<p>We investigated 1) how objects come to serve as landmarks in spatial memory and more specifically how they form part of an allocentric cognitive map and 2) how humans encode multiple connected spatial environments. In both sets of experiments, participants performing a virtual driving task incidentally learned the layout of a town and locations of objects or stores in that town. Their spatial memory and recognition memory for the objects or stores were subsequently tested. To assess whether the objects were encoded allocentrically, we developed a new measurement, pointing consistency. We found that when participants had more limited experience of the environment spatial memory for objects at navigationally relevant locations was more consistent across tested viewpoints than for objects at navigationally less relevant locations. When participants’ attention was focused on the appearance of objects, the navigational relevance effect was eliminated, whereas when their attention was focused on the objects’ locations, this effect was enhanced, supporting the hypothesis that when objects are processed in the service of navigation, rather than merely being viewed as objects, they engage qualitatively distinct attentional systems and are incorporated into an allocentric spatial representation. The results were consistent with evidence from the neuroimaging literature that when objects are relevant to navigation, they not only engage the ventral “object processing stream”, but also the dorsal stream and medial temporal lobe memory system classically associated with allocentric spatial memory. Moreover, in the connected environments, our data were more consistent with the formation of local maps, regardless of whether the neighborhoods were learned together or separately. Only when all visible distinctions between neighborhoods were removed did people behave as if they formed one integrated map. These data are broadly consistent with evidence from rodent hippocampal place cell recordings in connected boxes, and with hierarchical models of spatial coding.</p> / Doctor of Philosophy (PhD)

La navigation et les modes de déplacement intéressent la communauté scientifique depuis maintenant près d'un demi siècle. Cependant, l’augmentation de l’incidence des troubles dégénératifs du système nerveux central chez l’homme rend plus prégnante la nécessité de compréhension de la navigation et de l’influence du temps sur celle-ci. S'il est connu chez l'homme comme chez le rongeur que l'avancée en âge affecte les capacités à se déplacer dans de vastes environnements, peu de données sont disponibles quant aux processus cognitifs impliqués dans ce type de comportement et leurs éventuelles modulations avec l'âge. La définition des stratégies utilisées, l’incidence respective des mécanismes allocentriques et égocentriques, la capacité de mise en œuvre d’une stratégie au moment demandé, lors d’un rappel à court ou à long délai, l’influence du temps qui passe sont autant de questions que nous avons abordées dans ce travail de thèse. Afin d’effectuer ces études, nous avons développé des tâches en environnements virtuels modélisés sur ordinateur et utilisé des tests neuropsychologiques nécessitant la mobilisation des compétences visuo-spatiales. Dans une première étude utilisant une épreuve de localisation spatiale, les résultats obtenus mettent en évidence chez les personnes âgées, une altération des aptitudes lors de la mise en œuvre d’une stratégie allocentrique, sans atteinte des performances égocentriques. La deuxième étude utilisant une version virtuelle du test de la piscine de Morris reconnu comme une tâche allocentrique chez le rongeur, conforte ces données. De façon similaire dans les deux études, les personnes âgées présentent une altération de la sélection et de l’exécution de la stratégie de déplacement qui s’avère optimale pour résoudre la tâche spatiale. Nous avons également mis en évidence une difficulté, chez ces mêmes participants, à utiliser une représentation mentale globale de l’espace, sans toutefois qu’il soit possible de distinguer si l’origine de cette difficulté vient d’une altération de la formation ou de la récupération de cette « carte cognitive ». Le temps pourrait également jouer son rôle de par le délai entre l'acquisition d'une information spatiale et le moment où il est nécessaire de l’utiliser à nouveau. En étudiant l’effet du délai sur la trace mnésique spatiale, nous avons observé que les sujets jeunes utilisant de façon prédominante une stratégie allocentrique voyaient leurs performances diminuer lors d’un rappel après quatre semaines alors que celles des sujets âgés restaient inchangées. Ceci soulève bien entendu la question de la différence d’encodage des informations entre les sujets jeunes et âgés, avec un versant plus détaillé chez les sujets jeunes, mais surtout s’intègre au sein du débat actuel sur l’existence d’une modification de la trace mnésique qui pourrait selon la théorie des traces multiples de la consolidation, évoluer vers un souvenir plus schématisé avec le délai. Les résultats d’une dernière étude dans laquelle nous manipulons le contexte environnemental de la piscine virtuelle de Morris, amène des arguments en faveur d’une « schématisation » du souvenir au cours de la consolidation, en mettant en évidence une absence de discrimination par les participants, d’un changement des repères spatiaux lors d’un rappel de l’information après six semaines de délai. Toutes ces données sont discutées dans le cadre du débat actuel de la consolidation, notamment sur la contribution de l’hippocampe dans le stockage et le rappel des informations anciennes. A la lumière de nos données, nous proposons une vue intégrative du fonctionnement de l’interface hippocampo-corticale lors des rappels après un court et