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The leit motif of this thesis is that binding of visual features is a process that begins with input of stimulation and ends with the emergence of an object in working memory so that it can be further manipulated for higher cognitive processes. The primary focus was on the binding process from 0 to 2500 ms, with stimuli defined by location, colour, and shape. The initial experiments explored the relative role of topdown and bottom-up factors. Task relevance was compared by asking participants to detect swaps in bindings of two features whilst the third was either unchanged, or made irrelevant by randomization from study to test, in a change detection task. The experiments also studied the differences among the three defining features across experiments where each feature was randomized, whilst the binding between the other two was tested. Results showed that though features were processed to different time scales, they were treated in the same way by Visual Working Memory processes. Relevant features were consolidated and irrelevant features were inhibited. Later experiments confirmed that consolidation was aided by iconic memory and the inhibitory process was primarily a post-perceptual active inhibition.

Accurate memory for an object or event requires that multiple diverse features are bound together and retained as an integrated representation. There is overwhelming evidence that healthy ageing is accompanied by an associative deficit in that older adults struggle to remember relations between items above any deficit exhibited in remembering the items themselves. However, the effect of age on the ability to bind features within novel objects (for example, their colour and shape) and retain correct conjunctions over brief intervals is less clear. The relatively small body of work that exists on this topic to-date has suggested no additional working memory impairment for conjunctions of features beyond a general age-related impairment in the ability to temporarily retain features. This is in stark contrast to the feature binding deficit observed in the early stages of Alzheimer’s disease. Nevertheless, there have been reports of age-related feature binding deficits in working memory under specific circumstances. Thus a major focus of the present work was to assess these potential boundary conditions. The change detection paradigm was used throughout this work to examine age-differences in visual working memory. Despite the popularity of this task important issues regarding the way in which working memory is probed have been left unaddressed. Chapter 2 reports three experiments with younger adults comparing two methods of testing recognition memory for features or conjunctions. Contrary to an influential study in the field, it appears that processing multiple items at test does not differentially impact on participants’ ability to detect binding changes. Chapters 3, 4, and 5 report a series of experiments motivated by previous findings of specific age-related feature binding deficits. These experiments, improving on previous methodology where possible, demonstrate that increasing the amount of time for which items can be studied (Chapter 3) or mixing feature-conjunction changes in trial-blocks with more salient changes to individual features (Chapters 4 and 5) does not differentially impact on healthy older adults’ ability to detect binding changes. Rather, the argument is made that specific procedural aspects of previous work led to the appearance of deficits that do not generalise. Chapter 5 also addresses the suggestion that healthy ageing specifically affects the retention of item-location conjunctions. The existing evidence for this claim is reviewed, and found wanting, and new data are presented providing evidence against it. To follow-up on the absence of a deficit for simple feature conjunctions, Chapter 6 contrasts two theoretically distinct binding mechanisms: one for features intrinsic to an object and another for extrinsic, contextual features. Preliminary evidence is reported that the cost associated with retaining pairings of features is specifically pronounced for older adults when the features are extrinsic to each other. In an attempt to separate out the contribution of working memory capacity and lapses of attention to age-differences in overall task performance, Chapter 7 reports the results of an exploratory analysis using processing models developed in Chapter 2. Analysis of two data sets from Chapters 4 and 5 demonstrates that lapses of attention make an important contribution to differences in change detection performance. Chapter 8 returns to the issue of measurement in assessing the evidence for specific age-related deficits. Simulations demonstrate that the choice of outcome measure can greatly affect conclusions regarding age-group by condition interactions, suggesting that some previous findings of such interactions in the literature may have been more apparent than real. In closing the General Discussion relates the present work to current theory regarding feature binding in visual working memory and to the wider literature on binding deficits in healthy and pathological ageing.

