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Tese Camila Pegoraro.Pdf i UNIVERSIDADE FEDERAL DE PELOTAS Programa de Pós-Graduação em Agronomia Faculdade de Agronomia Eliseu Maciel Tese Bases moleculares da resposta a estresses ambientais em vegetais: arroz, tomate e pêssego Camila Pegoraro Pelotas, 2012 ii Camila Pegoraro Bases moleculares da resposta a estresses ambientais em vegetais: arroz, tomate e pêssego Tese apresentada ao Programa de Pós- Graduação em Agronomia da Universidade Federal de Pelotas, como requisito parcial à obtenção do título de Doutora em Ciências (área do conhecimento: Fitomelhoramento). Orientador: Dr. Antonio Costa de Oliveira – FAEM/UFPel Co-orientadores: Dr. Cesar Valmor Rombaldi – FAEM/UFPel Dr. Luciano Carlos da Maia – FAEM/UFPel Dr. Livio Trainotti – UNIPD Pelotas, 2012. iii Dados de catalogação na fonte: (Marlene Cravo Castillo – CRB-10/744) P376b Pegoraro, Camila Bases moleculares da resposta a estresses ambientais em vêgetais: arroz, tomate e pêssego / Camila Pegoraro; orientador Antonio Costa de Oliveira; co-orientadores Cesar Valmor Rombaldi, Luciano Carlos da Maia e Livio Trainotti - Pelotas, 2012. 229f.: il.- Tese (Doutorado) –Programa de Pós-Graduação em Agron omia. Faculdade de Agronomia Eliseu Maciel. Universidade Federal de Pelotas. Pelotas, 2012. 1.Expressão gênica 2.Oryza sativa 3.Solanum lycopersicum 4.Prunus persica 5.Estresses abióticos I.Oliveira, Antonio Costa de(orientador) I.Título. CDD 574.87 iv Banca Examinadora: Dr. Antonio Costa de Oliveira – FAEM/UFPel (presidente) Dr. Cesar Valmor Rombaldi – FAEM/UFPel Dra. Roberta Manica Berto – FAEM/UFPel Dr. Sandro Bonow – EMBRAPA Clima Temperado Dra. Vera Maria Quecini – EMBRAPA Uva e Vinho v Agradecimentos A Deus pela vida, força e proteção em todas os momentos. A toda minha família, principalmente meus pais Isaias e Iraci e meu irmão Cassiano pelo amor, incentivo e apoio. Ao meu noivo Marcos pelo amor, paciência e compreensão. Aos meus orientadores professores Antonio, Cesar e Luciano pela transmissão de conhecimentos, incentivo, apoio, ajuda, compreensão, amizade e confiança. Aos colegas do Centro de Genômica e Fitomelhoramento Adriana, Daisy, Daniel, Daniela, Diego, Elisane, Francieli, Frederico, Gabriela, Glacy, Henrique, Leomar, Letícia, Mateus, Monalise, Naciele, Rafael, Renata, Rodrigo, Sidney, Solange, Vivian, Viviane, Taciane, Taiane, Tatiane, em especial aos colegas Mariana e Railson pela amizade, convivência e troca de experiências. Aos estagiários do Centro de Genômica e Fitomelhoramento Bianca, Carolina, Cristiano, Daiana, Eder, Ederson, Fabiane, Fernando (In Memorian), Helaine, Michel, Pagiel, Raissa, Ricardo, Rodrigo e Victoria, pela amizade e convivência. Aos colegas do Laboratorio di Biologia Molecolare Alessandro, Alice, Anna, Nicola, Margueritta, Rahim e Rossela, pela receptividade, amizade, apoio e ajuda. Aos colegas Alice, Aline, Daniel, Liliane, Mariana, Railson, Roberta e Taciane pelo auxílio na realização das atividades. A todos meus amigos pelo apoio e incentivo. À professora da UFRGS Marcia Maria Auxiliadora Naschenveng Pinheiro Margis pela ajuda no trabalho de transformação genética. Ao Programa de Pós-graduação em Agronomia/FAEM-UFPel pela oportunidade de realização do curso de doutorado, e a Università degli Studi di Padova pela possibilidade de doutorado sanduíche. À CAPES pelo financiamento da bolsa de doutorado e ao CNPq pelo financiamento da bolsa de doutorado sanduíche. E a todos que de alguma forma contribuíram para a realização deste trabalho. vi Resumo PEGORARO, Camila. Bases moleculares da resposta a estresses ambientais em vegetais: arroz, tomate e pêssego. 2012. 226p. Tese (Doutorado) - Programa de Pós-Graduação em Agronomia. Universidade Federal de Pelotas, Pelotas. Estresses ambientais representam grandes ameaças à agricultura e, por consequência, à segurança alimentar, uma vez que limitam a produtividade da maioria das plantas cultivadas. Buscando identificar as respostas das plantas frente a estresses abióticos e suas relações com a tolerância sob diferentes condições ambientais, conceitos e técnicas de biologia molecular, biologia celular e genética vêm sendo continuamente utilizadas. A identificação de genes responsivos a estresses pode ser utilizada na obtenção de plantas tolerantes a condições ambientais adversas através de manipulação genética ou cruzamentos direcionados entre diferentes genótipos. Desta forma, este estudo teve como objetivo estabelecer o perfil de expressão de genes pertencentes às famílias ERF, HSP, HSF, TIM/TOM e TIC/TOC, em arroz e tomate sob condições de estresse por deficiência de O2. Além disso, buscou-se estabelecer o perfil transcricional de pêssegos sob condição de estresse por frio. Os resultados obtidos demonstram que em arroz, alguns membros da família ERF apresentam expressão específica de acordo