long délai, en fonction de l’âge. / While the detrimental effects of human aging on cognitive functions are well documented, how normal aging affects spatial memory processing and the organization of recent and long-term memories remains unclear. What are the cognitive strategies used when confronted to spatial navigation in large environments? How are the selection and use of these strategies affected by aging? How are recent and long-term remote memories organized as a function of aging during systems-level consolidation? These are the questions we sought to address during the course of this Ph.D. thesis by developing a series of virtual environments aimed at assessing spatial navigation and memory performance in young adults and aged participants. In a first series of experiments, participants were tested for object location memory in a virtual environment (a medieval castle) that enabled shifts in spatial viewpoints between study and test. Aged participants exhibited poor performance relative to young adults only in the shifted view conditions, thus providing strong evidence for a decline in allocentric, but not egocentric, spatial memory. In contrast to young adults, aged participants exhibited difficulties in processing efficiently distal cues of the environment and were less prone to adopt allocentric strategies. Manipulations of the spatial layout of the environment led us to the conclusion that aging seems to preferentially interfere with the capacity to form or use mental representations built upon all pieces of the environmental features which typically, are never in full view in real world large-scale environments. In a second set of experiments, participants were tested in an ecologically-relevant virtual version of the Morris water maze which mimics that classically used in rodents. Aged participants performed more poorly compared to middle-aged and young adults and formed a more schematic spatial memory. They favoured a directional single cue-based strategy to locate the hidden platform contrasting with young adults who formed complex geometrical relationships between distal cues of the environment. A neuropsychological test battery confirmed that binding of unrelated items and abilities to mentally manipulate information were two processes involved in solving the water maze task. Thus, upon acquisition, aged participants had difficulties in forming experientially detailed cognitive maps and in binding unrelated features of the environment into a cohesive spatial memory, possibly indicative of altered hippocampal-frontal circuitry. We next proceeded to examine the organization of spatial memory as a function of time. Long-term memory assessed 4 weeks after acquisition revealed that performance decreased more rapidly in young adults compared to elderly participants, suggesting that the passage of time differentially affects the content of spatial memory, richly detailed spatial memories being more vulnerable to decay than schematic ones. This concept of memory transformation (i.e. memories are not stored in the cortex in their original form) was supported by findings of a last experiment in which we provide evidence that participants failed in detecting changes in the spatial layout of the pool as memories matured over time. All these findings are discussed in the context of the current debate about the concept of memory consolidation which opposes the standard model of memory consolidation to the multiple trace theory, two views which make different predictions about the contribution of the hippocampus to remote memory storage and retrieval. In light of our own findings, we attempt to propose an integrative view of the functioning of the hippocampal-cortical interface during recent and remote memory retrieval as a function of normal aging.

La mémoire épisodique lie différents éléments dans un contexte spatial et temporel particulier. Il a été proposé que lors de la récupération d‟un épisode, la ré-instanciation d‟une cohérence entre les éléments néocorticaux soit opérée grâce à une représentation spatiale allocentrée stockée au niveau de la structure hippocampique (i.e., codage de la position des objets entre eux, indépendamment de la position de l‟individu, Burgess, Becker, King, & O'Keefe, 2001; Nadel & Moscovitch, 1998). Ce travail de thèse propose de traiter la mémoire épisodique et le sentiment de projection dans son passé (i.e., conscience autonoétique) comme une qualité attribuée à une dextérité relative dans le traitement spatial égocentré mis à jour (i.e., la position, orientation et le déplacement de son corps dans l‟environnement). Le rôle des traitements spatiaux allocentrés et égocentrés mis à jour dans la mémoire épisodique a été évalué expérimentalement. Les résultats suggèrent l‟existence d‟un lien causal entre le traitement de la mise à jour égocentré et les performances de mémoire épisodique. De plus, les études ont mis en évidence l‟existence de spécificités cérébrales et comportementales de la mise à jour égocentrée confirmant l‟adéquation de ce traitement au modèle théorique proposé. Par ailleurs, en référence à cette dissociation entre l‟information égocentrée mis à jour et allocentrée, des études neuropsychologiques ont révélé la présence de déficits de la mise à jour égocentrée, et d‟une préservation allocentrée dans l‟amnésie bihippocampique qu‟elle soit acquise ou développementale. Enfin, l‟évaluation des conséquences cérébrales lors de la récupération épisodique d‟un encodage maximisant le traitement égocentré mis à jour a permis de révéler une implication spécifique des structures temporo-pariétales. Ce travail de thèse a été organisé autour d‟un modèle théorique original du fonctionnement de la mémoire épisodique proposant de nouvelles prédictions expérimentales. Les approches