Mollusca: Cephalopoda) E Reavaliação Da Reação Espermatofórica Dos Cefalópodes

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Mollusca: Cephalopoda) E Reavaliação Da Reação Espermatofórica Dos Cefalópodes José Eduardo Amoroso Rodriguez Marian Estrutura e funcionamento dos espermatóforos de Doryteuthis plei (Mollusca: Cephalopoda) e reavaliação da reação espermatofórica dos cefalópodes Spermatophore structure and functioning in Doryteuthis plei (Mollusca: Cephalopoda), and a reappraisal of the cephalopod spermatophoric reaction São Paulo 2010 José Eduardo Amoroso Rodriguez Marian Estrutura e funcionamento dos espermatóforos de Doryteuthis plei (Mollusca: Cephalopoda) e reavaliação da reação espermatofórica dos cefalópodes Spermatophore structure and functioning in Doryteuthis plei (Mollusca: Cephalopoda), and a reappraisal of the cephalopod spermatoforic reaction Tese apresentada ao Instituto de Biociências da Universidade de São Paulo, para a obtenção de Título de Doutor em Ciências, na Área de Zoologia. Orientadora: Profa. Dra. Sônia Godoy Bueno Carvalho Lopes São Paulo 2010 Marian, José Eduardo Amoroso Rodriguez M 333e Estrutura e funcionamento dos espermatóforos de Doryteuthis plei (Mollusca: Cephalopoda) e reavaliação da reação espermatofórica dos cefalópodes / José Eduardo Amoroso Rodriguez Marian. -- São Paulo : J. E. A. R. M., 2010. 238p. : il. Tese (Doutorado) - Instituto de Biociências da Universidade de São Paulo. Departamento de Zoologia, 2010. 1. Cephalopoda – Morfologia 2. Loliginidae 3. Cephalopoda – Reprodução 4. Espermatóforo 5. Reação espermatofórica 6. Mollusca I. Universidade de São Paulo. Instituto de Biociências. Departamento de Zoologia II. Título LC QL 430.2 Comissão Julgadora: Prof(a). Dr(a). Prof(a). Dr(a). Prof(a). Dr(a). Prof(a). Dr(a). Profa. Dra. Sônia G. B. C. Lopes Orientadora Ao meu querido sobrinho, Guilherme. "I believe that in approaching our subject with the sensibilities of statisticians and dissectionists, we distance ourselves increasingly from the marvelous and spell-binding planet of imagination whose gravity drew us to our studies in the first place. This is not to say that we should cease to establish facts and to verify our information, but merely to suggest that unless those facts can be imbued with the flash of poetic insight then they remain dull gems; semi-precious stones scarcely worth the collecting." "Until we transform our mere sightings into genuine visions; until our ear is mature enough to order a symphony from the shrill pandemonium of the aviary; until then we may have a hobby, but we shall not have a passion." "A scientific understanding of the beautifully synchronized and articulated motion of an owl’s individual feathers during flight does not impede a poetic appreciation of the same phenomenon. Rather, the two enhance each other, a more lyrical eye lending the cold data a romance from which it has long been divorced." Alan Moore, Blood from the Shoulder of Pallas AGRADECIMENTOS Gostaria de expressar meus sinceros agradecimentos a todos que, de alguma forma, contribuíram para a concretização desta Tese. Em primeiro lugar, ao Prof. Dr. Osmar Domaneschi (in memoriam), pela dedicação e seriedade com que orientou este trabalho, desde o projeto inicial até pouco antes de seu falecimento, em março de 2008. Conheci seu entusiasmo pela busca e transmissão do conhecimento durante as aulas teóricas e práticas de Zoologia dos Invertebrados, o que me motivou a iniciar meus estudos acadêmicos em seu laboratório. A confiança depositada em mim para iniciar uma nova frente de pesquisa, com cefalópodes, foi uma das maiores motivações que tive ao longo desses anos de academia. A presente Tese estaria certamente muito superior se ele tivesse tido a oportunidade de acompanhar completamente o desenvolvimento do trabalho. Morfologista funcional habilidoso e experiente, Prof. Osmar teria contribuído enormemente nas fases finais do trabalho. Acredito que seus ensinamentos em vida, entretanto, foram suficientes para que eu pudesse completar mais essa etapa a contento. Sua conduta moral impecável, seu intenso espírito de luta e incansável esforço foram igualmente importantes para o engrandecimento desta Tese, mas principalmente para minha formação acadêmica e pessoal. À Profa. Dra. Sônia Godoy Bueno Carvalho Lopes, pela coragem de assumir a orientação de um projeto de Doutorado em fase avançada. Agradeço a confiança, o respeito e a liberdade a mim concedida, fundamentais para concluir meu trabalho. Sua generosidade nunca será esquecida, principalmente por ter proporcionado distintas oportunidades que enriqueceram minha formação profissional e pessoal. À Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP – processo 04/13212-7), pela concessão de bolsa de Doutorado Direto e de auxílio financeiro (Reserva Técnica), indispensável para custeio das despesas relacionadas à pesquisa. À Pró-Reitoria de