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Differentiation and Genetic Variability in Cork Oak Populations (Quercus Suber L.)

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Differentiation and Genetic Variability in Cork Oak Populations (Quercus Suber L.) UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS DEPARTAMENTO DE BIOLOGIA ANIMAL Differentiation and genetic variability in cork oak populations (Quercus suber L.) Joana Seabra Pulido Neves da Costa MESTRADO EM BIOLOGIA HUMANA E AMBIENTE Lisboa 2011 UNIVERSIDADE DE LISBOA FACULDADE DE CIÊNCIAS DEPARTAMENTO DE BIOLOGIA ANIMAL Differentiation and genetic variability in cork oak populations (Quercus suber L.) Joana Seabra Pulido Neves da Costa Dissertação orientada por: Prof. Doutor Octávio Fernando de Sousa Salgueiro Godinho Paulo Doutora Dora Cristina Vicente Batista Lyon de Castro MESTRADO EM BIOLOGIA HUMANA E AMBIENTE Lisboa 2011 Nota prévia A presente tese de mestrado encontra-se escrita na língua Inglesa uma vez que esta é considerada a língua científica universal. Por esta razão, o conhecimento e treino da sua escrita apresentam uma importância considerável para quem tenciona seguir uma carreira em investigação científica em Biologia. Com a escrita da tese em Inglês pretende-se também acelerar o processo de elaboração dos manuscritos e subsequentes publicações científicas. As referências bibliográficas foram elaboradas segundo os parâmetros da revista científica internacional, “Trends in Ecology and Evolution” (www.cell.com/trends/ecology- evolution/authors). Esta é uma das revistas mais relevantes na área em que esta tese foi desenvolvida e possui um sistema de citações cómodo para a leitura de textos de revisão científica. Adicionando o seu elevado factor de impacto na sociedade científica, pareceu apropriada a escolha desta revista como referência para a apresentação da bibliografia. O estudo elaborado nesta tese foi desenvolvido no âmbito do projecto PTDC/AGR- GLP/104966/2008, “Avaliação dos recursos genéticos e genómicos do sobreiro: bases para uma gestão prospectiva”, financiado pela Fundação para a Ciência e Tecnologia (FCT). III Foreword The present master thesis is written in English. This is considered as the universal scientific language and, therefore, is of the upmost importance the practice of its writing and grammar for those who intend to follow a career in Biology and scientific investigation. Also, the writing of the present thesis in the English language allows to accelerate the process of submission of the manuscripts for further publication. The bibliographic references were elaborated following the parameters of the international scientific journal “Trends in Ecology and Evolution” (www.cell.com/trends/ecology- evolution/authors). This is one of the most relevant journals in the area where this thesis was developed, with an elevated impact factor in the scientific society. Also it possesses a confortable citations system for the reading long texts. This study is part of the project PTDC/AGR-GLP/104966/2008, “Avaliação dos recursos genéticos e genómicos do sobreiro: bases para uma gestão prospectiva”, funded by Fundação para a Ciência e Tecnologia (FCT). IV Agradecimentos No terminar desta tese surge a necessidade de agradecer a todos aqueles que de alguma forma a tornaram possível. O primeiro agradecimento é devido aos meus orientadores, Octávio Paulo e Dora Batista. Ao Professor Octávio pelo incentivo e voto de confiança que depositou em mim desde o início. À Dora pela proposta do tema de mestrado e pelo despertar do meu interesse pelas plantas. À Professora Deodália Dias pelas oportunidades que me proporcionou e pelo apoio incondicional quando os problemas fogem ao nosso controlo e não dependem de nós. Agradeço em particular à Professora Helena Almeida do Instituto Superior de Agronomia pelo acesso à Herdade Monta da Fava de onde vieram algumas das populações de sobreiro mais importantes para o desenvolvimento deste trabalho. A todos os que directa ou indirectamente foram importantes para a recolha das amostras. Ao CoBiG2 pelo grupo que se formou e pelos bons momentos. Ao Francisco Pina-Martins e à Vera Nunes por me terem criado nos momentos iniciais da minha vida de laboratório. À Sofia Seabra pela sua calma natural e amizade. Ao Eduardo Marabuto pelo seu bom humor, muito necessário em tempos difíceis. À Sara Ema, por incrível que te possa parecer acho que stressas mais que eu e isso é uma ajuda enorme, assim como estares comigo até ao dia da entrega… mesmo de moletas. Ao Diogo Silva pelo partilhar de alguns momentos difíceis com os orientadores a afins. À Catarina Dourado, Ana Sofia, Patrícia Brás, Renata Martins, Inês Modesto e Bruno Vieira que me foram ajudando com as usuais dificuldades de uma tese. Aos restantes membros do CoBiG2, assim como a antigos membros e às mais recentes aquisições, muito obrigada! À Rita Oliveira e Raquel Vaz, não fazem parte do CoBiG2, mas