Yes / Human memory is content addressable—i.e., contents of the memory can be accessed using partial information about the bound features of a stored item. In this study, we used a cross-feature cuing technique to examine how the human visual system encodes, binds, and retains information about multiple stimulus features within a set of moving objects. We sought to characterize the roles of three different features (position, color, and direction of motion, the latter two of which are processed preferentially within the ventral and dorsal visual streams, respectively) in the construction and maintenance of object representations. We investigated the extent to which these features are bound together across the following processing stages: during stimulus encoding, sensory (iconic) memory, and visual shortterm memory. Whereas all features examined here can serve as cues for addressing content, their effectiveness shows asymmetries and varies according to cue–report pairings and the stage of information processing and storage. Position-based indexing theories predict that position should be more effective as a cue compared to other features. While we found a privileged role for position as a cue at the stimulus-encoding stage, position was not the privileged cue at the sensory and visual short-term memory stages. Instead, the pattern that emerged from our findings is one that mirrors the parallel processing streams in the visual system. This stream-specific binding and cuing effectiveness manifests itself in all three stages of information processing examined here. Finally, we find that the Leaky Flask model proposed in our previous study is applicable to all three features.

Feature binding refers to the fundamental challenge of the brain to integrate sensory information registered by distinct brain regions to form a unified neural representation of a stimulus. While the human cognitive literature has established that attentional processes in a frontoparietal cortical network support feature binding, the neurochemical contributions to this attentional process remain unknown. Using systemic administration of the cholinergic muscarinic receptor antagonist scopolamine and a digging-based rat feature binding task that used both odor and texture stimuli, it was demonstrated that blockade of acetylcholine (ACh) at the muscarinic receptors impaired rats’ ability to feature bind at encoding, and it was proposed that ACh may support the attentional processes necessary for feature binding (Botly & De Rosa, 2007). This series of experiments further investigated a role for ACh and the cholinergic basal forebrain (BF) in feature binding. In Experiment 1, a cross-species experimental design was employed in which rats under the systemic influence of scopolamine and human participants under divided-attention performed comparable feature binding tasks using odor stimuli for rats and coloured-shape visual stimuli for humans. Given the comparable performance impairments demonstrated by both species, Experiment 1 suggested that ACh acting at muscarinic receptors supports the attentional processes necessary for feature binding at encoding. Experiments 2-4 investigated the functional neuroanatomy of feature binding using bilateral quisqualic acid excitotoxic (Experiment 2) and 192 IgG-saporin cholinergic immunotoxic (Experiments 3 and 4) brain lesions that were assessed for completeness using histological and immunohistological analyses. Using the crossmodal digging-based rat feature binding task, Experiment 2 revealed that the nucleus basalis magnocellularis (NBM) of the BF is critically involved in feature binding, and Experiment 3 revealed that cholinergic neurons in the NBM are necessary for feature binding at encoding. Lastly, in Experiment 4, rats performed visual search, the standard test of feature binding in humans, with touchscreen-equipped operant chambers. Here it was also revealed that cholinergic neurons in the NBM of the BF are critical for efficient visual search. Taken together, these behavioural, pharmacological, and brain-lesion findings have provided insights into the neurochemical contributions to the fundamental attentional process of feature binding.

Feature binding refers to the fundamental challenge of the brain to integrate sensory information registered by distinct brain regions to form a unified neural representation of a stimulus. While the human cognitive literature has established that attentional processes in a frontoparietal cortical network support feature binding, the neurochemical contributions to this attentional process remain unknown. Using systemic administration of the cholinergic muscarinic receptor antagonist scopolamine and a digging-based rat feature binding task that used both odor and texture stimuli, it was demonstrated that blockade of acetylcholine (ACh) at the muscarinic receptors impaired rats’ ability to feature bind at encoding, and it was proposed that ACh may support the attentional processes necessary for feature binding (Botly & De Rosa, 2007). This series of experiments further investigated a role for ACh and the cholinergic basal forebrain (BF) in feature binding. In Experiment 1, a cross-species experimental design was employed in which rats under the systemic influence of scopolamine and human participants under divided-attention performed comparable feature binding tasks using odor stimuli for rats and coloured-shape visual stimuli for humans. Given the comparable performance impairments demonstrated by both species, Experiment 1 suggested that ACh acting at muscarinic receptors supports the attentional processes necessary for feature binding at encoding. Experiments 2-4 investigated the functional neuroanatomy of feature binding using bilateral quisqualic acid excitotoxic (Experiment 2) and 192 IgG-saporin cholinergic immunotoxic (Experiments 3 and 4) brain lesions that were assessed for completeness using histological and immunohistological analyses. Using the crossmodal digging-based rat feature binding task, Experiment 2 revealed that the nucleus basalis magnocellularis (NBM) of the BF is critically involved in feature binding, and Experiment 3 revealed that cholinergic neurons in the NBM are necessary for feature binding at encoding. Lastly, in Experiment 4, rats performed visual search, the standard test of feature binding in humans, with touchscreen-equipped operant chambers. Here it was also revealed that cholinergic neurons in the NBM of the BF are critical for efficient visual search. Taken together, these behavioural, pharmacological, and brain-lesion findings have provided insights into the neurochemical contributions to the fundamental attentional process of feature binding.

