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Instituto Nacional De Pesquisas Da Amazônia – Inpa Programa De Pós-Graduação Em Biologia (Ecologia) – Ppg-Eco Diferença INSTITUTO NACIONAL DE PESQUISAS DA AMAZÔNIA – INPA PROGRAMA DE PÓS-GRADUAÇÃO EM BIOLOGIA (ECOLOGIA) – PPG-ECO DIFERENÇAS INTER E INTRASSEXUAIS NA APRENDIZAGEM ESPACIAL DE UM PEIXE AMAZÔNICO SEXUALMENTE DIMÓRFICO KALEBE DA SILVA PINTO MANAUS-AM, AMAZONAS FEVEREIRO, 2020 KALEBE DA SILVA PINTO DIFERENÇAS INTER E INTRASSEXUAIS NA APRENDIZAGEM ESPACIAL DE UM PEIXE AMAZÔNICO SEXUALMENTE DIMÓRFICO Orientador: Dr. Jansen Zuanon Coorientador: Dr. Tiago Henrique da Silva Pires Dissertação apresentada ao Instituto Nacional de Pesquisas da Amazônia como parte dos requisitos para obtenção do título de Mestre em Biologia (Ecologia). MANAUS-AM, AMAZONAS FEVEREIRO, 2020 ii iii Sinopse: Neste estudo, analisamos diferenças sexuais na aprendizagem espacial em um peixe amazônico sexualmente dimórfico (Crenuchus spilurus). Também investigamos como características morfológicas (ornamentos) e traços de personalidade (neofobia) afetam o desempenho individual na tarefa espacial. Além disso, testamos como a escolha feminina atua na seleção de machos com melhor desempenho em tarefas de aprendizagem espacial. Palavras-chave: Aprendizagem espacial, cognição, demanda conflitante, diferenças sexuais, escolha de parceiros, labirinto complexo, neofobia, peixe, traços de personalidade. iv Project Cecília v Agradecimentos Aos meus orientadores Jansen Zuanon e Tiago Pires, pela confiança depositada em mim e pelo incentivo para realizar este trabalho. Agradeço ao PPG-ECO/INPA pela oportunidade de realizar este trabalho; ao CNPq e JICA/JST pelos financiamentos de bolsa e equipamentos; e ao PPG- CIPET/UFAM pelo espaço físico para realização dos experimentos. A todos os membros do Laboratório de Ecologia Comportamental e Evolutiva - LECE pela ajuda na manutenção e cuidados com os peixes utilizados nesse trabalho. Agradeço ao Élio Borghezan, Adriely Melo e Álvaro Lima pela ajuda nos experimentos. Ao David Saenz pela sua imensa contribuição pessoal com este trabalho. Agradeço ao Tiago Pires por ter me dado um grande apoio pessoal fora do laboratório. Mais que um orientador, um grande amigo. Agradeço à Waleska Barbosa pela confiança, paciência e companheirismo durante todo o período em que realizei esse trabalho. Também agradeço pelo seu apoio emocional e familiar que foi essencial para chegar até aqui. Um agradecimento à minha família, por todo incentivo dado a buscar meus sonhos e objetivos. Por fim, agradeço a todos que de alguma forma contribuíram com este trabalho e com meu crescimento acadêmico e profissional. vi “Try not. Do... or do not. There is no try.” Yoda vii Resumo Os processos individuais de adquirir, armazenar e usar informações para resolver tarefas cognitivas podem variar entre os indivíduos de uma mesma espécie. Isso ocorre, por exemplo, quando existem variações comportamentais consistentes, decorrentes de dimorfismo sexual ou por variações morfológicas dentro de um único sexo. Isso pode gerar diferentes suscetibilidades à predação em indivíduos mais ornamentados, ou preferência por indivíduos com melhores habilidades cognitivas, o que contribui para a evolução da cognição. Nesse contexto, quando pressões seletivas atuam de maneira distinta entre os indivíduos, é esperado o surgimento de variações na tomada de decisões ecologicamente relevantes. Adicionalmente, é sugerida uma ligação funcional entre a variação individual cognitiva e a personalidade animal, gerando demandas conflitantes entre precisão e velocidade de decisão. Neste estudo, investigamos a neofobia, o aprendizado espacial e a escolha feminina no peixe amazônico Crenuchus spilurus, uma espécie sexualmente dimórfica. Não encontramos diferença sexual na taxa de aprendizagem da tarefa espacial. Contudo, quando avaliadas as diferenças intrassexuais, machos mais ornamentados mostraram melhor desempenho na tarefa, apresentando maior precisão para resolver o labirinto no último dia de testes do que machos menos ornamentados e fêmeas em geral. Machos mais ornamentados são mais neofóbicos e levaram mais tempo para resolver a tarefa espacial na primeira tentativa. Isso sugere que esses machos se comportam de acordo com o princípio de proteção de ativos, quando machos mais atraentes devem ser mais cautelosos como resultado de maiores retornos futuros decorrentes de suas características físicas. Além disso, fêmeas mostraram preferência por machos mais ornamentados, que foram os indivíduos que apresentaram melhor desempenho na tarefa espacial. Com estes resultados, propomos que a diferença sexual na cognição pode emergir da variação intrassexual, causada pela pressão seletiva que ocorre sobre machos mais ornamentados. Adicionalmente, como o princípio de proteção de ativos decorre da pressão de predação, a seleção sexual por meio da escolha feminina proporciona uma troca