Lívia Cuquetto Leite

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Lívia Cuquetto Leite Lívia Cuquetto Leite “MORFOLOGIA DO APARELHO REPRODUTOR MASCULINO E DOS ESPERMATOZOIDES DE Pachycondyla (HYMENOPTERA: FORMICIDAE: PONERINAE)” “MORPHOLOGY OF THE MALE REPRODUCTIVE APPARATUS AND THE SPERMATOZOA OF Pachycondyla (HYMENOPTERA: FORMICIDAE: PONERINAE)” Campinas, 2015 Dedico este trabalho a meus grandes amigos Amanda, Isadora e Marllonn (in memoriam) e a meus pais, que sempre me apoiaram e guiaram minhas escolhas. “Persiga um sonho, mas não deixe ele viver sozinho. Descubra-se todos os dias, deixe-se levar pelas vontades, mas não enlouqueça por elas. Procure, sempre procure o fim de uma história, seja ela qual for. Dê um sorriso para quem esqueceu como se faz isso. Acelere seus pensamentos, mas não permita que eles te consumam. Olhe para o lado, alguém precisa de você. Abasteça seu coração de fé, não a perca nunca. Mergulhe de cabeça nos seus desejos e satisfaça-os. Procure os seus caminhos, mas não magoe ninguém nessa procura. Arrependa-se, volte atrás, peça perdão! Não se acostume com o que não o faz feliz, revolte-se quando julgar necessário. Alague seu coração de esperanças, mas não deixe que ele se afogue nelas. Se achar que precisa voltar, volte! Se perceber que precisa seguir, siga! Se estiver tudo errado, comece novamente. Se estiver tudo certo, continue. Se sentir saudades, mate-a. Se perder um amor, não se perca! Se achá-lo, segure-o!” (Fernando Pessoa) AGRADECIMENTOS Meus sinceros agradecimentos: À Deus, primeiramente, por me permitir concluir esta vitória e ter paciência nos momentos difíceis, sempre guiando minha vida e norteando minhas decisões. À meus pais, Edilmar Leite e Rosana Cuquetto Leite, exemplos de amor, força e sabedoria, que desde minha infância me incentivaram a estudar e dar o melhor de mim. À minha querida orientadora Dr.ª Karina Mancini. Muito obrigada por me orientar nessa jornada com sua dedicação, compreensão, amizade e paciência nos momentos mais críticos. Karina, sou muito grata a todo conhecimento compartilhado. À adorável Drª. Mary Anne Heidi Dolder, que, com sua paciência me coorientou e auxiliou nos momentos mais necessários, especialmente no manuseio do Microscópio Eletrônico de Transmissão. Ao professor Dr. Paulo S. Oliveira e seus alunos, que me auxiliam em inúmeras coletas realizadas. Muito obrigada pelos artigos e conselhos de campo. À meu querido amigo Michel, que mesmo estando distante compartilhou de minhas angústias e alegrias. Muito obrigada também pelas loucuras nossas e cumplicidade. Às queridíssimas Bruna e Celina, que me acolheram de braços abertos na UNICAMP, me ajudando na rotina do laboratório e me mostrando a vida de Campinas. Ao Laboratório de Ecologia e Sistemática de Borboletas e ao Mário, pelo auxílio técnico e de cladística. Ao programa de pós-graduação em Biologia Celular e Estrutural e à agência de fomento CNPq, por todo suporte técnico, científico e financeiro oferecido nesses dois anos. Em suma, a todos que contribuíram direta e indiretamente para a conquista deste trabalho. Muito Obrigada! RESUMO Apesar de Formicidae constituir um dos grupos animais mais significativos em riqueza de espécies no Brasil, suas relações filogenéticas permanecem controversas. Uma notável fonte de caracteres para análises taxonômicas e filogenéticas é a caracterização espermática e do aparelho reprodutor. Tal caracterização tem se mostrado uma ferramenta na clarificação de questões da biologia reprodutiva e sistemática de diversos grupos de insetos. Desta forma, o presente estudo analisou, através de microscopia de luz e eletrônica de transmissão, a morfologia espermática de quatro espécies do gênero Pachycondyla (Formicidae: Ponerinae), bem como o aparelho reprodutor masculino: P. marginata, P. termitofane comutata, P. harpax e P. striata. A ultraestrutura dos espermatozoides de tais espécies é semelhante à descrita para outros Hymenoptera por apresentar: (1) acrossomo cônico do tipo bicamada; (2) adjunto do centríolo situado entre o núcleo e ambos os derivados mitocondriais e (3) flagelo constituído por dois derivados mitocondriais alongados, dois corpos acessórios triangulares e um axonema de padrão 9+9+2. Entretanto, as mesmas compartilham inúmeras características cujas evidências indicam evoluir como homoplasia via paralelismo, não conferindo, pois, suporte para uma resolução filogenética entre tais espécies e demais clados próximos. Dentre estes caracteres se destacam a organização espacial dos componentes acrossomais, o caráter cromatínico nuclear, a configuração da extremidade da transição núcleo-flagelo, além da condição simétrica do adjunto do centríolo. Além dessas, várias outras características corroboram a definição do estado parafilético do respectivo gênero, tais como o aspecto cromatínico, a condição simétrica do adjunto do centríolo