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Visual attention can be voluntarily focused on a location or automatically attracted by salient features in a visual scene. Studies using functional Magnetic Resonance Imaging (fMRI) suggest two networks of visual attention involved in these complementary mechanisms: a dorsal frontoparietal network and a ventral frontoparietal network of visuospatial attention respectively. However, most studies so far have applied non‐natural schematic stimuli. The present study investigates visual attention in images of natural environmental scenes. Adopting previously used eye‐tracker paradigms, we study the influence of task instruction and luminance contrast modifications in pictures on both eye movements and neural activity using Eye‐Tracking and functional Magnetic Resonance Imaging simultaneously. We expect increased top‐down control of attention in a search task compared to a free viewing condition visible in enhanced neural activation in the intraparietal sulcus (IPS) as part of the dorsal frontoparietal network. Strong modifications of luminance contrast should foster bottom‐up processing activating the temporoparietal junction (TPJ) a crucial area in the ventral frontoparietal network of visual attention. Although the obtained eye‐tracking data shows the expected shift of fixations towards locations of increased luminance contrast, we do not find an influence of luminance contrast modifications on neural processing. Comparison of instructions reveals diverse results across participants possibly due to the long presentation duration of stimuli which allowed participant’s attention to wander independently of task instruction. We find bilateral activation in IPS and parahippocampal place area (PPA) as well as bilateral deactivation in the TPJ region independent of task context. This might indicate similar contributions of these areas to free viewing of and search in visual scenes. However, dissociation of target detection and attention during search by deconvolution analysis of data obtained in this study might reveal a more detailed picture of functional involvement of the IPS and TPJ region in processes of visual attention. Remarkably, results show robust activation of the PPA in both tasks, suggesting that the PPA region might not only be activated by houses and open scenes but by narrow scenes (bushes, leaves) of natural outdoor environment as well.

Visual attention can be voluntarily focused on a location or automatically attracted by salient features in a visual scene. Studies using functional Magnetic Resonance Imaging (fMRI) suggest two networks of visual attention involved in these complementary mechanisms: a dorsal frontoparietal network and a ventral frontoparietal network of visuospatial attention respectively. However, most studies so far have applied non‐natural schematic stimuli. The present study investigates visual attention in images of natural environmental scenes. Adopting previously used eye‐tracker paradigms, we study the influence of task instruction and luminance contrast modifications in pictures on both eye movements and neural activity using Eye‐Tracking and functional Magnetic Resonance Imaging simultaneously. We expect increased top‐down control of attention in a search task compared to a free viewing condition visible in enhanced neural activation in the intraparietal sulcus (IPS) as part of the dorsal frontoparietal network. Strong modifications of luminance contrast should foster bottom‐up processing activating the temporoparietal junction (TPJ) a crucial area in the ventral frontoparietal network of visual attention. Although the obtained eye‐tracking data shows the expected shift of fixations towards locations of increased luminance contrast, we do not find an influence of luminance contrast modifications on neural processing. Comparison of instructions reveals diverse results across participants possibly due to the long presentation duration of stimuli which allowed participant’s attention to wander independently of task instruction. We find bilateral activation in IPS and parahippocampal place area (PPA) as well as bilateral deactivation in the TPJ region independent of task context. This might indicate similar contributions of these areas to free viewing of and search in visual scenes. However, dissociation of target detection and attention during search by deconvolution analysis of data obtained in this study might reveal a more detailed picture of functional involvement of the IPS and TPJ region in processes of visual attention. Remarkably, results show robust activation of the PPA in both tasks, suggesting that the PPA region might not only be activated by houses and open scenes but by narrow scenes (bushes, leaves) of natural outdoor environment as well.

