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Prospecção De Marcadores Citogenéticos Em Grandes Peixes Pelágicos Marinhos

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Prospecção De Marcadores Citogenéticos Em Grandes Peixes Pelágicos Marinhos UNIVERSIDADE FEDERAL DO RIO GRANDE DO NORTE CENTRO DE BIOCIÊNCIAS Programa de Pós-Graduação em Biotecnologia – RENORBIO Prospecção de marcadores citogenéticos em grandes peixes pelágicos marinhos RODRIGO XAVIER SOARES Natal-RN 2017 UNIVERSIDADE FEDERAL DO RIO GRANDE DO NORTE CENTRO DE BIOCIÊNCIAS Programa de Pós-Graduação em Biotecnologia - RENORBIO Prospecção de marcadores citogenéticos em grandes peixes pelágicos marinhos Rodrigo Xavier Soares Tese apresentada ao Programa de Pós- Graduação em Biotecnologia da Universidade Federal do Rio Grande do Norte, como parte dos requisitos para obtenção do título de Doutor em Biotecnologia. Orientador: Dr. Wagner Franco Molina Natal-RN 2017 Universidade Federal do Rio Grande do Norte - UFRN Sistema de Bibliotecas - SISBI Catalogação de Publicação na Fonte. UFRN - Biblioteca Setorial Prof. Leopoldo Nelson – Centro de Biociências – CB Soares, Rodrigo Xavier. Prospecção de marcadores citogenéticos em grandes peixes pelágicos marinhos / Rodrigo Xavier Soares. - Natal, 2017. 114 f.: il. Tese (Doutorado) - Universidade Federal do Rio Grande do Norte. Centro de Biociências. Programa de Pós-graduação em Biotecnologia. Orientador: Prof. Dr. Wagner Franco Molina. 1. Evolução cromossômica - Tese. 2. Cromossomos sexuais - Tese. 3. Citogenética de peixes - Tese. I. Molina, Wagner Franco. II. Universidade Federal do Rio Grande do Norte. III. Título. RN/UF/BSECB CDU 575 Prospecção de marcadores citogenéticos em grandes peixes pelágicos marinhos Rodrigo Xavier Soares Tese apresentada ao Programa de Pós- Graduação em Biotecnologia da Universidade Federal do Rio Grande do Norte, como parte dos requisitos para obtenção do título de Doutor em Biotecnologia. Comisão examinadora Dr. Allyson Santos de Souza Examinador Externo à Instituição - UFRN Dr. Gideão Wagner Werneck Felix da Costa Examinador Externo à Instituição - UFRN Dr. Paulo Augusto de Lima Filho Examinador Externo à Instituição - IFRN Dr. Paulo Sergio Marinho Lucio Examinador Externo ao Programa - UFRN Dr. Wagner Franco Molina Examinador Interno - UFRN Dedico esta tese aos meus pais, a minha família e minha namorada pelo apoio e paciência a mim prestados. “Talvez fosse um pecado ter matado o peixe. Suponho que sim, embora a carne fosse para me conservar a vida e para alimentar muita gente. Mas então tudo é pecado. Não pense no pecado, meu velho. [...] “Mas você não matou o peixe apenas para conservar-se vivo e o vender para alimento”, pensou ele. “Matou-o por orgulho e porque é um pescador. Amava o peixe quando estava vivo, afinal ainda o ama morto. Se o ama, com certeza que não foi pecado matá-lo. Ou será ainda pior?” Ernest Hemingway “O Velho e o Mar” (trad. Fernando de C. Ferro) AGRADECIMENTOS Venho agradecer a todos que fizeram parte da formação pessoal e acadêmica por mim adquirida desde o início da vida até chegar a realização desta tese de doutorado. Aos meus pais, Josenaide e Francisco Canindé, pela educação de berço e pelos princípios morais a mim ensinados e por todo apoio prestado sempre. Todo o