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Universidade Federal Do Rio Grande Do Norte Centro De UNIVERSIDADE FEDERAL DO RIO GRANDE DO NORTE CENTRO DE BIOCIÊNCIAS PROGRAMA DE PÓS-GRADUAÇÃO EM PSICOBIOLOGIA CLARISSA DE ALMEIDA MOURA APRENDIZAGEM ESPAÇO-TEMPORAL E EFEITOS DAS CONDIÇÕES LUMINOSAS NO APRENDIZADO DO PEIXE PAULISTINHA NATAL 2016 1 CLARISSA DE ALMEIDA MOURA APRENDIZAGEM ESPAÇO-TEMPORAL E EFEITOS DAS CONDIÇÕES LUMINOSAS NO APRENDIZADO DO PEIXE PAULISTINHA DEFESA DA DISSERTAÇÃO APRESENTADA AO PROGRAMA DE PÓS-GRADUAÇÃO EM PSICOBIOLOGIA DA UNIVERSIDADE FEDERAL DO RIO GRANDE DO NORTE, COMO REQUISITO PARA A OBTENÇÃO DO TÍTULO DE MESTRE EM PSICOBIOLOGIA (ÁREA: COMPORTAMENTO ANIMAL). Orientadora: Profa. Dra. Ana Carolina Luchiari NATAL 2016 2 Catalogação da Publicação na Fonte. UFRN / Biblioteca Setorial do Centro de Biociências Moura, Clarissa de Almeida. Aprendizagem espaço-temporal e efeitos das condições luminosas no aprendizado do peixe paulistinha / Clarissa de Almeida Moura. – Natal, RN, 2016. 86 f.: il. Orientadora: Profa. Dra. Ana Carolina Luchiari. Dissertação (Mestrado) – Universidade Federal do Rio Grande do Norte. Centro de Biociências. Programa de Pós-Graduação em Psicobiologia. 1. Aprendizagem espaço-temporal. – Dissertação. 2. Memória. – Dissertação. 3. Ritmo circadiano. – Dissertação. I. Luchiari, Ana Carolina. II. Universidade Federal do Rio Grande do Norte. III. Título. RN/UF/BSE-CB CDU 159.953 3 APRENDIZAGEM ESPAÇO-TEMPORAL E EFEITOS DAS CONDIÇÕES LUMINOSAS NO APRENDIZADO DO PEIXE PAULISTINHA CLARISSA DE ALMEIDA MOURA Natal, 03 de março de 2016. Banca Avaliadora ______________________________ Profª Dra. Ana Carolina Luchiari Departamento de Fisiologia- UFRN Orientadora ____________________________________ Prof. Dr. Mário André Leocadio Miguel Departamento de Fisiologia- UFRN Membro Interno ____________________________________ Prof. Dr. Rodrigo Egydio Barreto Departamente de Fisiologia – UNESP Botucatu Membro externo 4 AGRADECIMENTOS Ao mestre Jesus por guiar meus passos e iluminar minha mente e minhas mãos, me fazendo pensar e agir de maneira sensata em cada detalhe dessa pesquisa. À minha orientadora Ana Carolina Luchiari, que acreditou em meu potencial, abriu as portas para meu caminho na ciência, me preparou não só como pesquisadora, mas como futura docente, e não mediu esforços para me ajudar sempre que precisei. Às minhass amigas e companheiras de laboratório, Bábara, Mayara, Priscilla e Diana por toda força, palavras de carinho e incetivo nas horas certas. Aos meus ajudantes na pesquisa, Vanessa, Jéssica Pollyana, Mara, Celisa, Fred e Joanny por cada tempo dedicado e esforço em me ajudar com a coleta de dados. Sem vocês, esse trabalho jamais estaria completo! À Bruna Del Vechio, por ajudar nas correções e com os conhecimentos da Cronobiologia, e ao professor Mário Miguel, pela sua disponibilidade e cordialidade em me ajudar com as análises. À Tales, meu companheiro no amor e nos estudos, por sua companhia nos finais de semana de coleta, dias de estudos, e sua ajuda com os gráficos. À minha família, que por torcer tanto pelo meu sucesso, não medem esforços para me ver alcançar os objetivos. Mãe, pela paciência e preocupação. Pai, pelo esforço em ajudar e incentivo para que eu jamais desistisse. E, antes de tudo, pela educação que vocês me deram, pois para mim, não há nada de mais precioso na vida! 5 RESUMO GERAL Para processar a informação ambiental e perceber o tempo, os indivíduos utilizam-se de pistas ambientais, como luz e temperatura, que servem como guias para o relógio interno. O mecanismo temporizador endógeno é chamado relógio circadiano endógeno, o qual comanda uma grande variedade de ritmos diários bioquímicos, fisiológicos e comportamentais presentes nos organismos. Com isso, os animais podem antecipar eventos espaço-temporalmente distribuídos e usar essa informação para organizar as atividades diárias, o que é uma vantagem adaptativa para os indivíduos, já que muitos fatores ambientais apresentam variação circadiana. Aprendizagem espaço-temporal (do inglês: "time-place learning’’-TPL) é a habilidade de associar lugares com importantes eventos biológicos em diferentes horas do dia. Em nosso estudo utilizamos como modelo o peixe paulistinha (Danio rerio), conhecido por ser altamente social, para testar aprendizagem espaço-temporal baseada em reforço social. Além disso, objetivamos averiguar os efeitos das condições de claro constante e escuro constante na aprendizagem espaço-temporal, e se nessas condições, a atividade do peixe paulistinha é alterada. Para isso, testamos três diferentes condições (n=10): grupo claro-escuro (CE), grupo claro constante (CC) e grupo escuro constante (EE) durante 30 dias da seguinte maneira: diariamente, um grupo de 5 peixes paulistinha foi introduzido em um recipiente localizado no compartimento da manhã (um dos lados do