atomoxetine modulates the orientation of visuospatial attention according to the context, adjusting the rate of sensory accumulation or the impact of image saliency on attention orientation. Over time, atomoxetine adjusts the relationship between the sensitivity to discriminate a target among distractors and the animal’s response bias.In summary, my results demonstrate that norepinephrine influences both the spatial and temporal facets of attention and suggests an action through an adjustment of sensory information processing and an adjustment of attention control to the context

Extreme obstacle racing is a brand new sporting industry that is currently developing most not only in Europe but also around the world. I focused mainly on introducing this new sport in order to find out different level of the motor skills of beginners and professionals. In the theoretical part of my work, I defined what is characteristic for this sport and why it is so famous and wanted today. In the practical part, using the proposed test battery, I tested all the motor skills of beginners and professionals in order to identify the real physical differences between them. As a result of these facts, methods of testing, observation, processing, evaluation, analysis and statistics of data, along with the study of professional literature, were used. Based on the established hypotheses, I asked questions that I had in my opinions and attitudes to the beginners and professionals of this thriving sport. Through research and testing, we have found that professionals are certainly more dominant in motor skills than beginners with overall predominance in all underlying motor tests that included speed, endurance, strength and coordination tests. The benefit of this work is the analysis and introduction of a new sporting industry, including the main discovery of real differences between beginners and...

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The purpose of this thesis is to apply the adventure formulas laid out by John G. Cawelti and the dramatis personae written by Vladimir Propp to the traditional adventure books Le Tour du monde en quatre-vingts jour (1874) and Treasure Island (1883) and the interactive fictions The Secret of Monkey Island (1990) and 80 Days (2014), both in the adventure genre. After they have been applied, I will compare Le Tour du monde en quatre-vingts jour with 80 Days and Treasure Island with The Secret of Monkey Island by how the formulas and dramatis personae work and evolve in the fictions. To start things off, I will present the four works and then go through the relevant parts of Cawelti’s formula and Propp’s dramatis personae. Thereafter, I will explain what interactive fiction might be by using Espen J. Aarseth’s Cybertext. Important concepts will be clarified as well.    The analysis in this thesis starts afterwards. Le Tour du monde en quatre-vingts jour and 80 Days will be the first two works to be analysed after Cawelti and thereafter Propp. When that is done, I will conclude what I have found. The same process will be done with Treasure Island and The Secret of Monkey Island. After that, I will make a concluding comparison between 80 Days and The Secret of Monkey Island and finally conclude whether Cawelti and Propp can be applied to interactive fiction.

The movements of two vervet monkey troops were studied to determine whether they optimize their rate of food intake in relation to seasonal energy availability. The effect of variation in habitat structure on the troops’ foraging strategies while utilizing temporally and spatially distributed resources was determined. Troop home range boundaries were delineated, the various plant communities and species utilised by the troops identified and classified, and variations in home range and vegetation structure were reported. The diets of the troops were determined and compared. Effects of coexistence on competition were assessed. Vervet food trees were randomly selected, marked and seasonal phenological data collected. Samples of food items constituting the two troops diets were collected for energy analysis. Using geostatistical interpolation techniques, monthly energy values were extrapolated onto home range grids for the two vervet monkey troops. Grids were stored as database files that were interrogated through GIS simulation models. Using the stochastic processes inherent in Markov chain theory, a series of non-returning random walks were simulated for comparison to original routes taken by the two troops. Results from comparisons of home range energy, day range lengths and areas, shortest route energy to actual route energy, time spent in high energy areas, and energy utilisation from actual and randomly generated routes indicated that the two troops optimize resource energy available to them by adopting flexible foraging strategies. In environments where temporal and spatial variations in habitat structure affect the distribution of resources, it is essential that animals develop optimal foraging strategies to survive. For the two troops investigated, foraging strategies fluctuate between being time minimizers in more heterogeneous environments where resources are abundant, and energy maximisers in homogeneous environments where resources are constrained by low diversity and seasonality. / Environmental Sciences (Department) / D.Litt et Phil (Environmental Management)

