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JÚLIO DE MESQUITA FILHO” Unesp INSTITUTO DE BIOCIÊNCIAS – RIO CLARO UNIVERSIDADE ESTADUAL PAULISTA “JÚLIO DE MESQUITA FILHO” unesp INSTITUTO DE BIOCIÊNCIAS – RIO CLARO PROGRAMA DE PÓS-GRADUAÇÃO EM CIÊNCIAS BIOLÓGICAS (BIOLOGIA CELULAR E MOLECULAR) A glândula metapleural e suas secreções em formigas cultivadoras (Attini) e não cultivadoras de fungos (Myrmicini, Blepharidattini e Ectatommini) (Hymenoptera: Formicidae) Alexsandro Santana Vieira Tese apresentada ao Instituto de Biociências do Campus de Rio Claro, Universidade Estadual Paulista, como parte dos requisitos para obtenção do título de Doutor em Ciências Biológicas (Biologia Celular e Molecular). Rio Claro Julho - 2012 Alexsandro Santana Vieira A glândula metapleural e suas secreções em formigas cultivadoras (Attini) e não cultivadoras de fungos (Myrmicini, Blepharidattini e Ectatommini) (Hymenoptera: Formicidae) Tese apresentada ao Instituto de Biociências do Campus de Rio Claro, Universidade Estadual Paulista Júlio de Mesquita Filho, como parte dos requisitos para obtenção do título de Doutor em Ciências Biológicas (Biologia Celular e Molecular). Orientadora: Profa. Dra. Maria Izabel Souza Camargo Co-orientador: Prof. Dr. Odair Corrêa Bueno Rio Claro Julho - 2012 595.796 Vieira, Alexsandro Santana V657g A glândula metapleural e suas secreções em formigas cultivadoras (Attini) e não cultivadoras de fungos (Myrmicini, Blepharidattini e Ectatommini) (Hymenoptera: Formicidae) / Alexsandro Santana Vieira. - Rio Claro : [s.n.], 2012 188 f. : il., figs., gráfs., tabs. Tese (doutorado) - Universidade Estadual Paulista, Instituto de Biociências de Rio Claro Orientador: Maria Izabel Souza Camargo Co-Orientador: Odair Corrêa Bueno 1. Formiga. 2. Formigas cortadeiras. 3. Histoquímica. 4. Ultraestrutura. 5. Cromatografia em fase gasosa. 6. Compostos antibióticos. I. Título. Ficha Catalográfica elaborada pela STATI - Biblioteca da UNESP Campus de Rio Claro/SP i Aos meus pais Antonio Santana Vieira e Maria José Santana, que me deram a vida e me ensinaram a vivê-la com dignidade e dedicação para o trabalho. Aos meus irmãos Admilson Santana Vieira e André Santana Vieira pelo apoio moral e, à minha noiva Fernanda Carafa, que muitas vezes foi compreensiva aos meus atos. Dedico esta tese. ii Agradecimentos Agradecimento especial à (Fundação de Amparo à Pesquisa do Estado de São Paulo), pelo apoio financeiro concedido, o qual possibilitou a realização deste trabalho (Processo nº 08/54301-3). À minha orientadora, Bel (Maria Izabel Souza Camargo), a quem devo meu progresso científico, de quem vou guardar os seus ensinamentos pelo resto da vida. Agradeço de forma muito especial por me aceitar como orientado, desde o primeiro contato, e pela sua sabedoria, incentivo e exemplo de coragem e trabalho. Ao meu co-orientador, Odair Corrêa Bueno, de quem vou guardar as suas discussões científicas, bem como sua sabedoria e incentivo de ousadia na pesquisa. À todos os docentes do Departamento de Biologia da UNESP de Rio Claro por estarem sempre dispostos ao esclarecimento de dúvidas. Aos técnicos de laboratório Antonio Teruyoshi Yabuki, Gerson Mello Souza, Mônica Iamonte pelos serviços técnicos prestados e amizade construída durante todos estes anos. À todos os funcionários do Departamento de Biologia, Seção de Pós- Graduação e Biblioteca pela ajuda concedida. À todos os meus amigos de pós-graduação, companheiros de todas as horas, pela grande amizade construída durante todos estes anos, Luís Flávio da Silva Costa, Pablo Henrique Nunes, Sandra Denardi, Gabriela Ortiz, Érika, Patrícia, Gislaine e Karim pela confiança, dedicação e atenção, pelo companheirismo, por dividirem comigo suas alegrias e tristezas. E às pessoas fantásticas que passaram em meu caminho nestes anos, inclusive E. D. Morgan que foi meu supervisor do estágio realizado na Keele University (England-UK), auxiliou no estudo cromatográfico em fase gasosa da composição química da secreção das glândulas metapleurais. Meu muito obrigado! iii SUMÁRIO RESUMO E ABSTRACT ................................................................................................. 1 I. INTRODUÇÃO ............................................................................................................. 4 II. OBJETIVOS ............................................................................................................... 11 III. MATERIAL E MÉTODOS ....................................................................................... 13 III. 1. Análise Morfológica .............................................................................................. 15 III. 1.2. Microscopia Eletrônica de Varredura (MEV) .................................................... 15 III. 1.3. Técnica da Hematoxilina de Harris – Eosina Aquosa ........................................ 