Les systèmes sensoriels encodent l’information sur notre environnement sous la forme d’impulsions électriques qui se propagent dans des réseaux de neurones. Élucider le code neuronal – les principes par lesquels l’information est représentée dans l’activité des neurones – est une question fondamentale des neurosciences. Cette thèse constituée de 3 études (E) s’intéresse à deux types de codes, la synchronisation et l’adaptation, dans les neurones du cortex visuel primaire (V1) du chat. Au niveau de V1, les neurones sont sélectifs pour des propriétés comme l’orientation des contours, la direction et la vitesse du mouvement. Chaque neurone ayant une combinaison de propriétés pour laquelle sa réponse est maximale, l’information se retrouve distribuée dans différents neurones situés dans diverses colonnes et aires corticales. Un mécanisme potentiel pour relier l’activité de neurones répondant à des items eux-mêmes reliés (e.g. deux contours appartenant au même objet) est la synchronisation de leur activité. Cependant, le type de relations potentiellement encodées par la synchronisation n’est pas entièrement clair (E1). Une autre stratégie de codage consiste en des changements transitoires des propriétés de réponse des neurones en fonction de l’environnement (adaptation). Cette plasticité est présente chez le chat adulte, les neurones de V1 changeant d’orientation préférée après exposition à une orientation non préférée. Cependant, on ignore si des neurones spatialement proches exhibent une plasticité comparable (E2). Finalement, nous avons étudié la dynamique de la relation entre synchronisation et plasticité des propriétés de réponse (E3). Résultats principaux — (E1) Nous avons montré que deux stimuli en mouvement soit convergent soit divergent élicitent plus de synchronisation entre les neurones de V1 que deux stimuli avec la même direction. La fréquence de décharge n’était en revanche pas différente en fonction du type de stimulus. Dans ce cas, la synchronisation semble coder pour la relation de cocircularité dont le mouvement convergent (centripète) et divergent (centrifuge) sont deux cas particuliers, et ainsi pourrait jouer un rôle dans l’intégration des contours. Cela indique que la synchronisation code pour une information qui n’est pas présente dans la fréquence de décharge des neurones. (E2) Après exposition à une orientation non préférée, les neurones changent d’orientation préférée dans la même direction que leurs voisins dans 75% des cas. Plusieurs propriétés de réponse des neurones de V1 dépendent de leur localisation dans la carte fonctionnelle corticale pour l’orientation. Les comportements plus diversifiés des 25% de neurones restants sont le fait de différences fonctionnelles que nous avons observé et qui suggèrent une localisation corticale particulière, les singularités, tandis que la majorité des neurones semblent situés dans les domaines d’iso-orientation. (E3) Après adaptation, les paires de neurones dont les propriétés de réponse deviennent plus similaires montrent une synchronisation accrue. Après récupération, la synchronisation retourne à son niveau initial. Par conséquent, la synchronisation semble refléter de façon dynamique la similarité des propriétés de réponse des neurones. Conclusions — Cette thèse contribue à notre connaissance des capacités d’adaptation de notre système visuel à un environnement changeant. Nous proposons également des données originales liées au rôle potentiel de la synchronisation. En particulier, la synchronisation semble capable de coder des relations entre objets similaires ou dissimilaires, suggérant l’existence d’assemblées neuronales superposées. / Sensory systems encode information about our environment into electrical impulses that propagate in networks of neurons. Understanding the neural code – the principles by which information is represented in neuronal activity – is one of the most fundamental issues in neuroscience. This thesis investigates in a series of 3 studies (S) two coding mechanisms, synchrony and adaptation, in neurons of the cat primary visual cortex (V1). In V1, neurons display selectivity for image features such as contour orientation, motion direction and velocity. Each neuron has at least one combination of features that elicits its maximum firing rate. Visual information is thus distributed among numerous neurons within and across cortical columns, modules and areas. Synchronized electrical activity between cells was proposed as a potential mechanism underlying the binding of related features to form coherent perception. However, the precise nature of the relations between image features that may elicit neuronal synchrony remains unclear (S1). In another coding strategy, sensory