Dans notre environnement, nous sommes constamment exposés à de multiples stimuli sensoriels que notre cerveau doit analyser. Afin d'interagir avec le monde qui nous entoure, nous devons intégrer ces différentes sources d'informations sensorielles. L'étude des processus d'intégration multisensorielle est essentielle pour comprendre comment le cerveau intègre les éléments séparés d'un objet défini par plusieurs composantes sensorielles pour former un percept unifié. Il est maintenant couramment admis que la présentation conjointe de plusieurs informations sensorielles de modalités différentes d'un même stimulus peut faciliter la perception. Cette facilitation multisensorielle semble être soumise à des règles particulières puisque certains facteurs l'influencent. Parmi eux, nous avons étudié, dans notre première étude, l'impact de trois facteurs que sont la saillance, la congruence sémantique et le changement de modalité sur les performances de détection de stimuli naturels chez l'homme et le singe. L'utilisation de stimuli naturels nous a permis de mettre en lumière l'influence des paramètres physiques des stimuli sur l'intégration multisensorielle. De plus, nous avons montré que les effets de ces facteurs sur des stimuli naturels diffèrent de ceux retrouvés avec des stimuli simples. Ces résultats convergent vers des effets multifactoriels sur la facilitation multisensorielle dont la force, les interdépendances et l'ordre varieraient en fonction de la tâche comportementale et de ce fait, de la charge cognitive. D'un point de vue anatomique et plus précisément au niveau cortical, les processus d'intégration multisensorielle paraissaient être jusqu'à récemment une caractéristique que seules possédaient les aires associatives situées au sommet de la hiérarchie du traitement de l'information. On sait maintenant que des aires corticales de bas niveau, pensées jusque-là comme étant unisensorielles, sont impliquées dans les processus multisensoriels, soulevant ainsi la question des aires sous-corticales. Des études anatomiques ont mis en évidence l'existence de noyaux thalamiques qui, par leurs connexions, pourraient permettre un transfert rapide et même une intégration des informations sensorielles. Cette nouvelle littérature témoigne de la grande complexité des réseaux cérébraux de la multisensorialité. Dans deux études électrophysiologiques chez le singe, nous avons examiné les propriétés multisensorielles de deux structures, le gyrus cingulaire postérieur et le pulvinar médian, qui n'avaient jamais été explorées auparavant dans un contexte multisensoriel. Nous avons montré que ces structures sont non seulement multisensorielles mais également intégratives et qu'elles pourraient appartenir à un même système fonctionnel. Ces travaux de thèse ont apporté des éléments supplémentaires quant à notre compréhension des processus d'intégration multisensorielle au niveau comportemental et des réseaux cérébraux sous-jacents et particulièrement ceux liés à l'intégration de stimuli naturels. / In our environment, we are constantly exposed to multiple sensory stimuli that our brain has to analyze. To interact with the surrounding world, we have to integrate these different sources of sensory information. The study of the processes of multisensory integration are essential in understanding how our brain integrates the individual parts of an object defined by several sensory components to arrive at a unified percept. It is now widely accepted that the concurrent presentation of several sensory information about the same stimulus in different modalities can facilitate its perception. This multisensory facilitation seems to be subjected to specific rules since some factors influence it. Amongst them, we have studied, in our first experiment, the impact of three factors, namely saliency, semantic congruency and modality switch, on the detection of natural stimuli in humans and monkeys. Using natural stimuli enabled us to highlight the influence of the physical parameters of stimuli on multisensory integration. Moreover, we showed that the effect of these factors on natural stimuli are different from those found with simple stimuli. These results point toward multifactorial effects on multisensory facilitation, of which the force, the interdependency and the order would vary as a function of the behavioral task, and, thus as a function of the cognitive load. From an anatomical point of view and more specifically at the cortical level, the integration mechanism appeared to be, until recently, a characteristic possessed only by associative areas at the top of the hierarchy of information processing. We now know that low level cortical areas, thought up to then to be only unisensory, are implicated in multisensory processes, thus raising the question about subcortical areas. Anatomical studies have shown the existence of thalamic nuclei which, through their connectivity, could allow for a rapid transfer and even an integration of sensory information. This new literature demonstrates the high complexity of the multisensory cerebral networks. In two electrophysiological studies in the monkey, we examined the multisensory properties of two structures, the posterior cingulate gyrus and the median pulvinar, which had never been explored before in a multisensory context. We not only showed that these structures are multisensory, but also integrative and that they could be part of the same functional network. This thesis has brought additional elements towards a better understanding of multisensory integration processes at the behavioral level and about the underlying brain networks, in particular those linked with the integration of natural stimuli.