Une variété de modèles sur le processus de prise de décision dans divers contextes présume que les sujets accumulent les évidences sensorielles, échantillonnent et intègrent constamment les signaux pour et contre des hypothèses alternatives. L'intégration continue jusqu'à ce que les évidences en faveur de l'une des hypothèses dépassent un seuil de critère de décision (niveau de preuve exigé pour prendre une décision). De nouveaux modèles suggèrent que ce processus de décision est plutôt dynamique; les différents paramètres peuvent varier entre les essais et même pendant l’essai plutôt que d’être un processus statique avec des paramètres qui ne changent qu’entre les blocs d’essais. Ce projet de doctorat a pour but de démontrer que les décisions concernant les mouvements d’atteinte impliquent un mécanisme d’accumulation temporelle des informations sensorielles menant à un seuil de décision. Pour ce faire, nous avons élaboré un paradigme de prise de décision basée sur un stimulus ambigu afin de voir si les neurones du cortex moteur primaire (M1), prémoteur dorsal (PMd) et préfrontal (DLPFc) démontrent des corrélats neuronaux de ce processus d’accumulation temporelle. Nous avons tout d’abord testé différentes versions de la tâche avec l’aide de sujets humains afin de développer une tâche où l’on observe le comportement idéal des sujets pour nous permettre de vérifier l’hypothèse de travail. Les données comportementales chez l’humain et les singes des temps de réaction et du pourcentage d'erreurs montrent une augmentation systématique avec l'augmentation de l'ambigüité du stimulus. Ces résultats sont cohérents avec les prédictions des modèles de diffusion, tel que confirmé par une modélisation computationnelle des données. Nous avons, par la suite, enregistré des cellules dans M1, PMd et DLPFc de 2 singes pendant qu'ils s'exécutaient à la tâche. Les neurones de M1 ne semblent pas être influencés par l'ambiguïté des stimuli mais déchargent plutôt en corrélation avec le mouvement exécuté. Les neurones du PMd codent la direction du mouvement choisi par les singes, assez rapidement après la présentation du stimulus. De plus, l’activation de plusieurs cellules du PMd est plus lente lorsque l'ambiguïté du stimulus augmente et prend plus de temps à signaler la direction de mouvement. L’activité des neurones du PMd reflète le choix de l’animal, peu importe si c’est une bonne réponse ou une erreur. Ceci supporte un rôle du PMd dans la prise de décision concernant les mouvements d’atteinte. Finalement, nous avons débuté des enregistrements dans le cortex préfrontal et les résultats présentés sont préliminaires. Les neurones du DLPFc semblent beaucoup plus influencés par les combinaisons des facteurs de couleur et de position spatiale que les neurones du PMd. Notre conclusion est que le cortex PMd est impliqué dans l'évaluation des évidences pour ou contre la position spatiale de différentes cibles potentielles mais assez indépendamment de la couleur de celles-ci. Le cortex DLPFc serait plutôt responsable du traitement des informations pour la combinaison de la couleur et de la position des cibles spatiales et du stimulus ambigu nécessaire pour faire le lien entre le stimulus ambigu et la cible correspondante. / A variety of models of the decision-making process in many different contexts suggest that subjects sample, accumulate and integrate sensory evidence for and against different alternative choices, until one of those signals exceeds a decision criterion threshold. Early models assumed that this process is static and does not change during a trial or even between trials, but only between blocks of trials when task demands such as speed versus accuracy change. However, newer models suggest that the decision-making process is dynamic and factors that influence the evidence accumulation process might change both between trials in a block and even during a trial. This thesis project aims to demonstrate that decisions about reaching movements emerge from a mechanism of integration of sensory evidence to a decision criterion threshold. We developed a paradigm for decision-making about reach direction based on ambiguous sensory input to search for neural correlates of the decision-making process in primary motor cortex (M1), premotor cortex (PMd) and dorsolateral prefrontal cortex (DLPFc). We first tested several versions of the task with human subjects before developing a task (“Choose and Go”) that showed ideal behavior from the subjects to test our hypothesis. The task required subjects to choose between two color-coded targets in different spatial locations by deciding the predominant color of a central “decision cue” that contained different amounts of colored squares of the two target colors. The strength of the evidence was manipulated by varying the relative numbers of squares of the two colors. The response times and error rates both increased in parallel as the strength of the sensory evidence in the decision cue (its color bias) became increasingly weaker. Computational modelling showed that the choice behaviour of the subjects