Visual functions in the healthy and schizophrenic brain : from stimulus control to illusory perceptions using electrical neuroimaging

ABSTRACT (FRENCH)Ce travail de thèse basé sur le système visuel chez les sujets sains et chez les patients schizophrènes, s'articule autour de trois articles scientifiques publiés ou en cours de publication. Ces articles traitent des sujets suivants : le premier article présente une nouvelle méthode de traitement des composantes physiques des stimuli (luminance et fréquence spatiale). Le second article montre, à l'aide d'analyses de données EEG, un déficit de la voie magnocellulaire dans le traitement visuel des illusions chez les patients schizophrènes. Ceci est démontré par l'absence de modulation de la composante PI chez les patients schizophrènes contrairement aux sujets sains. Cette absence est induite par des stimuli de type illusion Kanizsa de différentes excentricités. Finalement, le troisième article, également à l'aide de méthodes de neuroimagerie électrique (EEG), montre que le traitement des contours illusoires se trouve dans le complexe latéro-occipital (LOC), à l'aide d'illusion « misaligned gratings ». De plus il révèle que les activités démontrées précédemment dans les aires visuelles primaires sont dues à des inférences « top- down ».Afin de permettre la compréhension de ces trois articles, l'introduction de ce manuscrit présente les concepts essentiels. De plus des méthodes d'analyses de temps-fréquence sont présentées. L'introduction est divisée en quatre parties : la première présente le système visuel depuis les cellules retino-corticales aux deux voix du traitement de l'information en passant par les régions composant le système visuel. La deuxième partie présente la schizophrénie par son diagnostic, ces déficits de bas niveau de traitement des stimuli visuel et ces déficits cognitifs. La troisième partie présente le traitement des contours illusoires et les trois modèles utilisés dans le dernier article. Finalement, les méthodes de traitement des données EEG seront explicitées, y compris les méthodes de temps-fréquences.Les résultats des trois articles sont présentés dans le chapitre éponyme (du même nom). De plus ce chapitre comprendra les résultats obtenus à l'aide des méthodes de temps-fréquenceFinalement, la discussion sera orientée selon trois axes : les méthodes de temps-fréquence ainsi qu'une proposition de traitement de ces données par une méthode statistique indépendante de la référence. La discussion du premier article en montrera la qualité du traitement de ces stimuli. La discussion des deux articles neurophysiologiques, proposera de nouvelles d'expériences afin d'affiner les résultats actuels sur les déficits des schizophrènes. Ceci pourrait permettre d'établir un marqueur biologique fiable de la schizophrénie.ABSTRACT (ENGLISH)This thesis focuses on the visual system in healthy subjects and schizophrenic patients. To address this research, advanced methods of analysis of electroencephalographic (EEG) data were used and developed. This manuscript is comprised of three scientific articles. The first article showed a novel method to control the physical features of visual stimuli (luminance and spatial frequencies). The second article showed, using electrical neuroimaging of EEG, a deficit in spatial processing associated with the dorsal pathway in chronic schizophrenic patients. This deficit was elicited by an absent modulation of the PI component in terms of response strength and topography as well as source estimations. This deficit was orthogonal to the preserved ability to process Kanizsa-type illusory contours. Finally, the third article resolved ongoing debates concerning the neural mechanism mediating illusory contour sensitivity by using electrical neuroimaging to show that the first differentiation of illusory contour presence vs. absence is localized within the lateral occipital complex. This effect was subsequent to modulations due to the orientation of misaligned grating stimuli. Collectively, these results support a model where effects in V1/V2 are mediated by "top-down" modulation from the LOC.To understand these three articles, the Introduction of this thesis presents the major concepts used in these articles. Additionally, a section is devoted to time-frequency analysis methods not presented in the articles themselves. The introduction is divided in four parts. The first part presents three aspects of the visual system: cellular, regional, and its functional interactions. The second part presents an overview of schizophrenia and its sensoiy-cognitive deficits. The third part presents an overview of illusory contour processing and the three models examined in the third article. Finally, advanced analysis methods for EEG are presented, including time- frequency methodology.The Introduction is followed by a synopsis of the main results in the articles as well as those obtained from the time-frequency analyses.Finally, the Discussion chapter is divided along three axes. The first axis discusses the time frequency analysis and proposes a novel statistical approach that is independent of the reference. The second axis contextualizes the first article and discusses the quality of the stimulus control and direction for further improvements. Finally, both neurophysiologic articles are contextualized by proposing future experiments and hypotheses that may serve to improve our understanding of schizophrenia on the one hand and visual functions more generally.