neurons display transient changes of their response properties following prolonged exposure to an appropriate stimulus (adaptation). In adult cat V1, orientation-selective neurons shift their preferred orientation after being exposed to a non-preferred orientation. How the adaptive behavior of a neuron is related to that of its neighbors remains unclear (S2). Finally, we investigated the relationship between synchrony and orientation tuning in neuron pairs, especially how synchrony is modulated during adaptation-induced plasticity (S3). Main results — (S1) We show that two stimuli in either convergent or divergent motion elicit significantly more synchrony in V1 neuron pairs than two stimuli with the same motion direction. Synchronization seems to encode the relation of cocircularity, of which convergent (centripetal) and divergent (centrifugal) motion are two special instances, and could thus play a role in contour integration. Our results suggest that V1 neuron pairs transmit specific information on distinct image configurations through stimulus-dependent synchrony of their action potentials. (S2) We show that after being adapted to a non-preferred orientation, cells shift their preferred orientation in the same direction as their neighbors in most cases (75%). Several response properties of V1 neurons depend on their location within the cortical orientation map. The differences we found between cell clusters that shift in the same direction and cell clusters with both attractive and repulsive shifts suggest a different cortical location, iso-orientation domains for the former and pinwheel centers for the latter. (S3) We found that after adaptation, neuron pairs that share closer tuning properties display a significant increase of synchronization. Recovery from adaptation is accompanied by a return to the initial synchrony level. Synchrony therefore seems to reflect the similarity in neurons’ response properties, and varies accordingly when these properties change. Conclusions — This thesis further advances our understanding of how visual neurons adapt to a changing environment, especially regarding cortical network dynamics. We also propose novel data about the potential role of synchrony. Especially, synchrony appears capable of binding various features, whether similar or dissimilar, suggesting superimposed neural assemblies.

Les systèmes sensoriels encodent l’information sur notre environnement sous la forme d’impulsions électriques qui se propagent dans des réseaux de neurones. Élucider le code neuronal – les principes par lesquels l’information est représentée dans l’activité des neurones – est une question fondamentale des neurosciences. Cette thèse constituée de 3 études (E) s’intéresse à deux types de codes, la synchronisation et l’adaptation, dans les neurones du cortex visuel primaire (V1) du chat. Au niveau de V1, les neurones sont sélectifs pour des propriétés comme l’orientation des contours, la direction et la vitesse du mouvement. Chaque neurone ayant une combinaison de propriétés pour laquelle sa réponse est maximale, l’information se retrouve distribuée dans différents neurones situés dans diverses colonnes et aires corticales. Un mécanisme potentiel pour relier l’activité de neurones répondant à des items eux-mêmes reliés (e.g. deux contours appartenant au même objet) est la synchronisation de leur activité. Cependant, le type de relations potentiellement encodées par la synchronisation n’est pas entièrement clair (E1). Une autre stratégie de codage consiste en des changements transitoires des propriétés de réponse des neurones en fonction de l’environnement (adaptation). Cette plasticité est présente chez le chat adulte, les neurones de V1 changeant d’orientation préférée après exposition à une orientation non préférée. Cependant, on ignore si des neurones spatialement proches exhibent une plasticité comparable (E2). Finalement, nous avons étudié la dynamique de la relation entre synchronisation et plasticité des propriétés de réponse (E3). Résultats principaux — (E1) Nous avons montré que deux stimuli en mouvement soit convergent soit divergent élicitent plus de synchronisation entre les neurones de V1 que deux stimuli avec la même direction. La fréquence de décharge n’était en revanche pas différente en fonction du type de stimulus. Dans ce cas, la synchronisation semble coder pour la relation de cocircularité dont le mouvement convergent (centripète) et divergent (centrifuge) sont deux cas particuliers, et ainsi pourrait jouer un rôle dans l’intégration des contours. Cela indique que la synchronisation code pour une information qui n’est pas présente dans la fréquence de décharge des neurones. (E2) Après exposition à une orientation