could be captured by different variants of the drift-diffusion model for accumulation of sensory evidence to a decision threshold. We then recorded cells from M1, PMd and DLPFc in 2 macaques while they performed the task. Behavioral data showed that response times and error rates increased with the amount of ambiguity of the decision cues. M1 cells discharged in correlation with movement onset and were not influenced by the ambiguity of the decision cues. In contrast, the discharge of PMd cells increased more slowly with increased ambiguity of the decision cues and took increasingly more time to signal the movement direction chosen by the monkeys. The changes in activity reflected the monkeys’ reach choices. These data support a role for PMd in the choice of reach direction. DLPFc data are preliminary but reveal a stronger effect of the color-location conjunction rule in the neuronal discharge than in PMd. Our conclusion is that PMd is involved in the evaluation of evidence for and against different alternatives and about target spatial location independent of the color of the targets. DLPFC neurons play a greater role in processing information about the color and location of the spatial targets and decision cue to resolve the color-location conjunction rule required to decide on the reach target direction.

Le cortex cérébral primate a subi des modifications majeures pendant l'évolution qui ont permis le développement de fonctions cognitives supérieures. Un accroissement massif a eu lieu avec l'extension spécifique des couches supragranulaires et une forte expansion tangentielle. Le cortex primate ne possède pas uniquement davantage de neurones, comparé au rongeur, mais aussi des différences qualitatives. Ceci suggère des différences qualitatives pendant le développement du cortex.Une zone proliférative corticale supplémentaire a été identifiée chez le singe macaque: la zone subventriculaire externe (OSVZ) supposée être impliquée dans l'expansion du cortex primate. Mais les propriétés des précurseurs de l'OSVZ restent mal connues. Des techniques de microscopie en temps réel et d'immunofluorescence ont permis de réaliser une description exhaustive des précurseurs de l'OSVZ et de leurs propriétés chez le singe macaque.Nos résultats mettent en évidence des différences primates/rongeurs majeures. Les observations en temps réel révèlent des capacités prolifératives bien plus importantes des précurseurs. Les précurseurs primates de l'OSVZ présentent des taux de prolifération variables pendant la corticogenèse liés à la cinétique du cycle cellulaire. Nos enregistrements ont permis la génération d'une grande base de données de propriétés et lignages de précurseurs et la mise en évidence d’une diversité morphologique inattendue. 5 types ont été identifiés. Impliqués dans des lignages complexes, chaque type a la capacité de s'auto-renouveler et de générer directement des neurones. Parallèlement, nous avons développé une méthode de classification non supervisée des précurseurs corticaux. Cette technique a identifié les mêmes 5 types de précurseurs.Les résultats de cette thèse apportent de nouveaux éléments dans la compréhension des spécificités de la corticogenèse primate qui contribuent à l'expansion corticale et au développement de capacités cognitives supérieures. / The primate cerebral cortex underwent major modifications during evolution that enabled the development of high cognitive functions. A massive enlargement occurred with the specific expansion of the supra granular layers and the apparition of new frontal areas. Not only quantitative differences are found compared to the rodent but also qualitative differences. This points to potential qualitative differences in primate cortical development. An extra proliferating zone had already been identified during macaque corticogenesis: the outer subventricular zone (OSVZ). This zone is assumed to play a key role in the expansion of the primate cortex but the cellular and functional properties of OSVZ precursors remain elusive. We used quantitative long-term time-lapse video-microscopy (TLV) and immunofluorescence in and ex vivo to perform a detailed and exhaustive description of OSVZ precursor types and proliferative abilities at different stages of macaque cortical development. Our results highlight major rodent/primate differences. TLV observations revealed a much higher proliferative potential of OSVZ compared to the rodent SVZ. We report variable rates of proliferation linked to cell-cycle duration in a stage-specific manner. TLV recordings allowed the formation of a large database of primate precursor properties and lineages. This dataset unravelled an unexpectedly high diversity of OSVZ precursor morphologies. Five precursor types were identified. Involved in complex lineages, each precursor type can self-renew and directly generate neurons. In a parallel approach, we developed an unbiased clustering tool to automatically classify cortical precursors. This technique returned the same five precursor types as the morphological categorization. The results of this PhD thesis provide new insights into primate specificities during corticogenesis that contribute to cortical expansion and to the development of higher cognitive abilities.