Restricted stimulus control in stimulus control shaping with a capuchin monkey

Teaching the first instances of arbitrary matching-to-sample to nonhumans can prove difficult and time consuming. Stimulus control relations may develop that differ from those intended by the experimentereven when stimulus control shaping procedures are used. We present, in this study, efforts to identify sources of shaping program failure with a capuchin monkey. Procedures began with a baseline of identity matching. During subsequent shaping trials, compound comparison stimuli had two componentsone identical to and another different from the sample. The identical component was eliminated gradually by removing portions across trials (i.e., subtracting stimulus elements). The monkey performed accurately throughout shaping. At a late stage in the program, probe tests were conducted: (1) arbitrary matching trials that had all elements of the identical comparison removed and (2) other trials that included residual elements. During the test, the monkey performed at low levels on the former trials and higher levels on the latter. These results suggested that higher accuracy was due merely to continued control by the residual elements: the target arbitrary matching relations had not been learned. Thus, it appears that procedures that gradually transform identity matching baselines into arbitrary matching can fail by inadvertently shaping restricted control by residual elements. Subsequent probes at the end of the shaping series showed a successful transfer of stimulus control from identity to arbitrary matching after further programming steps apparently overcame the restricted stimulus control.

OBSERVING BEHAVIOR AND ATYPICALLY RESTRICTED STIMULUS CONTROL

Restricted stimulus control refers to discrimination learning with atypical limitations in the range of controlling stimuli or stimulus features In the study reported here 4 normally capable individuals and 10 individuals with Intellectual disabilities (ID) performed two-sample delayed matching to sample Sample stimulus observing was recorded with an eye tracking apparatus High accuracy scores indicated stimulus control by both sample stimuli for the 4 nondisabled participants and 4 participants with ID and eye tracking data showed reliable observing of all stimuli Intermediate accuracy scores indicated restricted stimulus control for the remaining 6 participants Their eye tracking data showed that errors were related to failures to observe sample stimuli and relatively brief observing durations Five of these participants were then given interventions designed to improve observing behavior For 4 participants the interventions resulted initially in elimination of observing failures increased observing durations and Increased accuracy For 2 of these participants contingencies sufficient to maintain adequate observing were not always sufficient to maintain high accuracy subsequent procedure modifications restored It however For the 5th participant initial improvements in observing were not accompanied by improved accuracy in apparent Instance of observing without attending accuracy improved only after an additional intervention that imposed contingencies on observing behavior Thus interventions that control observing behavior seem necessary but may not always be sufficient for the remediation of restricted stimulus control

Operant measurement of auditory threshold in prelingually deaf users of cochlear implants: II

Two experiments evaluated an operant procedure for establishing stimulus control using auditory and electrical stimuli as a baseline for measuring the electrical current threshold of electrodes implanted in the cochlea. Twenty-one prelingually deaf children, users of cochlear implants, learned a Go/No Go auditory discrimination task (i.e., pressing a button in the presence of the stimulus but not in its absence). When the simple discrimination baseline became stable, the electrical current was manipulated in descending and ascending series according to an adapted staircase method. Thresholds were determined for three electrodes, one in each location in the cochlea (basal, medial, and apical). Stimulus control was maintained within a certain range of decreasing electrical current but was eventually disrupted. Increasing the current recovered stimulus control, thus allowing the determination of a range of electrical currents that could be defined as the threshold. The present study demonstrated the feasibility of the operant procedure combined with a psychophysical method for threshold assessment, thus contributing to the routine fitting and maintenance of cochlear implants within the limitations of a hospital setting.

Learning processes and the neural analysis of conditioning

Classical and operant conditioning principles, such as the behavioral discrepancy-derived assumption that reinforcement always selects antecedent stimulus and response relations, have been studied at the neural level, mainly by observing the strengthening of neuronal responses or synaptic connections. A review of the literature on the neural basis of behavior provided extensive scientific data that indicate a synthesis between the two conditioning processes based mainly on stimulus control in learning tasks. The resulting analysis revealed the following aspects. Dopamine acts as a behavioral discrepancy signal in the midbrain pathway of positive reinforcement, leading toward the nucleus accumbens. Dopamine modulates both types of conditioning in the Aplysia mollusk and in mammals. In vivo and in vitro mollusk preparations show convergence of both types of conditioning in the same motor neuron. Frontal cortical neurons are involved in behavioral discrimination in reversal and extinction procedures, and these neurons preferentially deliver glutamate through conditioned stimulus or discriminative stimulus pathways. Discriminative neural responses can reliably precede operant movements and can also be common to stimuli that share complex symbolic relations. The present article discusses convergent and divergent points between conditioning paradigms at the neural level of analysis to advance our knowledge on reinforcement.

