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2013 Cesardanielpetroli.Pdf UNIVERSIDADE DE BRASÍLIA Instituto de Ciências Biológicas Departamento de Biologia Celular TESE DE DOUTORADO CARACTERIZAÇÃO GENÔMICA DE MARCADORES DArT COM BASE EM MAPEAMENTO GENÉTICO E FÍSICO E DETECÇÃO DE QTLs EM Eucalyptus CÉSAR DANIEL PETROLI BRASILIA – DF Abril, 2013 UNIVERSIDADE DE BRASÍLIA Instituto de Ciências Biológicas Departamento de Biologia Celular CARACTERIZAÇÃO GENÔMICA DE MARCADORES DArT COM BASE EM MAPEAMENTO GENÉTICO E FÍSICO E DETECÇÃO DE QTLs EM Eucalyptus OriEntador: Dr. Dario Grattapaglia Tese apresentada ao Departamento de Biologia Celular do Instituto de Biologia, da Universidade de Brasília, como requisito parcial para obtenção do grau de Doutor em Ciências Biológicas, Área de Concentração: Biologia Molecular BRASILIA – DF Abril, 2013 ii TERMO DE APROVAÇÃO Tese apresentada ao Departamento de Biologia Celular da Universidade de Brasília, como requisito parcial para obtenção de grau de Doutor em Ciências Biológicas, área de concentração Biologia Molecular. Tese defendida e aprovada em 26/04/2013 por: Dr. Dario Grattapaglia - Orientador Embrapa Recursos Genéticos e Biotecnologia Dr. Marcio Elias Ferreira Embrapa Recursos Genéticos e Biotecnologia Dr. Georgios Joannis Pappas Júnior Universidade de Brasília Dr. Márcio de Carvalho Moretzsohn Embrapa Recursos Genéticos e Biotecnologia Dr. Alan Carvalho Andrade Embrapa Recursos Genéticos e Biotecnologia iii A mEus grandEs amorEs Caro E nossa princesa IsabElla DEDICO iv AGRADECIMENTOS A realização dessa tese somente foi possível pelo apoio recebido de diversas pessoas e instituições, mas gostaria de agradecer de maneira especial: À Universidade de Brasília (UnB), e especialmente ao programa de pós-graduação em Biologia Molecular, representados por todos os professores e funcionários, pela oportunidade de realização deste curso. À Coordenação de Aperfeiçoamento de Pessoal de Nivel superior (CAPES) pelo apoio financeiro que tornou possível a realização desse trabalho. À EMBRAPA Recursos Genéticos e Biotecnologia, pela infra-estrutura de trabalho. A todas as empresas florestais e Instituições que facilitaram as amostras e os recursos necessários para a realização deste trabalho. Devo muito à orientação do Dr. Dario Grattapaglia, com quem estabeleci uma relação profissional e pessoal, a qual desejo seja duradoura após a finalização deste trabalho. É em pessoas como ele que eu me espelho e busco inspiração para o meu próprio desenvolvimento profissional e pessoal. Ao Dr. Andrzej Kilian, director da empresa Diversity Arrays Technology, por ter acreditado no projeto e aberto as portas do DArT para recebernos com muito carinho em Canberra, Australia. Pelos grandes conhecimentos transmitidos que são os pilares para o meu futuro profissional. Aos membros da banca de avaliação desta tese, o Dr. Marcio Elias Ferreira, Dr. Georgios J. Pappas Júnior, Dr. Marcio de Carvalho Moretzsohn e o Dr. Alan Carvalho Andrade, pelos comentários, sugestões e questionamentos que foram essenciais para a melhoria e o estabelecimento do formato final. v Aos pesquisadores e técnicos do Laboratorio de Genética Vegetal da Embrapa Recursos Genéticos e Biotecnología, Marcão, Marcio, Vânia, Glaucia, Zilneide, Lorena, Peter e muito especialmente a minha grande amiga