Primatas da subespécie A. g. clamitans, popular bugio, possuem um dimorfismo sexual evidenciado na fase adulta com machos de cor ruiva e fêmeas de cor castanho com nuanças avermelhadas. Em estudos de cativeiro com esta subespécie descobriu-se uma secreção epidérmica avermelhada, semelhante à cor da pelagem dos MA. A partir desta constatação diferentes hipóteses têm sido levantadas sobre a função e a origem desta secreção. Assim, o presente estudo objetivou: 1- Analisar se esta secreção é responsável pela coloração do pêlo dos animais e se a água é capaz de descolorir os pêlos 2- Verificar se a liberação de secreção sofre influencia das temperaturas corpórea e ambiente; 3- Identificar através de microscopia óptica e eletrônica as características morfológicas das glândulas produtoras de secreção, 4- Mapear as regiões do corpo dos animais que possuem tal glândula, 5- Definir qual sexo e faixa etária possui a glândula; 6- Determinar se a secreção é capaz de corar o pêlo mesmo após estocada a -4C; 7- Avaliar a cor dos campos cromatogênicos, durante um ano em animais adultos, sub adultos e juvenis; 8- Analisar a composição da secreção; 9- Dosar o hormônio testosterona em animais de diferentes sexo e faixas etárias e 10- Verificar uma possível relação entre comportamento social em bugios silvestres e a cor observada. Constatou-se que MA, MSA, MJ , FA e FSA liberam secreção. A liberação em quase todas as regiões do corpo sofre influência da temperatura corpórea, diferindo apenas na região do hióide em MA e da mandíbula em MSA, regiões que a temperatura corpórea possuem maior influência. Verificou-se que pêlos de MSA quando mantidos em estufa, com a secreção fresca liberada por este animal, muda de cor, de escuro para ruivo, igualando a cor do pêlo de MA; uma vez pigmentado a água não muda a cor do pêlo,. A secreção estocada muda apenas à cor da região central do pêlo. Identificou-se a GPP (glândula produtora de secreção colorida), na região do osso hióide e mandíbula, em MA e MSA. Em FA e FSA identificou-se estrutura glandular menos desenvolvida, denominada de GPPi (glândula produtora de pigmento em fase intermediária) nas regiões do hióide, mandíbula, esterno e região inguinal. As GPPi estão também presentes na região inguinal e nuca de MA e MSA. Os animais juvenis e infantes apresentam somente glândulas sudoríparas normais em todas as regiões do corpo. Na secreção colorida não existem carotenóides. As secreções de MA, FA, MAS e MJ possuem diferentes concentrações de ferro. A secreção de MA apresentou maior concentração de ferro quando comparada às secreções dos animais de outro sexo e/ou idade. A microscopia eletrônica confirmou a característica secretora da GPP e evidenciou estruturas cristalinas dentro das células do ácino semelhantes as inclusões de ferro observadas no citoplasma de células de outros organismos. Os animais que sofreram maior variação de cor nos campos cromatogênicos foram os MSA, e foram os animais que tiveram maiores índices de esfregação, sugerindo um mecanismo de espalhamento da secreção Os níveis de testosterona, foram proporcionais ao sexo e idade dos animais, sendo os maiores valores encontrados em MA. O estudo do comportamento social mostrou que os MA ruivos possuem um elevado status hierárquico e desempenham um papel de guardião do grupo. Os resultados indicam que a secreção colorida, liberada na epiderme de A. g. clamitans, são as responsáveis pela coloração do pêlo de MA devido à concentração de ferro. Um cromóforo que contém ferro seria um dos agentes responsáveis pela coloração. A diferença de cor nesta subespécie é uma característica sexual secundária, e parece ocorrer por um mecanismo ativacional do hormônio testosterona, ativando o desenvolvimento das glândulas GPP. O nível de testosterona, a presença de GPP e a coloração ruiva intensa possuem