Occasion setting in Pavlovian ambiguous target discriminations

Rats were trained in a Pavlovian serial ambiguous target discrimination, in which a target cue was reinforced if it was preceded by one stimulus (P -> T+) but was not reinforced if it was preceded by another stimulus (N -> T-). Test performance indicated that stimulus control by these features was weaker than that acquired by features trained within separate serial feature positive (P -> T+, T-) and serial feature negative (N -> W-, W+) discriminations. The form of conditioned responding and the patterns of transfer observed suggested that the serial ambiguous target discrimination was solved by occasion setting. The data are discussed in terms of the use of retrospective coding strategies when solving Pavlovian serial conditional discriminations, and the acquisition of special properties by both feature and target stimuli. (C) 2008 Published by Elsevier B.V.

Relational learning in children with deafness and cochlear implants

This four-experiment series sought to evaluate the potential of children with neurosensory deafness and cochlear implants to exhibit auditory-visual and visual-visual stimulus equivalence relations within a matching-to-sample format. Twelve children who became deaf prior to acquiring language (prelingual) and four who became deaf afterwards (postlingual) were studied. All children learned auditory-visual conditional discriminations and nearly all showed emergent equivalence relations. Naming tests, conducted with a subset of the: children, showed no consistent relationship to the equivalence-test outcomes.. This study makes several contributions: to the literature on stimulus equivalence. First; it demonstrates that both pre- and postlingually deaf children-can: acquire auditory-visual equivalence-relations after cochlear implantation, thus demonstrating symbolic functioning. Second, it directs attention to a population that may be especially interesting for researchers seeking to analyze the relationship. between speaker and listener repertoires. Third, it demonstrates the feasibility of conducting experimental studies of stimulus control processes within the limitations of a hospital, which these children must visit routinely for the maintenance of their cochlear implants.

Relational learning in children with deafness and cochlear implants

This four-experiment series sought to evaluate the potential of children with neurosensory deafness and cochlear implants to exhibit auditory-visual and visual-visual stimulus equivalence relations within a matching-to-sample format. Twelve children who became deaf prior to acquiring language (prelingual) and four who became deaf afterwards (postlingual) were studied. All children learned auditory-visual conditional discriminations and nearly all showed emergent equivalence relations. Naming tests, conducted with a subset of the: children, showed no consistent relationship to the equivalence-test outcomes.. This study makes several contributions: to the literature on stimulus equivalence. First; it demonstrates that both pre- and postlingually deaf children-can: acquire auditory-visual equivalence-relations after cochlear implantation, thus demonstrating symbolic functioning. Second, it directs attention to a population that may be especially interesting for researchers seeking to analyze the relationship. between speaker and listener repertoires. Third, it demonstrates the feasibility of conducting experimental studies of stimulus control processes within the limitations of a hospital, which these children must visit routinely for the maintenance of their cochlear implants.

Operant measurement of auditory threshold in prelingually deaf users of cochlear implants: II

Two experiments evaluated an operant procedure for establishing stimulus control using auditory and electrical stimuli as a baseline for measuring the electrical current threshold of electrodes implanted in the cochlea. Twenty-one prelingually deaf children, users of cochlear implants, learned a Go/No Go auditory discrimination task (i.e., pressing a button in the presence of the stimulus but not in its absence). When the simple discrimination baseline became stable, the electrical current was manipulated in descending and ascending series according to an adapted staircase method. Thresholds were determined for three electrodes, one in each location in the cochlea (basal, medial, and apical). Stimulus control was maintained within a certain range of decreasing electrical current but was eventually disrupted. Increasing the current recovered stimulus control, thus allowing the determination of a range of electrical currents that could be defined as the threshold. The present study demonstrated the feasibility of the operant procedure combined with a psychophysical method for threshold assessment, thus contributing to the routine fitting and maintenance of cochlear implants within the limitations of a hospital setting.

Análise comportamental das instruções e perguntas de uma professora universitária em sala de aula de uma turma de matemática

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Análise de contingências no ensino e na aprendizagem de descritores de desempenho escolar no ensino fundamental

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Um estudo sobre o conceito de prestar atenção na análise do comportamento de B. F. Skinner

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Evolução da nomeação após fortalecimento de relações auditivo-visuais em crianças com deficiência auditiva e implante coclear