e conselheira Marília. Aos amigos que tive o prazer de conhecer no Laboratorio de Genética Vegetal, Bruna (e Daniel), Thaisa, Marília (Georgios e meu príncipe Nicolas), Dione (Andrê e Ian), Ediene e Rodrigo, Tati (Bruno e Arthur), Natália, Mariana, Marco (Bia e Teo), Tulio (e Cecilia) e Pedro pela amizade de tantos anos. A muitos de vocês considero irmãos do coração que a vida me presenteu e tenho certeza que nem a distância ou o tempo vai destruir essa amizade tão linda e verdadeira. Aos meus amigos do laboratório DarT na Austrália, Colleen, Michael, Kasia, Grzegorz, Jason, Ling, Eric, Damian, Puthick, Vanessa, Cina, Cleare, Frank, Gosia, Hang e Adriane, por todo o apoio e carinho durante minha estadia em Canberra, onde passei dois anos maravilhosos da minha vida, cehio de experiências inesquecíveis. Aos meus irmãos brasileiros Eduardo, Genny, Arnon, Vinicius, Leo, Gabriel, William e Reinaldo, você sempre estarão presentes. A meus pais, Elena e Cacho, os principais responsáveis por minha educação e formação como pessoa. Sou eternamente agradecida pelos ensinamentos, apoio incondicional e incentivo para meu crescimento pessoal e profissional. Tudo o que eu sou eu devo a vocês. Ao meu irmão David e sua esposa Sonia pelo apoio, carinho e principalmente por ter me presenteado com duas crianças maravilhosas: Sofia e Tisiano, amos muito vocês! Ao meu irmão Gastón, pelo carinho e alegria de sempre. Apesar da juventude você é um exemplo para mim. vi À família que me acolheu como mais um filho: Juan, Norma, Adrian, Sole, Vale, Flor e Vicky, tios e primos, pelo carinho e apoio recebido sempre. À Carol, meu grande amor e companheira de vida, pelo seu apoio, amizade, compreensão e amor incondicional. Este trabalho é compartilhado 100%. Obrigado por ser a melhor esposa do mundo e por ter cumprido meu grande sonho de formar juntos, uma família maravilhosa. E finalmente a pessoa que conseguiu mudar completamente o sentido da minha vida, nossa princessa Isabella. Nunca imaginei que meu coração tinha tanto amor para dar. Cada dia com um simples sorriso você me faz o papai mais feliz do mundo. Te amo filha! Também para a pessoa que está chegando, você iluminara nossas vidas tanto como a sua irmazihna. vii ÍNDICE Resumo 1 Abstract 2 1. INTRODUÇÃO 4 2. REVISÃO BIBLIOGRÁFICA 5 2.1. O gênero Eucalyptus 5 2.2. Atualidade e impacto econômico do Eucalyptus no Brasil 6 2.3. Mapeamento genético 8 2.4. Analise de QTLs (Quantitative Trait Loci) 10 2.5. Melhoramento genético de Eucalyptus 11 2.6. Desenvolvimento de marcadores moleculares em Eucalyptus 13 2.7. Estudos de genética de populações e evolução em Eucalyptus 15 2.8. Desenvolvimento de mapas genéticos em Eucalyptus 15 2.9. Identificação de QTLs em Eucalyptus 17 2.10. Marcadores DArT (Diversity Arrays Technology) 20 3. CAPÍTULO 1: CARACTERIZAÇÃO GENÔMICA DE MARCADORES DArT 24 BASEADA NA ANÁLISE DE MAPEAMENTO POR LIGAÇÃO E FÍSICO NO GENOMA DE Eucalyptus 3.1. INTRODUÇÃO 24 3.2. OBJETIVOS 28 3.3. MATERIAL E MÉTODOS 29 3.3.1. Material vegetal 29 3.3.2. Genotipagem de microssatélites 29 3.3.3. Genotipagem de marcadores DArT (Diversity Arrays Technology) 29 3.3.4. Construção do mapa genético 31 3.3.5. Análise comparativa entre o mapa de ligação e a montagem de sequência 32 do genoma. 