uma forte correlação com o status hierárquico de MA. / Adult brown howler monkeys, Alouatta guariba clamitans, have a clear sexual dimorphism, with red-haired males and chestnut females with red nuances. Captivity studies with this subspecies revealed an epidermic secretion of red coloration, similar to the coat color of adult males. Different hypotheses have been proposed on the function and origin of this secretion. The present work aimed to: 1 analyze if fresh secretion is responsible for hair coloration in this animals and if water is capable to decolorize hair; 2 verify if secretion release is affected by body and atmospheric temperatures; 3 identify by optic and electronic microscopy the morphologic characteristics of the secretion-producing glands; 4 map the areas of the animal body that have such glands; 5 define which sex-age categories have this gland; 6 determine if secretion is able to color hair after stored at -4º C; 7 evaluate during one year the coloration of cromatogenic fields in adult, subadults and juvenile animals; 8 analyze the secretion composition; 9 measure testosterone levels in animals of different sex-age categories; 10 verify the possible relationship between the social behavior of wild animals and the observed coloration. It was found that adult, subadult and juvenile males and adult and subadult females release secretion. The release in almost all body areas is under influence of the corporal temperature that is greater in adult male hyoid and subadult male jaw areas. The hair of subadult males of A. g. clamitans maintained in a stove treated with fresh secretion changes from dark to reddish color, similar to the adult males hair color. The water does not change the color of the hair, once pigmented. The stored secretion changes only the color of the central area of hair. The gland producing colored secretion (GPP) was identified in the area of the hyoid bone and jaw in adult and subadult males. In adult and subadult females a less developed glandular structure was found, denominated GPPi in the hyoid, jaw, breastbone and inguinal areas. GPPi are also present in the inguinal area and nape of adult and subadult males. The juvenile and infant animals presented only normal sweat glands in all body areas. There are no carotenoids in the colored secretion. The secretions of adult, subadult and juvenile males and adult females have different concentrations of iron and adult males presented larger concentration of iron in their secretions when compared to other sex and age categories. The electronic microscopy confirmed the secretory features of GPP and evidenced crystalline structures inside the acinus cells that resemble iron-containing structures evidenced in other cellular structures. Subadult males suffered the greater degree in color variation in cromatogenic fields, and they were also the animals that presented larger rubbing index, suggesting that this would be the mechanism of secretion dispersal. The testosterone levels were proportional to the sex and age categories of the animals, with the largest values found in adult males. The study of the social behavior showed that the reddish adult males have a superior hierarchic status and play the role of group guardians. Our results indicate that the colored secretions, released in the epidermis of A. g. clamitans, are responsible for the hair coloration in adult males due to iron concentration. An iron-containing chromophore would be one of the agents promoting the coloration. The color difference in this subspecies is a secondary sexual trait that seems to occur through a testosterone-activated mechanism, promoting the development of GPP glands. Testosterone levels, the presences of GPP and the intense red-haired coloration have a strong correlation with the hierarchical status of adult males.