Pós-graduação em Psicologia do Desenvolvimento e Aprendizagem - FC

Procedimentos de treino e teste de relações entre estímulos em Cebus apella

A aprendizagem complexa abrange o controle por relações abstratas após a exposição a relações diretamente treinadas. Neste trabalho, uma seqüência de experimentos investigou a aprendizagem de relações entre estímulos em dois macacos prego (Cebus apella) envolvendo o treino direto de relações de identidade, testes sucessivos de identidade generalizada, o treino direto de relações arbitrárias e teste de relações simétricas como indicação da formação de classes de equivalência. Três bases teóricas foram seguidas: 1) A formação de classes de equivalência é uma função básica de estímulo 2) A teoria da coerência de topografia de controle de estímulos, e 3) Uma versão do sistema personalizado de instrução como uma alternativa para estabelecer repertórios relacionais em organismos não-verbais. Os sujeitos apresentaram altos níveis de desempenho em discriminações condicionais de identidade e arbitrárias. Os sujeitos expostos a testes de identidade generalizada demonstraram essa relação abstrata; o sujeito exposto ao treino arbitrário apresentou resultados negativos no teste de desempenhos simétricos. A maioria dos problemas de aprendizagem de relações entre estímulos encontrados aqui com não-humanos parece similar aos problemas apresentados por indivíduos com desenvolvimento retardado. Os procedimentos de ensino para indivíduos com retardo foram suficientes para solucionar grande parte das deficiências de repertório de não-humanos, as quais geralmente são atribuídas a limitações da espécie.

Relações ordinais sob controle contextual em crianças surdas

Dificuldades apresentadas por crianças surdas na aprendizagem da matemática têm conduzido educadores ao desenvolvimento de procedimentos especiais de ensino. O paradigma de equivalência tem sido útil na explicação de comportamentos complexos, como comportamentos conceituais numéricos. Uma expansão desse paradigma envolve a formação de classes de estímulos equivalentes em seqüência. A emergência de novas relações através do responder ordinal já foi documentada em estudos com contingências de reforçamento de três termos. Há necessidade de verificar se esses resultados se mantém estáveis sob contingências de quatro e cinco termos. Três experimentos foram programados com o objetivo de investigar a emergência de relações ordinais com controle discriminativo simples, sob controle condicional (sem e com randomização das tentativas) e sob controle contextual em crianças surdas. No Experimento 1 participaram cinco crianças surdas, matriculadas numa Escola Pública Especializada. Um microcomputador com um software (REL 4.0, utilizado nos Experimentos 1, 2a e 2b e atualizado para a versão 5.0 no Estudo 3) foi utilizado. Nesse estudo foi ensinado aos participantes seqüências de pares de estímulos sobrepostos. Em seguida, foram realizados testes de transitividade e conectividade. Todos os participantes alcançaram o critério de acerto e responderam aos testes. Os resultados replicaram estudos da literatura confirmando a eficiência do procedimento de ensino por sobreposição de estímulos no estabelecimento de relações ordinais. No Experimento 2a, quatro novos participantes e um com história experimental, foram ensinados a selecionar estímulos, aos pares, na ordem crescente na presença da cor verde e na ordem decrescente na presença da cor vermelha. Foram aplicados testes de transitividade e conectividade sob controle condicional. Em seguida foi conduzido um teste de generalização com estímulos do ambiente escolar. Todos os participantes alcançaram o critério de acerto e responderam aos testes de transitividade e conectividade. Nos testes de generalização, três participantes responderam consistentemente aos novos estímulos, um respondeu parcialmente e um não respondeu ao teste. Os resultados corroboraram a eficiência do procedimento de ensino por sobreposição de estímulos sob controle condicional em crianças surdas. O Experimento 2b envolveu os mesmos participantes do Experimento 1 com história experimental e a randomização das tentativas com os estímulos condicionais. Todos os participantes alcançaram o critério de acerto. Nos testes demonstraram um responder consistente com a linha de base. No Experimento 3 participaram três crianças dos Experimento 1 e duas do Experimento 2a, que foram expostas ao procedimento de ensino por pares sobrepostos sob controle contextual de duas formas círculo e triângulo e sob controle condicional das cores verde e vermelha (ex. A1A2, na presença do círculo e da cor verde; ou A2A1, na presença do círculo e da cor vermelha). Todos os participantes alcançaram o critério de acerto e responderam aos testes de transitividade e conectividade. Os resultados indicaram a eficiência do procedimento de ensino por sobreposição de estímulos sob controle contextual, sugerindo que o ensino por contingências de reforçamento simples e sob controle condicional foram pré-requisitos para a emergência de classes ordinais sob controle contextual. Uma extensão deste estudo deve ampliar o número de membros na seqüência e investigar a emergência de novas relações ordinais com seqüências mais longas, e verificar se a ordem de treino em que a seqüência é ensinada interfere sobre o responder ordinal.