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Barbara Dos Santos Correia As Ómicas Ligadas À Ecofisiologia Do Universidade de Aveiro Departamento de Biologia 2017 Barbara dos Santos As ómicas ligadas à ecofisiologia do Eucalipto: Correia desvendando a tolerância ao stress numa espécie florestal Linking omics and ecophysiology in Eucalyptus: unravelling stress tolerance in a forest species This page has been left blank intentionally Universidade de Aveiro Departamento de Biologia 2017 Barbara dos Santos As ómicas ligadas à ecofisiologia do Eucalipto: Correia desvendando a tolerância ao stress numa espécie florestal Linking omics and ecophysiology in Eucalyptus: unravelling stress tolerance in a forest species Tese apresentada à Universidade de Aveiro para cumprimento dos requisitos necessários à obtenção do grau de Doutor em Biologia, realizada sob a orientação científica da Doutora Glória Catarina Cintra da Costa Pinto, professora auxiliar convidada do Departamento de Biologia da Universidade de Aveiro e coorientação dos Doutores Luis Valledor González, investigador de pós-doutoramento do Programa Ramón y Cajal na Universidade de Oviedo, e Robert Douglas Hancock, investigador principal no Instituto James Hutton. Apoio financeiro do FEDER no âmbito do programa COMPETE – Programa Operacional Fatores de Competitividade, e da FCT – Fundação para a Ciência e Tecnologia, através do projeto Ecophysiolyptus (PTDC/AGR- CFL/112996/2009) e bolsa SFRH/BD/86448/2012. o júri the jury presidente president Prof. Doutor Rui Luis Andrade Aguiar professor catedrático da Universidade de Aveiro arguente examiner Prof. Doutor Carlos Manuel Correia professor associado do Departamento de Biologia e Ambiente, Escola de Ciências da Vida e do Ambiente, Universidade de Trás-os-Montes e Alto Douro arguente examiner Doutor Herlânder Anselmo Queirós Pereira Azevedo investigador auxiliar do CIBIO-InBIO, Universidade do Porto arguente examiner Prof. Doutora Maria Margarida Branco de Brito Tavares Tomé professora catedrática do Departamento de Engenharia Florestal, Instituto Superior de Agronomia, Universidade de Lisboa arguente examiner Doutor Artur Jorge da Costa Peixoto Alves equiparado a investigador principal do Departamento de Biologia, Universidade de Aveiro orientadora supervisor Doutora Glória Catarina Cintra da Costa Pinto professora auxiliar convidada do Departamento de Biologia, Universidade de Aveiro agradecimentos Mais do que um extenso trabalho científico, esta tese representa um percurso, acknowledgments começou bem antes do seu tempo, e espero que possa perdurar. E como é difícil expressar tudo o que devo a tantos que foram cruzando este caminho. More than an extensive scientific work, this thesis represents a route, it started long before its time, and I hope it will persist. And how difficult it is to express what I owe so many that have crossed this path. À mãe Isabel e ao pai Baltasar, os melhores orientadores que a vida me podia ter dado, devo os melhores ensinamentos, conselhos e incentivos, o amor e a compreensão. Devo-lhes a força e a coragem que me movem. À Dra Glória Pinto, a grande comandante desta tese e exemplo ímpar de resiliência, devo a confiança, a paciência, o respeito e tantos ensinamentos científicos. Obrigada, Professora! Ao Dr Luís Valledor devo a instrução, o carinho e a boa disposição. Agradeço todos os desafios e, principalmente, a confiança de que os superaria! I owe Dr Rob Hancock the trust, the constant corrections, teaching, and many fancy beers. How fortunate I am that you have accepted to be my supervisor! A outros devo também azos de orientação: Dra Marta Pintó-Marijuan, Dra Mónica Meijón, Engº Armando Costa, Dr Bruno Castro e Dr Artur Alves. A todos os colegas do FunPlant devo inumeráveis discussões frutíferas e gargalhadas partilhadas. Aos colegas de Oviedo devo instrução, amizade e boas gargalhadas. Ao Chus e à Mónica E. devo também a receita da tortilla! I owe all colleagues at the James Hutton Institute the scientific shares and the warm affection, so important when we are far from home. À Dra Lucinda Neves, bem como à Altri Florestal e ao Engº Luís Leal, devo disponibilidade, simpatia e o material vegetal utilizado. Aos amigos devo tanto. À Isabel, à Tânia, à Rosa, à Susana e à Ni devo o apoio e a honestidade de uma amizade tão longa. Ao David, devo a compreensão e o respeito de uma amizade inabalável. À Fernanda, ao Nuno, à Verónica, ao Fernando, ao Pedro e ao Bruno devo as risadas, a descontração e a harmonia de uma amizade tão leve. À Cláudia e à Joana, devo muito mais: devo uma amizade e uma lealdade sem fronteiras, devo horas de confidência e incentivo que não me deixaram desistir de caminhar. A todos os outros amigos, que encontrei no MicroLab, no Cibio e por tantos outros lugares, devo também muitas horas felizes. Ao Golias devo a singularidade de amor e companhia