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DISCRIMINATORY PROFILE of Rdna SITES and TREND for ACROCENTRIC CHROMOSOME FORMATION in the GENUS TRACHINOTUS LACÉPÈDE, 1801 (PERCIFORMES, CARANGIDAE)

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DISCRIMINATORY PROFILE of Rdna SITES and TREND for ACROCENTRIC CHROMOSOME FORMATION in the GENUS TRACHINOTUS LACÉPÈDE, 1801 (PERCIFORMES, CARANGIDAE) PARÂMETROS CITOGENÉTICOS DE ESPÉCIES COMERCIAIS DE CARANGIDAE (PERCIFORMES), COM VISTAS A SUA EMPREGABILIDADE NA CONSERVAÇÃO BIOLÓGICA E PISCICULTURA MARINHA UEDSON PEREIRA JACOBINA Tese de doutorado apresentada ao Programa de Pós Graduação – Rede Nordeste de Biotecnologia (RENORBIO), como parte dos requisitos para obtenção do Título de Doutor em Biotecnologia Orientador- Dr. Wagner Franco Molina NATAL-RN 2012 Seção de Informação e Referência Catalogação da Publicação na Fonte. UFRN / Biblioteca Central Zila Mamede Jacobina, Uedson Pereira Parâmetros citogenéticos de espécies comerciais de carangidae (perciformes), com vistas a sua empregabilidade na conservação biológica / Uedson Pereira Jacobina. – Natal, RN, 2012. 150 f. : il. Orientador: Wagner Franco Molina. Tese (Doutorado) – Universidade Federal do Rio Grande do Norte. Rede Nordeste de Biotecnologia. 1. Piscicultura Marinha – Tese. 2. Marcadores Cromossônicos – Tese. 3. Software Lekin – Tese. I. Molina, Wagner Franco. II. Universidade Federal do Rio Grande do Norte. III. Título. RN/UF/BCZM CDU 639.2.04 Dedico este trabalho a meu pai Lídio Jacobina Vieira Santos “Nem tudo que se enfrenta pode ser modificado, mas nada pode ser modificado até que seja enfrentado. A tradição é a personalidade dos imbecis.” Albert Einstein AGRADECIMENTOS Agradeço a Universidade Federal do Rio Grande do Norte e ao programa Rede Nordeste de Biotecnologia (RENORBIO) pela oportunidade de realizar este trabalho. Ao meu orientador Professor Dr. Wagner Franco Molina, pelo apoio e confiança depositada em mim, que foram fundamentais para a realização deste trabalho e oportunidade de trabalhar com grupo, objeto de estudo, “fantástico”, os Carangídeos. Aos professores Dr Luiz Antonio Carlos Bertollo, Marcelo de Bello Cioffi e Marcelo Vicari pela amizade, parceria e dicas na melhoria deste trabalho. Aos Professores Dr. Darlio Teixeira, Carlos Blaha e Paulo Sérgio Marinho pelas críticas e sugestões para a melhoria do trabalho. Aos Professores Dr. Orlando Moreira-Filho, Vitor de Oliveira Lunardi, Carlos Alfredo Galindo Blaha e Liliane de Lima Gurgel pelas críticas essenciais e fundamentais na melhoria desse trabalho. Ao Conselho Nacional de Pesquisa (CNPQ) pela bolsa de pesquisa concebida em 2009. Ao Conselho de Aperfeiçoamento Superior pela bolsa REUNI concebida de 2010 a 2012, fundamental na minha permanência e continuidade deste trabalho em Natal. As Professoras Kattia Scortecci e Adriana Uchôa pelos ensinamentos, dicas e bate papos descontraídos nas aulas de Biologia Celular fundamentais para o meu aprendizado e amadurecimento como um futuro docente. A minha família, os irmãos Wilton, Wendell e José Welton e principalmente minha mãe Beatriz Alencar Jacobina por todo amor e carinho e confiança depositada em mim. Ao grande amigo Pablo Martinez pela convivência, parceria nos trabalhos e claro parceiro das melhores baladas em Natal, e por compartilhar os momentos difíceis. Aos grandes amigos Layse, Calado, Washington, Valdir, Silvio e Valério pelo apoio e por tornarem esta convivência mais descontraída e bem mais feliz em minha estadia de quatro anos em Natal. Aos amigos de Guamaré Layse, Kelly, Carol, Pedro, Calado, Acarilton, Érico, Iraê, Gustavo, Diogo, Felipe, Daniel, Aline, Lucas e Léo pela amizade, apoio, compreensão e por tornar a conviência mais fácil nos dias de labuta em Guamabeach. Ao amigo Garcia Júnior pelo apoio e suporte na identificação das espécies. Aos colegas do laboratório de Genética de Recursos Marinhos Gideão, Rodrigo Pantera, Eurico, Allysson, Amanda, George, Clóvis e Paulo por participarem direta ou indiretamente na realização deste trabalho. Em geral gostaria de agradecer a todas as pessoas que de certa forma contribuíram na realização deste trabalho. Resumo Entre os peixes marinhos, as famílias Carangidae e Rachycentridae se apresentam como grupos de grande importância comercial pela pesca e potencial para piscicultura marinha. Entretanto, bases genéticas que possam alicerçar o futuro cultivo destas espécies, sobretudo seus aspectos citogenéticos são incipientes. Os padrões cromossômicos têm fornecido dados básicos para a prospecção de processos biotecnológicos de manipulação cromossômica voltados ao melhoramento genético, como a indução a poliploidia, ginogênese e androgênese, assim como obtenção de estoques monossexo e hibridizações interespecíficas. Neste trabalho é apresentado um amplo levantamento citogenético em 10 espécies, sendo sete da família Carangidae e de Rachycentron canadum, espécie monotípica da família Rachycentridae. A caracterização citogenética clássica e mapeamento in situ de sequências multigênicas foram empregadas. Adicionalmente em espécies do gênero Selene e em morfótipos de Caranx lugubris foram realizadas comparações através de morfometria geométrica. Em geral, as espécies exibiram um marcante conservadorismo cromossômico numérico (2n=48). Apesar de apresentar, em grande parte, fórmulas cariotípicas diferenciadas, conservam diversas características cromossômicas típicas da Ordem Perciformes, como elevado número de elementos monobraquiais, sítios Ag-RONs/ DNAr 18S simples e heterocromatina reduzida, preferencialmente centromérica. Os principais mecanismos envolvidos na diversificação cariotípica são as inversões pericêntricas, com ação secundária de fusões cêntricas. Além do mapeamento físico e detalhamento cromossômico para as espécies são apresentados e discutidos padrões de variabilidade intra e diversificação interespecíficas, com identificação de marcadores citotaxonômicos. Este conjunto dos dados propicia um melhor conhecimento dos padrões carioevolutivos destes grupos e condições para o desenvolvimento de protocolos biotecnológicos baseados na manipulação cromossômica para estas espécies Atlânticas. Abstract Worldwide, families Carangidae and Rachycentridae represent one of the groups most important commercial fish, used for food, and great potential for marine aquaculture. However, the genetic bases that can underpin the future cultivation of these species, cytogenetic between these aspects are very weak. The chromosomal patterns have provided basic data for the exploration of biotechnological processes aimed at handling chromosomal genetic improvement, such as induction of polyploidy, androgenesis and ginogenesis, as well as obtaining monosex stocks and interspecific hybridizations. This paper presents a comprehensive cytogenetic survey in 10 species, seven of the family Carangidae and the monotypic family Rachycentridae. Classical cytogenetic analysis and in situ mapping of multigene sequences were employed, and additionally for the genus Selene and morphotypes of Caranx lugubris, comparisons were made using geometric morphometrics. In general, conservative species exhibit a marked chromosome number (2n=48). Although present in large part, different karyotypic form, retain many characteristics typical of chromosomal Order Perciformes, the high number of elements monobrachyal, Ag-NORs/18S rDNA sites and heterochromatin simply reduced, preferably centromeric. The main mechanisms involved in karyotypic diversification are the pericentric inversions, with secondary action of centric fusions. In addition to physical mapping and chromosome detail for the species are presented and discussed patterns of intra-and interspecific diversity, cytotaxonomic markers. This data set provides a better understanding of these patterns caryoevolutyonary groups and conditions for the development of protocols based on Biotechnology for chromosomal manipulation Atlantic these species. Lista de Figuras e Tabelas Introdução Figura-1. Relações filogenéticas baseadas em dados morfológicos (Gushiken, 1988) para Carangoidei (a) e de tribos da família Carangidae (adaptado de Reed et al., 2002) a partir de sequências Cit B (b). Pag-18 Tabela-1. Dados citogenéticos da família Carangidae. Pag-19 Capítulo 2 Figura 1. Cariótipos de Rachycentron canadum. Coloração convencional (a) destacando os sítios Ag- NORs no par cromossômico no. 2; (b) bandamento C, destacando as RONs como regiões + - heterocromáticas CMA3 /DAPI ; (c) bandas de replicação, evidenciando os sítios de rDNA 18S (par 2) e 5S (pares 3 e 13) como de replicação inicial; (d) double-FISH com sondas de DNAr 18S (rosa) e 5S (verde), evidenciando a localização dos sítios de DNAr 18S no par no. 2 e dos sítios de DNAr 5S nos pares 3 e 13; (e) FISH com seqüências (TTAGGG)n evidenciando a localização de sítios teloméricos nos cromossomos (laranja). Barra = 5 µm. Pag-69 Figura 2. Idiograma representativo do complemento cromossômico de Rachycentron canadum, exibindo o mapeamento citogenético das sequências ribossomais, Ag-RONs, regiões heterocromáticas e bandas de replicação dos cromossomos. Pag-70 Capítulo 3 Figura 1. Cariótipos das espécies Trachinotus goodei (a, d, g), Trachinotus carolinus (b, e, h) e Trachinotus falcatus (c, f, i). Coloração convencional (a, b, c) destacando os pares portadores de sítios Ag-RONs; bandamento C (d, e, f); em destaque os pares organizadores nucleolares sob + - coloração por CMA3 /DAPI . Dupla coloração em FISH com sondas de 18S DNAr (verde) e 5S DNAr (vermelho) (f, g, h). Barra = 5 µm. Pag-78 Figura 2. Árvore filogenética de parcimônia em algumas espécies da tribo Trachinotini (a), adaptado de Reed et al. (2002). As relações moleculares confrontam com uma fórmula cromossômica das espécies de Trachinotus analisadas. Abaixo, uma representação esquematica de uma nova possibilidade de “acrocentrização” (b) e condição derivada de sítios de DNAr múltiplos exclusivo em T. goodei (c). Em destque
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