aquário), às 8:00h e retirado às 9:00h, e em outro recipiente do compartimento da tarde (lado oposto do aquário), às 17:00h e removido às 18:00h, servindo como estímulo para que o peixe experimental ocupasse o compartimento onde o grupo fosse colocado. O comportamento foi filmado nos dois horários, 15 minutos antes e durante os 60 minutos de exposição ao estímulo, no 15º e no 30ª dia, porém neste último, os peixes foram filmados sem a presença do estímulo a fim de averiguarmos a aprendizagem espaço- temporal. Por fim, para saber a influência das três condições luminosas na atividade dos peixes, filmamos os últimos 6 dias de teste, para registrar o padrão de atividade. Nossos resultados mostraram que em ciclo claro-escuro (CE) o peixe paulistinha apresenta TPL, bem como é capaz de antecipar a hora e local do estímulo (grupo de coespecíficos), enfatizando a importância do estímulo social para a aprendizagem. Em condições de claro constante e escuro constante, o peixe paulistinha não apresentou aprendizagem espaço-temporal. Ademais, após 30 dias em condições luminosas constantes (claro constante e escuro constante), o peixe paulistinha mantém ritmo circadiano, porém em claro constante sua atividade é aumentada e seu ritmo atividade-repouso é alterado, através de um padrão de atividade distribuída homogeneamente ao longo das 24h, ao invés de concentrada na subjetiva fase clara, como nos grupos de ciclo claro-escuro e escuro constante, os quais conservam o padrão de atividade diurno da espécie. Palavras chave: aprendizagem espaço-temporal, memória, estímulo social, ritmo circadiano. 6 SUMÁRIO 1. INTRODUÇÃO GERAL...................................................................................... 8 Sistema temporizador endógeno e sincronizadores.................................................. 8 Ritmos em peixes...................................................................................................... 11 Ritmo no peixe paulistinha........................................................................................ 12 Pistas sociais como sinalizadores.............................................................................. 14 Aprendizagem espaço-temporal................................................................................ 15 2. OBJETIVOS ........................................................................................................ 20 Objetivo Geral........................................................................................................ 20 Objetivos Específicos ............................................................................................... 20 3. ARTIGO 1.......................................................................................................... 21 ABSTRACT .......................................................................................................... 21 INTRODUCTION..................................................................................................... 22 MATERIAL AND METHODS................................................................................ 23 Animals and procedures............................................................................................ 23 Experimental design……………………………………………………………….. 24 Statistical analysis..................................................................................................... 26 RESULTS................................................................................................................. 26 Responsiveness on day 15......................................................................................... 26 Determination of learning on day 30........................................................................ 27 DISCUSSION........................................................................................................... 29 REFERENCES........................................................................................................... 33 4. ARTIGO 2.............................................................................................................. 45 ABSTRACT............................................................................................................... 45 INTRODUCTION...................................................................................................... 46 MATERIAL AND METHODS.................................................................................. 48 Animals and procedures............................................................................................. 48 Experimental design................................................................................................... 48 Activity registry.........................................................................................................
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