agradecimento a eles seria pouco. A minha namorada, Amanda, companheira de todas as aventuras, que me acompanhou e me ajudou como colega e amiga em todos os momentos desse doutorado, participando ativamente de minhas pesquisas. A minha irmã Renata e todos os meus parentes que torceram pelo meu sucesso em cada etapa da minha formação. As famílias Xavier, Soares, Torres, Borges, Gomes e “MAQUE”. A Wagner Molina, que desde a iniciação científica me orienta com paciência e eficiência contribuindo para formação do meu conhecimento científico. A CAPES e CNPQ pelo apoio material e bolsa de estudo fornecido para o desenvolvimento desta tese, Secirm - Marinha do Brasil pelo apoio logístico, ICMBio e IBAMA pelas licenças concedidas e pelo apoio logístico. Assim como a disponibilização de exemplares cultiváveis pelo Daniel Benetti, RSMAS Universidade de Miami. Aos membros e suplentes da banca de qualificação e defesa que aceitaram o convite e contribuíram com suas correções para o melhoramento desta tese. Aos colegas do LGRM que contribuiram ativamente ou passivamente para a conclusão deste trabalho. Dentre eles: Gideão, Paulo, Clóvis, Inailson, Allyson, Karlla, Rafael, Cida, Juliana, Washington, Jeane, Josy, Juliane, Calado, Eurico, Cristiane, Divana, Guinga, Roberta, Wanessa, Davi, Cadu e todos aqueles que não estão citados aqui mas que esteve presente no laboratório algum momento. Aos amigos que torceram por esta realização: Maria Vitória, Janaína, Wania e família, Geomar Maia, Cézar, Rômulo, Equipe China e todos os funcionários do CB. A todos aqueles que participaram das expedições e ajudaram nas coletas, pescarias e processamentos assim como a Transmar (Mestres Jonas e Aristides e os demais tripulantes) e todos da Lancha Isabel. SUMÁRIO Lista de Figuras e Tabelas IX Lista de Abreviaturas XII Resumo XIII Abstract XIV INTRODUÇÃO 15 OBJETIVOS 28 MATERIAL E MÉTODOS 29 REFERÊNCIAS 38 RESULTADOS E DISCUSSÃO Capítulo I Chromosomal evolution in large pelagic oceanic apex predators, the 48 barracudas (Sphyraenidae, Percomorpha) Capítulo II Padrões carioevolutivos em quatro espécies pelágicas de Carangidae 63 (Carangiformes): o papel do fluxo gênico Capítulo III Sistema de cromossomos sexuais múltiplos na espécie circuntropical 82 Coryphaena hippurus (Linnaeus, 1758) Capítulo IV Caracterização citogenômica das espécies pelágicas Megalops atlanticus 99 (Tarpão) e Istiophorus albicans (Agulhão-de-vela), alvos da pesca esportiva CONCLUSÕES 113 LISTA DE FIGURAS E TABELAS Introdução Figura 1. Dados de produção mundial de pesca e aquacultura em milhões 18 de toneladas, com projeção para o ano de 2025 (FAO, 2016). Figura 2. Produção pesqueira mundial de Coryphaena hippurus em 24 tonelada (k = x1000 toneladas) por ano (FAO, 2017b). Figura 3. Exemplares de Sphyraena barracuda (a), Sphyraena guachancho 30 (b) e Sphyraena picudilla (c) (Sphyraenidae, Istiophoriformes). Barra = 5cm. Figura 4. Exemplares de Carangidae (Carangiformes): Elagatis bipinnulata 31 (a), Seriola rivoliana (b), Gnathanodon speciosus (c) e Trachinotus carolinus (d). Barra = 5cm. Figura 5. Exemplares de Coryphaena hippurus (Coryphaenidae, 32 Carangiformes). Macho (a) e fêmea (b). Barra = 5cm. Figura 6. Exemplares de Istiophorus albicans (Istiophoridae) (Fonte: 32 Ilmaredamare.com) (a) e Megalops atlanticus (Megalopidae) (b). Barra = 5cm. Figura 7. Mapa com pontos de coletas e espécies coletadas: RN = Litoral do 33 Rio Grande do Norte, FN = Arquipélago de Fernando de Noronha, ASPSP = Arquipélago de São Pdero e São Paulo, FL = Flórida. Ch = Coryphaena hippurus, Eb = Elagatis bipinnulata, Sr = Seriola rivoliana, Sp = Sphyraena picudilla, Sb = Sphyraena barracuda, Sg = Sphyraena guachancho, Ia = Istiophorus albicans, Ma = Megalops atlanticus, Tc = Trachinotus carolinus. Capítulo I Figura 8. Karyotypes of the species Sphyraena picudilla, Sphyraena 53 barracuda and Sphyraena guachancho. Staining with Giemsa (Ag- NOR and MM+/DAPI- sites are highlighted) (a) C-banding (b); double-FISH with 18S rDNA (red) and 5S rDNA (green) probes (c); and distribution of the (TTAGGG)n telomeric sequences (d). Scale bar = 5 μm. Figura 9. Phylogenetic perspective from karyotype patterns and 18S/5S 55 rDNA arrangements in species of Sphyraena (adapted from Milana et al. 2014 and Santini et al. 2014). Capítulo II Figura 10. Cariótipos com coloração Giemsa (sítios Ag-RONs e MM+/DAPI- 68 nas caixas) das espécies Elagatis bipinnulata (a, c, e) e Seriola rivoliana (b, d, f). Bandamento C (c, d) e double-FISH com sondas DNAr 18S (vermelho) e DNAr 5S (verde) (e, f). Barra de escala = 5 μm. Figura 11. Cariótipos com coloração Giemsa (sítios Ag-RONs e MM+/DAPI- 69 nas caixas) das espécies Gnathanodon speciosus (a, c, e) e Trachinotus carolinus (b, d, f). Bandamento C (c, d) e double-FISH com sondas DNAr 18S (vermelho) e DNAr 5S (verde) (e, f). Barra de escala = 5 μm. Figura 12. Diversidade cromossômica entre espécies de Naucratini 74 (Carangidae), em relação à Gnathanodon speciosus (Carangini) e Trachinotus carolinus (Trachinotini), sob perspectiva filogenética. A barra indica datação dos processos evolutivos em milhões de anos (adaptado de Santini & Carnevale, 2015). Capítulo III Figura 13. Cariótipos, de fêmeas (a) e machos (b) de Coryphaena hippurus 87 com cromossomos corados com Giemsa e bandamento C. Em destaque nos quadros, cromossomos portadores de sítios Ag- RONs e correspondente região MM+/DAPI. Hibridização in situ de duas cores (c) com sondas DNAr 18S (vermelho) e DNAr 5S (verde) em fêmea e macho respectivamente. Cromossomos X1X2Y estão destacados no cariótipo dos machos. Barra=5 µm. Figura 14. Hibridização in situ evidenciando a distribuição de sequências 88 teloméricas (TTAGGG)n (a, b) e de elementos transponíveis Tol2 (c, d), respectivamente em fêmeas (a, c) e machos (b, d) de C. hippurus. A seta indica o cromossomo Y. Barra = 5 µm. Figura 15. Ideograma dos padrões cromossômicos de machos e fêmeas de 89 C. hippurus, evidenciando as regiões 18S rDNA/Ag-RONs/GC+/AT-, DNAr 5S, sequências teloméricas (TTAGGG)n e distribuição de elementos transponíveis Tol2. Figura 16. Relações filogenéticas entre Coryphaenidae, Rachycentridae e 91 Echeneidae e origem estimada do sistema de cromossomos sexuais múltiplos X1X1X2X2/X1X2Y em Coryphaena, em milhões de anos (adaptado de Santini & Carnevale, 2015). Capítulo IV Figura 17. Mapa de distribuição geográfica de Istiophorus albicans (a) e 101 Megalops atlanticus (b) (Adaptado de Froese & Pauly, 2017). Figura 18. Cariótipos de Megalops atlanticus (a) e Istiophorus albicans (b). 104
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