We can affirm to apprehend a system in its totality only when we know how it behaves under its natural operating conditions. However, in the face of the complexity of the world, science can only evolve by simplifications, which paradoxically hide a good deal of the very mechanisms we are interested in. On the other hand, scientific enterprise is very tightly related to the advances in technology and the latter inevitably influences the manner in which the scientific experiments are conducted. Due to this factor, experimental conditions which would have been impossible to bring into laboratory not more than 20 years ago, are today within our reach. This thesis investigates neuronal integrative processes by using a variety of theoretical and experimental techniques wherein the approximation of ecologically relevant conditions within the laboratory is the common denominator. The working hypothesis of this thesis is that neurons and neuronal systems, in the sensory and higher cortices, are specifically adapted, as a result of evolutionary processes, to the sensory signals most likely to be received under ecologically relevant conditions. In order to conduct the present study along this line, we first recorded movies with the help of two microcameras carried by cats exploring a natural environment. This resulted in a database of binocular natural movies that was used in our theoretical and experimental studies. In a theoretical study, we aimed to understand the principles of binocular disparity encoding in terms of spatio-temporal statistical properties of natural movies in conjunction with simple mathematical expressions governing the activity levels of simulated neurons. In an unsupervised learning scheme, we used the binocular movies as input to a neuronal network and obtained receptive fields that represent these movies optimally with respect to the temporal stability criterion. Many distinctive aspects of the binocular coding in complex cells, such as the phase and position encoding of disparity and the existence of unbalanced ocular contributions, were seen to emerge as the result of this optimization process. Therefore we conclude that the encoding of binocular disparity by complex cells can be understood in terms of an optimization process that regulates activities of neurons receiving ecologically relevant information. Next we aimed to physiologically characterize the responses of the visual cortex to ecologically relevant stimuli in its full complexity and compare these to the responses evoked by artificial, conventional laboratory stimuli. To achieve this, a state-of-the-art recording method, voltage-sensitive dye imaging was used. This method captures the spatio-temporal activity patterns within the millisecond range across large cortical portions spanning over many pinwheels and orientation columns. It is therefore very well suited to provide a faithful picture of the cortical state in its full complexity. Drifting bar stimuli evoked two major sets of components, one coding for the position and the other for the orientation of the grating. Responses to natural stimuli involved more complex dynamics, which were locked to the motion present in the natural movies. In response to drifting gratings, the cortical state was initially dominated by a strong excitatory wave. This initial spatially widespread hyper-excitatory state had a detrimental effect on feature selectivity. In contrast, natural movies only rarely induced such high activity levels and the onset of inhibition cut short a further increase in activation level. An increase of 30% of the movie contrast was estimated to be necessary in order to produce activity levels comparable to gratings. These results show that the operating regime within which the natural movies are processed differs remarkably. Moreover, it remains to be established to what extent the cortical state under artificial conditions represents a valid state to make inferences concerning operationally more relevant input. The primary visual cortex contains a dense web of neuronal connections linking distant neurons. However the flow of information within this local network is to a large extent unknown under natural stimulation conditions. To functionally characterize these long-range intra-areal interactions, we presented natural movies also locally through either one or two apertures and analyzed the effects of the distant visual stimulation on the local activity levels. The distant patch had a net facilitatory effect on the local activity levels. Furthermore, the degree of the facilitation was dependent on the congruency between the two simultaneously presented movie patches. Taken together, our results indicate that the ecologically relevant stimuli are processed within a distinct operating regime characterized by moderate levels of excitation and/or high levels of inhibition, where facilitatory cooperative interactions form the basis of integrative processes. To gather better insights into the motion locking phenomenon and test the generalizability of the local cooperative processes toward larger scale interactions, we resorted to the unequalized temporal resolution of EEG and conducted a multimodal study. Inspired from the temporal properties of our natural movies, we designed a dynamic multimodal stimulus that was either congruent or incongruent across visual and auditory modalities. In the visual areas, the dynamic stimulation unfolded neuronal oscillations with frequencies well above the frequency spectrum content of the stimuli and the strength of these oscillations was coupled to the stimuli's motion profile. Furthermore, the coupling was found to be stronger in the case where the auditory and visual streams were congruent. These results show that the motion locking, which was so far observed in cats, is a phenomenon that also exists in humans. Moreover, the presence of long-range multimodal interactions indicates that, in addition to local intra-areal mechanisms ensuring the integration of local information, the central nervous system embodies an architecture that enables also the integration of information on much larger scales spread across different modalities. Any characterization of integrative phenomena at the neuronal level needs to be supplemented by its effects at the behavioral level. We therefore tested whether we could find any evidence of integration of different sources of information at the behavioral level using natural stimuli. To this end, we presented to human subjects images of natural scenes and evaluated the effect of simultaneously played localized natural sounds on their eye movements. The behavior during multimodal conditions was well approximated by a linear combination of the behavior under unimodal conditions. This is a strong indication that both streams of information are integrated in a joint multimodal saliency map before the final motor command is produced. The results presented here validate the possibility and the utility of using natural stimuli in experimental settings. It is clear that the ecological relevance of the experimental conditions are crucial in order to elucidate complex neuronal mechanisms resulting from evolutionary processes. In the future, having better insights on the nervous system can only be possible when the complexity of our experiments will match to the complexity of the mechanisms we are interested in.