3.3.6. Análise comparativa entre o mapa de ligação e a montagem da sequência 32 do genoma 3.4. RESULTADOS 34 viii 3.4.1. Genotipagem dos marcadores DarT 34 3.4.2. Mapeamento de ligação 36 3.4.3. Distâncias de recombinação e física no genoma de Eucalyptus 43 3.4.4. Análises de redundância das sequências das sondas DArT 47 3.4.5. Alinhamento das sondas DArT no genoma de Eucalyptus 49 3.4.6. Cobertura do espaço gênico de Eucalyptus pelos marcadores DArT 50 3.5. DISCUSSÃO 54 3.5.1. Eficiência da genotipagem de marcadores DArT para análise genética em 55 Eucalyptus 3.5.2. Estimativas de redundância das sondas DarT 57 3.5.3. O mapa genético framework permite estimativas mais confiáveis da 58 relação kpb/cM no genoma de Eucalyptus 3.5.4. O alinhamento do mapa genético à sequência física sugere uma 61 característica pan-genômica do microarranjo DArT e a completude da montagem do genoma de Eucalyptus 3.5.5. O microarranjo DArT oferece uma cobertura uniforme do genoma e 62 amostra preferencialmente regiões ricas em genes 3.6. CONCLUSÃO 63 4. CAPÍTULO 2: MAPEAMENTO DE QTLs PARA CARACTERÍSTICAS DE 65 IMPORTÂNCIA ECONÔMICA E ANÁLISE DO CONTEUDO GÊNICO NOS INTERVALOS GENÔMICOS CORRESPONDENTES 4.1. INTRODUÇÃO 65 4.2. OBJETIVOS 68 4.3. MATERIAL E MÉTODOS 69 4.3.1. Material vegetal 69 4.3.2. Construção dos mapas e detecção de QTLs 69 4.4. RESULTADOS 70 4.4.1. Correlação entre as características fenotípicas 70 4.4.2. Mapas genéticos e detecção de QTLs (Quantitative Trait Loci) 71 4.4.3. Detecção de QTLs para densidade básica da madeira 75 4.4.4. Detecção de QTLs para crescimento em altura 78 ix 4.4.5. Detecção de QTLs para diâmetro à altura do peito 81 4.4.6. Detecção de QTLs para rendimento depurado de celulose 83 4.4.7. Detecção de QTLs para teor de lignina total da madeira 86 4.4.8. Detecção de QTLs para a relação Siringil/Guaiacil 91 4.4.9. Detecção de QTLs para densidade da madeira medida pela profundidade 94 de penetração do Pilodyn. 4.4.10. Co-localização de QTLs para diferentes características 97 4.4.11. Conteúdo de genes nos intervalos de QTLs 103 4.5. DISCUSSÃO 105 4.5.1. Análise comparativa com QTLs detectados em outros estudos de 105 Eucalyptus 4.5.2. Estimativas das proporções da variação explicadas pelos QTLs 109 4.5.3. Co-localização entre QTLs mapeados para diferentes características 111 4.5.4. De QTLs para genes e seu uso no melhoramento 112 4.6. CONCLUSÃO 114 5. BIBLIOGRAFIA 115 LISTA DE FIGURAS Figura 1: Procedimento de desenvolvimento de marcadores DArT. 23 Figura 2. Interpretação de dados calculado pelo programa DArTSoft mostrando a 24 diferença de intensidade entre genótipos. Figura 3. Distribuição do número e porcentagens de marcadores DArT que 36 passaram o limiar de filtragem adotado para reprodutibilidade (≥95%), qualidade (Q≥65%) e call rate (≥75%). Figura 4. Mapa de ligação framework DArT/microssatélites para Eucalyptus. 38 Figura 5. Alinhamento do mapa full (barras amarelas) com o mapa framework 41 (Fmwk) (barras verdes) para os 11 pseudo-cromossomos do Eucalyptus mostrando as conexões entre os mesmos locos em ambos os mapas. Figura 6. Distribuição de frequência das distâncias de recombinação de Kosambi 42 entre marcadores consecutivos ao longo das duas versões do mapa de x ligação. Figura 7. Alinhamento do mapa framework com o genoma de referência de 46 Eucalyptus grandis.
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