e a organização terminal flagelar, por serem compartilhadas entre determinadas espécies de Pachycondyla e grupos filogeneticamente distantes, evidenciando, pois, as divergências dentro da mesma espécie. Por fim, a análise morfológica do aparelho reprodutor de P. marginata e P. striata evidenciou a existência de um padrão morfológico reprodutivo para Formicidae, uma vez que tais espécies exibiram íntima similaridade às descrições realizadas à respectiva família. Palavras-chave: Ultraestrutura espermática, Morfologia reprodutiva, Ponerini. ABSTRACT Although Formicidae be one of the most important animal groups in species richness in Brazil, their phylogenetic relationships remain controversial. One notable source of characters for taxonomic and phylogenetic analysis is the characterization of the spermatozoa and male reproductive tract. This characterization has shown to be a tool in clarifying issues of reproductive biology and systematics of many insect groups. Thus, this study analyzed by light and transmission electron microscopies, the sperm morphology of four species of the genus Pachycondyla (Formicidae: Ponerinae), and their male reproductive system: P. marginata, P. termitofane comutata, P. harpax and P. striata. The ultrastructure of spermatozoa of these four species is similar to that described to other Hymenoptera: (1) bilayer tapered acrosome; (2) centriolar adjunct located between nucleus and both mitochondrial derivatives and (3) flagellum comprises two elongate mitochondrial derivatives, two triangular bodies and an axoneme of 9 + 9 + 2 pattern. However, they share a number of features whose evidence indicates evolve as homoplasy via parallelism, not giving, support for a phylogenetic resolution among species and other nearby clades. Among these characters stand out, the spatial organization of acrosomal components, the nuclear chromatin character, setting the end of the transition nucleus-flagellum, in addition to symmetrical condition centriolar adjunct. Besides these, various other features confirm the setting of the paraphyletic status of its genus, such as chromatin aspect, symmetrical condition centriolar adjunct and flagellar terminal organization, by being shared among certain species phylogenetically distant and Pachycondyla groups, highlighting, therefore, the differences within the same species. Finally, the morphological analysis of the reproductor system of P. striata and P. marginata highlighted the existence of a reproductive morphology pattern to Formicidae, since such species showed close similarity to the descriptions made to its family. Keywords: Sperm Ultrastructure, Reproductive Morphology, Ponerini. SUMÁRIO 1. Introdução ........................................................................................................................... 12 1.1. Ordem Hymenoptera .................................................................................................... 12 1.1.1. Composição e Filogenia ........................................................................................... 12 1.1.2. Relevância Ecológica ............................................................................................. 14 1.2. Formicidae ..................................................................................................................... 15 1.3. Ponerinae........................................................................................................................ 19 1.4. Aparelho Reprodutor Masculino e Espermatozoides em Hymenoptera e sua Relevância Filogenética ...................................................................................................... 23 1.4.1. Morfologia do Aparelho Reprodutor Masculino...................................................... 23 1.4.2. Espermatozoides de Hymenoptera ........................................................................... 24 1.4.3. Relevância da Morfologia Espermática ................................................................... 26 2. Objetivos ............................................................................................................................... 28 2.1. Objetivo Geral .............................................................................................................. 28 2.2. Objetivos Específicos .................................................................................................... 28 3. Referências Bibliográficas ................................................................................................... 29 Capítulo 1 - Características ultraestruturais dos espermatozoides de Pachycondyla harpax e Pachycondyla termitofane commutata (Hymenoptera: Formicidae) ...................................... 35 Capítulo 2 - Structural and Ultrastructural Characterization
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