comportementale, neuropsychologique et en imagerie fonctionnelle soulèvent à leur tour de nouvelles pistes de recherche sur le lien entre conscience de son corps et mémoire épisodique. / Episodic memory binds various elements in a specific spatial and temporal context. During retrieval, disparate neocortical elements can be re-associated into a coherent episode due to an allocentric spatial context maintained within the hippocampal formation (ie, coding for object-to-object relations, independently of the individual‟s position, Burgess, Becker, King, & O‟Keefe, 2001, Nadel & Moscovitch, 1998). Phenomenological experience is characteristic of episodic memory. In this thesis, it is described as an individual‟s attribution to a fluency in processing egocentric-updating spatial information (i.e., the position, orientation and movement of one‟s body) during retrieval. The function of egocentric-updating and of allocentric spatial processing in episodic memory was assessed experimentally. Results demonstrate the presence of a causal link between egocentric-updating and episodic memory performance. Moreover, experiments showed cerebral and behavioural specificities of egocentric-updating spatial processing supporting its involvement in episodic memory. Additionally, in line with this distinction between allocentric and egocentric-updating spatial processing, neuropsychological experiments revealed deficits in egocentric-updating with a preservation of allocentric spatial processing in both acquired and developemental bi-hippocampal amnesia. Finally, the assessment of cerebral consequences of encoding an episode while maximizing egocentric-updating processes revealed a higher involvement of temporo-parietal regions during the subsequent episodic retrieval. This thesis work was structured over an original theoretical model on episodic memory functioning allowing new experimental predictions. Combining behavioural, neuropsychological and neuroimaging approaches raised in turn new questions concerning links between episodic memory and self-consciousness.

La domanda "Che cos’è lo spazio?" è sempre stata un tema centrale per la filosofia, ed è diventata di interesse anche per la psicologia cognitiva e per le neuroscienze, con una domanda cruciale strettamente legata: "Dove sono io?". Lo sforzo per collegare le risposte a queste due domande mira proprio a comprendere la complessa relazione che esiste tra lo spazio interno ed esterno, che è l'obiettivo finale di questo lavoro. L'idea è che la nostra posizione nel mondo influenzi fortemente il modo in cui codifichiamo, archiviamo e recuperiamo dalla memoria un layout spaziale. Inoltre, questo layout spaziale serve da impalcatura che vincola tutte le informazioni relative al nostro passato, presente e futuro, e tutte le esperienze legate al nostro corpo. All’interno di un approccio enattivo, si suggerisce una sincronizzazione continua (cioè, il “mental frame syncing") tra una rappresentazione allocentrica indipendente dal punto di vista allocentrica (i.e. che include solo relazioni oggetto-oggetto astratte) e una rappresentazione allocentrica dipendente dal punto di vista (i.e. che include informazioni sulla nostra direzione egocentrica attuale) possa permettere di posizionare il corpo nello “spazio memorizzato” rendendo più semplice la traduzione di questo in un “lived space” di cui si necessita per navigare, per ricordare il passato e per sentire il corpo. Sulla base di queste premesse teoriche, quattro studi sperimentali saranno presentati per studiare il ruolo del mental frame syncing come un principio di allineamento centrato sull’osservatore nei processi di codifica e di recupero delle informazioni. / The question "What is space?" has always been a central topic for philosophy, and a closely linked crucial question becomes of interest for cognitive psychology and neuroscience, that is "Where am I?" The efforts to answer these two questions are means to better understanding of the complex relation between the outer and the inner space, which is the final goal of this work. The idea is that that our bodily position in the world strongly affects the way in which we encode, store and retrieve a spatial layout. Moreover, this spatial layout serves as a scaffold, binding all the information of our past, present, future and body-related experiences. Within an enactive approach, it is suggested that this continuous synchronization (namely, the “mental frame syncing”) of an allocentric viewpoint-independent representation (i.e. including only abstract object-to-object relations) and an allocentric viewpoint-dependent representation (i.e. comprising information about our current heading) may permit to place current bodily position in the “memorized space" making easy the translation of it into a “lived space” that it is needed to navigate, remember the past and feel the body. On these theoretical premises, four experimental studies will be presented to investigate the role of mental frame syncing as an alignment principle centred on observer the processes of encoding and retrieval of information

<p>Navigation is a very important area of spatial information research that presents researchers with a number of challenges. One of these challenges concerns the nature of spatial information encoding itself: is such encoding the result of a single mechanism system, a two-mechanism system or possibly a mixed system? One possible avenue of insight into this problem centers on the disorientation effect as described in Wang & Spelke (2000). A quick survey of basic findings, terminating with Waller & Hodgson (2006), indicates that there seem to be two systems at work. Moreover, the results obtained are based upon experiments carried out in actual reality. A virtual reality experiment was designed in an attempt to replicate the findings described in Waller & Hodgson (2006). The experiment is described in detail and its results are presented. These were found to be sufficiently reliable to justify pointing to a potentially rich field for future research, including such techniques as combining VR with fMRI to achieve more fine-grained results that cannot currently be obtained from the direct use of actual reality only. Underlying factors such as experimental control and data presentation are briefly described in the discussion section.</p> / Master of Science (MS)