non préférée, les neurones changent d’orientation préférée dans la même direction que leurs voisins dans 75% des cas. Plusieurs propriétés de réponse des neurones de V1 dépendent de leur localisation dans la carte fonctionnelle corticale pour l’orientation. Les comportements plus diversifiés des 25% de neurones restants sont le fait de différences fonctionnelles que nous avons observé et qui suggèrent une localisation corticale particulière, les singularités, tandis que la majorité des neurones semblent situés dans les domaines d’iso-orientation. (E3) Après adaptation, les paires de neurones dont les propriétés de réponse deviennent plus similaires montrent une synchronisation accrue. Après récupération, la synchronisation retourne à son niveau initial. Par conséquent, la synchronisation semble refléter de façon dynamique la similarité des propriétés de réponse des neurones. Conclusions — Cette thèse contribue à notre connaissance des capacités d’adaptation de notre système visuel à un environnement changeant. Nous proposons également des données originales liées au rôle potentiel de la synchronisation. En particulier, la synchronisation semble capable de coder des relations entre objets similaires ou dissimilaires, suggérant l’existence d’assemblées neuronales superposées. / Sensory systems encode information about our environment into electrical impulses that propagate in networks of neurons. Understanding the neural code – the principles by which information is represented in neuronal activity – is one of the most fundamental issues in neuroscience. This thesis investigates in a series of 3 studies (S) two coding mechanisms, synchrony and adaptation, in neurons of the cat primary visual cortex (V1). In V1, neurons display selectivity for image features such as contour orientation, motion direction and velocity. Each neuron has at least one combination of features that elicits its maximum firing rate. Visual information is thus distributed among numerous neurons within and across cortical columns, modules and areas. Synchronized electrical activity between cells was proposed as a potential mechanism underlying the binding of related features to form coherent perception. However, the precise nature of the relations between image features that may elicit neuronal synchrony remains unclear (S1). In another coding strategy, sensory neurons display transient changes of their response properties following prolonged exposure to an appropriate stimulus (adaptation). In adult cat V1, orientation-selective neurons shift their preferred orientation after being exposed to a non-preferred orientation. How the adaptive behavior of a neuron is related to that of its neighbors remains unclear (S2). Finally, we investigated the relationship between synchrony and orientation tuning in neuron pairs, especially how synchrony is modulated during adaptation-induced plasticity (S3). Main results — (S1) We show that two stimuli in either convergent or divergent motion elicit significantly more synchrony in V1 neuron pairs than two stimuli with the same motion direction. Synchronization seems to encode the relation of cocircularity, of which convergent (centripetal) and divergent (centrifugal) motion are two special instances, and could thus play a role in contour integration. Our results suggest that V1 neuron pairs transmit specific information on distinct image configurations through stimulus-dependent synchrony of their action potentials. (S2) We show that after being adapted to a non-preferred orientation, cells shift their preferred orientation in the same direction as their neighbors in most cases (75%). Several response properties of V1 neurons depend on their location within the cortical orientation map. The differences we found between cell clusters that shift in the same direction and cell clusters with both attractive and repulsive shifts suggest a different cortical location, iso-orientation domains for the former and pinwheel centers for the latter. (S3) We found that after adaptation, neuron pairs that share closer tuning properties display a significant increase of synchronization. Recovery from adaptation is accompanied by a return to the initial synchrony level. Synchrony therefore seems to reflect the similarity in neurons’ response properties, and varies accordingly when these properties change. Conclusions — This thesis further advances our understanding of how visual neurons adapt to a changing environment, especially regarding cortical network dynamics. We also propose novel data about the potential role of synchrony. Especially, synchrony appears capable of binding various features, whether similar or dissimilar, suggesting superimposed neural assemblies.