com o agente estressor, enquanto que outros apresentam sobreposição de expressão entre os estresses, além disso, dentro da família ERF alguns genes apresentam regulação complexa e outros regulação simples. Em arroz, alguns genes codificadores de proteínas pertencentes aos complexos TIM/TOM e TIC/TOC apresentaram altos níveis de expressão 24 horas após estresse por anoxia. Entretanto, após 72 horas sob condições de anoxia, ocorreu a inibição da expressão da maioria dos genes pertencentes a estes complexos, sugerindo que possa haver uma interrupção no transporte de proteínas para a mitocôndria e cloroplasto durante este período de estresse. Ao comparar cultivares de arroz tolerante e sensível ao alagamento, verificou-se que houve um acúmulo de transcritos dos genes codificadores de HSPs em ambas as cultivares. Entretanto, a maioria dos genes avaliados foi mais responsiva ao estresse por anoxia na cultivar sensível, do que na cultivar tolerante, sugerindo que genes HSPs têm relação importante na resposta ao estresse por anoxia, mas não estão diretamente envolvidos na tolerância. Em frutos de tomate armazenados sob hipoxia dois genes codificadores de sHSPs foram altamente vii induzidos por baixos níveis de oxigênio, indicando uma ação primária destes genes na manutenção da homeostase celular após este estresse. Em pêssegos, a aplicação de ácido giberélico atua na prevenção de danos ocasionados pelo frio através da indução de genes associados ao metabolismo de lipídios, à degradação da parede celular e à síntese de trealose. Palavras-chave: expressão gênica, Oryza sativa, Solanum lycopersicum, Prunus persica, estresses abióticos. viii Abstract PEGORARO, Camila. Molecular basis of response to environmental stresses in plants: rice, tomato and peach. 2012. 226p. Thesis (Doctoral degree) - Programa de Pós-Graduação em Agronomia. Universidade Federal de Pelotas, Pelotas. Abiotic stresses represent large threats to agriculture and food security, since they limit the productivity of the majority of cultivated plants. In order to identify the responses of plants to abiotic stresses and their associations with tolerance under different environmental conditions, concepts and techniques in molecular biology, cell biology and genetics have been used. The identification of genes responsive to stresses can be used to obtain plants tolerant to adverse environmental conditions by genetic transformation or directed crosses. Thus, this study had the objective the establishing of expression profiles of gene belonging to the ERF, HSP, HSF, TIM/TOM and TIC/TOC families in rice and tomato under low O2 stress. Also, the transcriptional profile of peaches under cold stress as obtained. The results indicate that in rice, some members of the ERF family present specific expression according to the stressing agent, while others have overlapping expression among different stresses. Besides, within the ERF family, some genes do present a complex regulation and others a simple regulation. In rice, some genes encode proteins belonging to the TIM/TOM and TIC/TOC complexes present high expression levels 24 hours after anoxia stress. However, after 72 hours under anoxia, the majority of genes were inhibited, suggesting an interruption on the transport of proteins to the mitochondria and chloroplast during the stress period. When the HSP expression profiles of flooding tolerant and sensitive rice cultivars were compared, transcript accumulation occurred for both cultivar types. However, the majority of genes evaluated were more responsive in the sensitive cultivar than in the tolerant, indicating that HSPs have an important association in the response to anoxia stress but are not directly involved in the tolerance. In tomato fruits stored under hypoxia, two sHSPs genes were highly induced by low oxygen levels, indicating a primary action of these genes on keeping cell homeostasis after the stress. In peaches, the application of gibberellic acid acts on preventing the damaged caused by cold by the induction of genes associated to lipid metabolism, cell wall degradation and trehalose synthesis. ix Keywords: gene expression, Oryza sativa, Solanum lycopersicum, Prunus persica, abiotic stress. x LISTA DE FIGURAS 3. CAPÍTULO II – Regulação e perfil de expressão de genes ERFs em condições de estresse Figura 1 Perfil de expressão de genes ERFs em plantas de arroz submetidas a diferentes estresses. O perfil de expressão é apresentado como heat map, gerado pelo programa Genevestigator. A: grupo 1; B: grupo 2 ....................................................................................................... 71 Figura 2 Especificidade de expressão para genes induzidos (A) e reprimidos (B). Os diagramas foram gerados utilizando-se o programa Venny .......... 72 Figura
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