fazem parte da família e merecem o devido reconhecimento e agradecimento pelo que “aturaram” da minha parte. Um agradecimento especial a quatro pessoas que devem ter sofrido muito comigo. Seriam precisas páginas de agradecimentos, mas como não o posso fazer fica a intenção. À Catarina Dourado um agradecimento em particular. Foi um longo caminho e fica o agradecimento pelo carinho, apoio e amizade. Ao Eduardo Marabuto pelos valiosos comentários, acima de tudo na Introdução. Tens razão em muitas coisas mas há que fazer compromissos. À Sofia Seabra pelas importantíssimas correcções, seria muito mais difícil sem ti. Ao Bruno Vieira pelas V horas infinitas que me ouviu queixar da vida em geral, da tese em particular. Sempre com muito amor e carinho! Obrigada aos quatro! À Diana Martins. Não estás sempre comigo mas estás sempre a pensar em mim e tens timings impecáveis para quando preciso mais de ti. E ao Pai, Mãe e Avós. Apesar de estarem no fim desta lista foram provavelmente as pessoas que mais contribuíram para que esta tese pudesse ser concluída. Claramente não estaria cá sem o apoio precioso da minha família. VI Resumo O ano 2011 foi designado como “O Ano Internacional das Florestas” pela Assembleia Geral das Nações Unidas, na tentativa de despertar o interesse público e promover a sustentabilidade da gestão e conservação florestal para o benefício das gerações futuras. Estimativas da FAO (Food and Agriculture Organization) para o ano de 2010 demonstraram que 31% da superfície terrestre ainda está coberta por florestas e que as árvores correspondem a 90% da biomassa terrestre, compreendendo um total de 60.000 a 100.000 taxa. Contudo, certas alterações induzidas pelo Homem, principalmente a desflorestação e as alterações climáticas elevaram o número de espécies ameaçadas de extinção para 10%. Nos últimos tempos as espécies florestais têm sido bastante usadas em estudos de genética populacional e evolutiva, assim como em estudos genómicos. As principais razões são as características particulares que estes modelos não-clássicos apresentam, visto resultarem de milhões de anos de divergência e diversificação, e assim apresentarem impressionantes níveis de diversidade morfológica, divergência evolutiva e diversidade ecológica. Apesar de o impacto que as alterações globais vão ter sobre estas espécies depender grandemente da sua capacidade de reacção e da dos seus ecossistemas, os estudos genéticos permitem-nos, até certo ponto, prever as consequências evolutivas das alterações uma vez que nos possibilitam aumentar o conhecimento da biodiversidade e evolução destas espécies. O conceito de “Filogeografia” foi apresentado por Avise et al. em 1987, e durante os últimos 25 anos teve um grande impacto na investigação, particularmente em animais. Nas plantas os resultados produzidos não têm sido tão explícitos, principalmente devido à falta de variabilidade genética aplicável à análise filogeográfica. Tem sido consideravelmente difícil encontrar um marcador genético em plantas com um poder de resolução semelhante ao DNA mitocondrial animal. No entanto a filogeografia em plantas tem-se desenvolvido bastante, principalmente nos últimos anos, com o crescimento do uso de marcadores moleculares nucleares e com a recolha de informação de fragmentos maiores do genoma cloroplastidial. O género Quercus (carvalhos) (Fagaceae) é um dos grupos mais importantes de angióspermicas lenhosas no hemisfério norte, nomeadamente em relação à diversidade de espécies, dominância ecológica e valor económico. O género é bastante antigo considerando que o fóssil mais antigo encontrado pertence ao Oligoceno (34-23 milhões de anos). Os VII carvalhos são os membros dominantes de uma grande variedade de habitats e pensa-se que existam 500-600 espécies na Terra. O sobreiro (Quercus suber L.) representa umas das espécies arbóreas mais importantes da região Oeste do Mediterrâneo, tanto económica como ecologicamente, onde define espaços florestais abertos (criados e mantidos pelo Homem) conhecidos em Portugal como “montados”. A área de distribuição do sobreiro, apesar de descontínua, vai desde a costa Atlântica do Norte de África e Península Ibérica até às regiões sudoeste de Itália, incluindo as ilhas Mediterrânicas Sicília e Sardenha, assim como as zonas costeiras do Mediterrâneo da Argélia e Tunísia. As florestas de sobreiro cobrem uma área total de cerca de 2,2 milhões de hectares, de onde são extraídas 340.000 toneladas/ano de cortiça. As maiores extensões de área coberta estão localizadas em Portugal com cerca de 700.000 hectares, correspondendo a 21% da área florestal Portuguesa e 30% da área mundial de produção de cortiça. O sobreiro tem sido usado desde a Antiguidade para a produção de cortiça e este produto natural apresenta um grande valor económico. As maiores ameaças contudo, são enfrentadas pelas populações naturais e marginais, que muitas vezes são pequenas
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