Pós-Graduação da Universidade de São Paulo, pelo auxílio financeiro concedido para participação no "Cephalopod International Advisory Council Symposium (CIAC 2006)" e no Workshop "Cephalopods as Experimental Animals", realizados na Tasmânia, Austrália. À CAPES PROAP-2006, pelo auxílio financeiro concedido no início do projeto, indispensável para realização das atividades relacionadas à pesquisa. A American Malacological Society (AMS Student Research Grant, 2005) e Houston Conchology Society (Constance E. Boone Grant to Malacology, 2005), pela concessão de auxílios financeiros. À Sociedade Brasileira de Malacologia (SBMa), pelo constante estímulo à pesquisa malacológica no Brasil. Ao "Programa de Pós-Graduação em Ciências" do IBUSP, pelo excelente curso de pós-graduação. Ao Instituto de Biociências da USP e, em especial, ao Departamento de Zoologia deste Instituto, pela qualidade e excelência da infra-estrutura destinada à pesquisa. Ao Centro de Biologia Marinha da USP (CEBIMar-USP), pela excelente infra- estrutura destinada à pesquisa em biologia marinha. Ao Diretor, docentes, funcionários e alunos do CEBIMar, estendo os agradecimentos pelo agradável e engrandecedor convívio durante minhas estadias neste Centro. Ao Prof. Dr. Álvaro Esteves Migotto (CEBIMar-USP), por ter permitido utilização irrestrita do laboratório de microscopia do CEBIMar. Agradeço a ele e ao M. Sc. Bruno C. Vellutini pelas valiosas orientações durante as filmagens da reação espermatofórica. Aos técnicos do CEBIMar-USP, Sr. Eduardo Honuma, Sr. Joseilto Medeiros de Oliveira, Sr. Joseph Marie Etienne Van Sebroeck e Sr. Elso Alves da Silva, e ao técnico aposentado deste Centro, Sr. Moisés Alexandre da Mota, agradeço o indispensável e incansável auxílio durante as coletas de lulas. Ao Prof. Dr. Cláudio Gonçalves Tiago e à Profa. Dra. Tagea Kristina Simon Björnberg, ambos do CEBIMar-USP, pela solicitude e assistência durante minhas estadias neste Centro. À Profa. Dra. Marta Antoniazzi, por ter permitido a utilização da infra-estrutura do "Laboratório de Biologia Celular do Instituto Butantan" para realização dos procedimentos de microtomia de materiais incluídos em historresina. Estendo os agradecimentos aos docentes, técnicos e alunos deste Laboratório, em especial às biólogas Adriana Giorgi Barsotti e Simone Gonçalves Silva Jared, pela assistência nos procedimentos de histologia. À Profa. Dra. Toshie Kawano (in memoriam), por ter permitido a utilização da infra- estrutura do "Laboratório de Parasitologia do Instituto Butantan", especificamente a utilização do microscópio de varredura laser confocal, e principalmente por ter sido um exemplo de humanidade a todos os que a conheceram. Ao Sr. Alexsander S. de Souza, pela assistência e operação do microscópio de varredura laser confocal do "Laboratório de Parasitologia do Instituto Butantan". Ao Prof. Dr. Alberto Ribeiro, por ter permitido a utilização do microscópio eletrônico de varredura e do micrótomo para historresina do "Laboratório de Biologia Celular do IBUSP". Estendo os agradecimentos ao M. Sc. Waldir Caldeira, pela orientação na utilização deste micrótomo. Ao Sr. Ênio Mattos e Sr. Eduardo Mattos (in memoriam), técnicos do Departamento de Zoologia, pela operação do microscópio eletrônico de varredura e pela solicitude e assistência durante o desenvolvimento desta Tese. Aos seguintes pesquisadores do IB-USP, por terem permitido a utilização da infra- estrutura de seus respectivos Laboratórios, indispensável para o desenvolvimento do trabalho: Prof. Dr. Carlos Eduardo Falavigna da Rocha (Laboratório de Carcinologia), Prof. Dr. Fernando Portella de Luna Marques (Laboratório de Helmintologia Evolutiva) e Prof. Dr. José Carlos de Freitas (Laboratório de Toxicologia e Produtos Naturais). Ao Dr. Richard E. Young (University of Hawaii, E.U.A.), pela leitura crítica dos manuscritos referentes aos Capítulos 3, 4 e 5 desta Tese e por todos os comentários e sugestões que permitiram o engrandecimento deste trabalho. Aos seguintes colegas pesquisadores, pelo auxílio na obtenção de bibliografia: Dr. Frank E. Anderson (Southern Illinois University, E.U.A.), Dr. Maurício Leme da Fonseca (Cenpes/Petrobras, Rio de Janeiro/RJ), M. Sc. Monica Lemos Ammon Fernandez (Fiocruz, Rio de Janeiro/RJ), Dr. Phil Pascoe (Plymouth Marine Laboratory, Inglaterra), M. Sc. Renata Souza de Oliveira (UNICAMP, Campinas/SP) e Dr. Vladimir Laptikhovsky (Government Fisheries Department, Ilhas Falkland). Aos colegas oriundos do Laboratório de Malacologia do IBUSP, entre eles Dra. Eliane Pintor Arruda, Dr. Flávio Dias Passos, Dra. Maria Júlia Estefânia Chelini, Dr. Maurício Leme da Fonseca e Pedro Ribeiro Piffer, pelo companheirismo e constante apoio durante o desenvolvimento desta Tese. Os respectivos Auxílios à Pesquisa e Reservas Técnicas de Bolsa de alguns destes colegas auxiliaram no engrandecimento da
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