15 III. 1.4. Morfometria ........................................................................................................ 15 III. 2. Análise Histoquímica ............................................................................................ 15 III. 2.1. Técnica do Azul de Bromofenol para detecção de proteínas ............................. 15 III. 2. 2. Técnica do PAS/Azul de Alcian para detecção de polissacarídeos ................... 16 III. 2. 3. Técnica do PAS/Verde Metila para detecção polissacarídeos e RNA .............. 16 III. 2. 4. Técnica do Azul de Nilo para detecção de lipídios ácidos ................................ 16 III. 2. 5. Técnica de Baker para detecção de lipídios totais ............................................. 17 III. 2. 6. Variante de CEC (Critical Electrolyte Concentration) para detecção de DNA e RNA ................................................................................................................................. 17 III. 3. Análise Ultraestrutural ........................................................................................... 17 III. 3. 1. Microscopia Eletrônica de Transmissão (MET) ................................................ 17 III. 4. Análise Química da Secreção ................................................................................ 18 III. 4. 1. Cromatografia em fase Gasosa – Espectrometria de Massas ........................... 18 IV. RESULTADOS ......................................................................................................... 20 CAPÍTULO 1 ................................................................................................................. 23 The functional morphology of the metapleural gland of the leaf-cutting ant Atta laevigata (Formicidae: Attini) CAPÍTULO 2 ................................................................................................................. 34 Ultrastructural profile of metapleural gland cells of the ant Atta laevigata (F. Smith, 1858) (Formicidae: Attini) CAPÍTULO 3 ................................................................................................................. 47 Secretory profile of metapleural gland cells of the leaf-cutting ant Acromyrmex coronatus (Formicidae: Attini) iv CAPÍTULO 4 ................................................................................................................. 57 Morphophysiological differences between the metapleural glands of fungus-growing and non-fungus-growing ants (Hymenoptera, Formicidae) CAPÍTULO 5 ................................................................................................................. 96 Comparative ultrastructure of the metapleural glands of fungus-growing (Attini) and non-fungus-growing ants (Blepharidattini and Ectatommini) (Hymenoptera: Formicidae) CAPÍTULO 6 ............................................................................................................... 132 Chemical composition of metapleural glands secretions of fungus-growing and non– fungus-growing ants (Hymenoptera, Formicidae) V. DISCUSSÃO GERAL .............................................................................................. 165 VI. CONCLUSÕES ....................................................................................................... 177 VII. REFERÊNCIAS ..................................................................................................... 180 Resumo e Abstract 2 Resumo e Abstract RESUMO Vieira, Alexsandro Santana. A glândula metapleural e suas secreções em formigas cultivadoras (Attini) e não cultivadoras de fungos (Myrmicini, Blepharidattini e Ectatommini) (Hymenoptera: Formicidae). 2012. 188p. Tese (Doutorado em Ciências Biológicas – Biologia Celular e Molecular). Instituto de Biociências, Universidade Estadual Paulista “Júlio de Mesquita Filho”, UNESP, Rio Claro, SP, 2012. A glândula metapleural é um órgão exclusivo das formigas e sua principal função seria a de produzir secreções antibióticas. O objetivo foi de investigar e comparar as possíveis modificações na morfologia interna da glândula metapleural, assim como a composição química das suas secreções nas formigas cultivadoras (Attini) e não cultivadoras de fungos (Ectatommini, Blepharidattini e Myrmicini). Ficou demonstrado que a glândula metapleural nos grupos Ectatommini, Myrmicini, Blepharidatiini e Attini (basal e derivada), apresentam suas porções: secretora formada por várias células secretoras, e armazenadora (reservatório), conectadas entre si por meio de canalículos (intra e extracitoplasmáticos). As Attini derivadas e basais (cultivadoras de fungos) diferiram das não cultivadoras de fungos, por possuírem as células secretoras com forma oval. Ainda, as células secretoras da glândula metapleural das formigas cortadeiras apresentaram
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