de recompensas e riscos, sobre a qual a seleção natural atua indiretamente aumentando habilidades de aprendizado. viii Abstract The individual processes of acquisition, retention and use of information to solve cognitive tasks can vary between individuals. Such cognitive variation may be explained by consistent behavioral differences, resulting from sexual dimorphism or morphological variations within a single sex. This can generate different susceptibilities to predation in individuals with higher degree of ornamentation, and/or mating preference for individuals with enhanced cognitive skills, which contributes to the evolution of cognition. In that context, when selective pressures act differently among individuals, variations in ecologically relevant decision-making are expected. Additionally, a functional link between individual cognitive variation and animal personality is suggested, generating speed-accuracy trade-offs. In a further step, we investigated the relationship between spatial learning to male ornamental degree and neophobia and finally female mate choice, in the Amazonian fish Crenuchus spilurus, a sexually dimorphic species. We found no sex differences in the learning rate of the spatial task. However, when assessing males’s individual differences, those with higher degree of ornamentation showed better performance in the learning task, displaying greater precision to solve the maze on the last day of test. The performance of these males was better than males with lower degree of ornamentation and females. Males with a higher degree of ornamentation were more neophobic and took longer to resolve the spatial task on the first attempt. This suggests that males with a higher degree of ornamentation behave according to the asset-protection principle, which proposes that more attractive males should be more cautious as a result of higher future returns consequences of their physical characteristics. In addition, females showed preference for males with higher degree of ornamentation, which consequently, were also the individuals who showed better performance in learning the spatial task. We propose that the sex differences in cognition can emerge from the intrassexual variation, caused by the selective pressure that occurs in males with higher degree of ornamentation. Additionally, as the asset-protection principle stems from the pressure of predation, sexual selection through female mate choice provides an exchange of rewards and risks, on which natural selection acts indirectly by increasing learning skills. ix Sumário Lista de figuras ......................................................................................................... xi Introdução geral ...................................................................................................... 13 Objetivos .................................................................................................................. 16 Geral ...................................................................................................................... 16 Específicos ............................................................................................................. 16 Capítulo 1 ................................................................................................................. 17 Resumo .................................................................................................................... 18 Introdução ................................................................................................................ 19 Material e métodos .................................................................................................. 20 Modelo animal ........................................................................................................ 20 Medição de características morfológicas ............................................................... 21 Aparato experimental ............................................................................................. 21 Procedimento experimental ................................................................................... 22 Análises estatísticas ............................................................................................... 23 Resultados ............................................................................................................... 24 Discussão ................................................................................................................ 26 Financiamento ....................................................................................................... 30 Agradecimentos ..................................................................................................... 30 Referências .............................................................................................................
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