que só os nossos animais sabem partilhar. Ao André devo metade desta tese. Devo dois anos de cumplicidade descomplicada. Like all magnificent things, it's very simple – Natalie Babbitt. um brinde à amizade! Cheers! https://www.youtube.com/watch?v=W8jaOQ639ds palavras-chave Árvore, Eucalyptus globulus, fisiologia, proteómica, metabolómica, expressão genética, seca, calor, recuperação resumo As plantações de eucalipto estão entre as mais produtivas do mundo inteiro. Em Portugal e Espanha, são amplamente utilizadas na produção de polpa e como fonte de energia. No entanto, o clima mediterrânico da região, com secas de verão cada vez mais severas, afeta negativamente o crescimento do eucalipto e aumenta a sua mortalidade. Esta tese de doutoramento tem como objetivo desvendar a tolerância à seca da espécie Eucalyptus globulus, investigando e interligando informação dos processos que medeiam o défice hídrico e a reidratação, desde a regulação genética e molecular até às respostas fisiológicas e desempenho da planta, utilizando dois genótipos distintos e diferentes ensaios experimentais. Esta tese está estruturada sob a forma de estudos científicos (capítulos 2, 3, 4 e 5), precedidos por uma introdução geral (capítulo 1), e termina com as notas finais (capítulo 6). O capítulo 2 descreve um ensaio de estufa e um défice hídrico imposto lentamente, e está dividido em 3 subcapítulos. Dois genótipos (AL-18 e AL-10) foram sujeitos a um período de stress hídrico de 3 semanas com duas intensidades diferentes (18% e 25% da capacidade de campo), seguido de uma semana de reidratação. A recuperação foi avaliada um dia e uma semana depois da reidratação. Várias fitohormonas foram monitorizadas nas folhas, seiva xilémica e raízes, 2 h, 4 h, 24 h e 168 h depois da reidratação. A falta de água reduziu a altura, a biomassa, o potencial hídrico e as trocas gasosas. Pelo contrário, os níveis de pigmentos, parâmetros da fluorescência da clorofila e MDA aumentaram. Os níveis de ABA e de ABA-GE aumentaram, enquanto o JA diminuiu nas folhas e aumentou na seiva xilémica. Durante a recuperação, a maioria das alterações fisiológicas e bioquímicas provocadas pelo stress reverteram. Análises comparativas do proteoma (analisado por eletroforese em gel diferencial) e do metaboloma (analisado por cromatografia gasosa com espetrometria de massa) permitiram a separação de 2031 pontos peptídicos, dos quais 217 foram identificados, e a deteção de 121 metabolitos polares. A análise do clone resiliente AL-18, que apresentou uma rede de resposta bem distinta do clone responsivo AL-10, reforçou o papel de proteínas específicas da fotossíntese e relacionadas com a defesa como intermediários chave na tolerância à seca e revelou novos intermediários: glutamina sintetase, malato desidrogenase e isoflavona redutase. O capítulo 3 diz respeito a um ensaio em câmara climática e a uma rápida escassez de água, e está dividido em 2 subcapítulos. A expressão relativa de 12 transcritos foi analisada por PCR quantitativo em dois clones com diferentes graus de tolerância (AL-18 e AL-13) depois de 7 e 11 dias sem qualquer rega e posterior reidratação. A rápida escassez de água foi mais prejudicial para as plantas do que o défice hídrico imposto lentamente, com maior visibilidade no clone AL-13 que revelou morte de algumas plantas. resumo (cont.) Indicadores moleculares potencialmente ligados a uma tolerância aumentada foram identificados: rubisco ativase (RCA), ferredoxina-NADP(H) oxidorredutase (FNR), malato desidrogenase mitocondrial (mMDH) catalase peroxissomal (CAT) e isoflavona redutase (IFR). De seguida, vários marcadores bioquímicos de stress oxidativo e padrões de metilação do DNA foram quantificados nas folhas do clone AL-18. As alterações detetadas utilizando indicadores globais e específicos refletiram a indução de complexas modificações redox e de metilação do DNA, que ocorrem paralelamente durante a imposição e interrupção do stress. O capítulo 4 reporta um ensaio de campo: o conjunto de indicadores de seleção de tolerância hídrica identificado anteriormente foi testado em AL-18 e AL-13 plantados no campo. Algumas das plantas foram regadas artificialmente (IR) e outras foram deixadas nas condições ambientais de precipitação reduzida (NI) durante seis semanas e meia antes de voltar a regar. O clone AL-18 mostrou pouca variação nas condições testadas, e as alterações encontradas no clone AL-13 realçaram a indução do metabolismo fotossintético e fotorespiratório após a reidratação artificial. Estes resultados mostraram que as respostas das plantas no campo não podem ser extrapoladas a partir do estudo de um stress aplicado individualmente, particularmente no contexto de encontrar marcadores de seleção. O capítulo 5 descreve um ensaio em câmara climática que testou o efeito isolado e combinado
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