<p>Multiple reference frames are used to interact with our surroundings. When these reference frames are in conflict, processing errors can occur. For tactile stimuli, this conflict is highlighted when the hands are crossed over the midline of the body. In this posture, vibrotactile temporal order judgments (TOJs) presented to the hands are impaired compared to an uncrossed posture. This decrease in temporal processing is known as the crossed-hands deficit. The deficit was explored in depth throughout this thesis. In Chapters 2, 3 and 4 different elements of the crossed-hands deficit were evaluated including its connections to the rod and frame test, individual and sex differences within the TOJ task, as well as the influence of vision and body position. These elements were framed with underlying goal of investigating the root cause of the deficit. The data presented here provided evidence for a conflict model of crossed hands processing. A conflict between the internal and external reference frames produced the deficit in temporal processing when the hands were crossed. The role of the body’s midline in understanding multisensory integration was further considered in Chapter 5 through the rubber hand illusion, which is a visuotactile phenomenon whereby an unseen real hand is mislocalized towards a seen rubber hand. When the real hand, rubber hand, or both were crossed over the midline the illusion did not occur. It was hypothesized that a failure to integrate the tactile information presented to the real hand with the visual rubber hand was responsible for the absence of the illusion. Taken together, the data presented in this thesis contribute to the greater understanding of how reference frame conflicts are resolved, particularly when the conflict occurs across the body’s midline.</p> / Doctor of Philosophy (PhD)

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Percevoir son espace d’évolution est une activité déterminante dans l’élaboration des relations spatiales que nous tissons avec notre environnement. En neurosciences comportementales, l’étude de ces relations a généralement été abordée selon deux perspectives théoriques. L’une d’elle s’attache à décrire les relations au monde au travers des processus de perception directe impliquant notamment la notion d’affordances (i.e. de possibilités d’actions naturellement offertes par l’environnement) ; tandis que d’autres s’intéressent d’avantage aux aspects cognitifs de la perception avec la mise en place de processus de représentation spatiale. Cette dernière reflète notamment l’existence d’état(s) représenté(s) qu’il est possible de décrire à travers de la combinaison d’espaces stables appelés référentiels spatiaux. L’objectif de ce travail de thèse vise à mieux comprendre la contribution du référentiel égocentré (i.e. corporel) dans la perception de l’espace géocentré (i.e. gravitaire). La question a notamment été abordée autour de deux axes de recherche interrogeant d’une part (i) l’origine de l’influence égocentrée préalablement observée dans le noir sur la perception géocentrée, et d’autre part (ii) la présence du phénomène égocentré dans un contexte visuel plus enrichi suite à l’ajout d’un flux optique. Pour ce faire quatre études centrées autour d’un paradigme d’estimation des possibilités de franchissement d’obstacles hauts ont été réalisées. Pris dans leur ensemble, les résultats expérimentaux soulignent le caractère particulièrement puissant et complexe du phénomène égocentré corporel observé sur la perception de l’espace gravitaire. Ces résultats, discutés en termes d’interpénétrabilité entre référentiels spatiaux offrent un support d’étude intéressant sur la manière dont les référentiels sont utilisés dans les processus de représentation spatiale. / Perceiving space is a relevant task in determining our relationships with the environment. In behavioral neuroscience, investigating this spatial relationship can classically be explored with two theoretical approaches. The first one uses direct perception to describe the spatial relationships, involving affordances (i.e. the action ability naturally offer by the environment). The other one investigates the cognitive aspect of perception implying the use of spatial representation process. The later one traduces the existence of represented states which can be described through the interaction of different stable states called spatial reference frames. The present work investigates the contribution of the egocentric reference frame (body-related) on the perception of the geocentric space (earth-based). This was questioned through two research lines, (i) the origin of egocentric influence previously observed in darkness upon geocentric perception, (ii) the existence of the egocentric phenomenon in an enriched visual scene. To answer these questions, four experiments were conducted where the paradigm of passing under high obstacles was used. Overall, these results stress the powerful and complex aspect of the egocentric phenomenon observed upon geocentric perception. This work, discussed in term of interpenetrability between reference frames, provide an interesting support on the way how spatial reference frames are used in perceiving space.