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“Luiz De Queiroz” Bactérias Associadas Às Cactáceas Da Caatinga

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“Luiz De Queiroz” Bactérias Associadas Às Cactáceas Da Caatinga Universidade de São Paulo Escola Superior de Agricultura “Luiz de Queiroz” Bactérias associadas às cactáceas da Caatinga: promoção de crescimento de plantas sob estresse hídrico Vanessa Nessner Kavamura Tese apresentada para obtenção do título de Doutor em Ciências. Área de concentração: Microbiologia Agrícola Piracicaba 2012 Vanessa Nessner Kavamura Bacharel em Ciências Biológicas Bactérias associadas às cactáceas da Caatinga: promoção de crescimento de plantas sob estresse hídrico Orientador: Prof. Dr. ITAMAR SOARES DE MELO Tese apresentada para obtenção do título de Doutor em Ciências. Área de concentração: Microbiologia Agrícola Piracicaba 2012 Dados Internacionais de Catalogação na Publicação DIVISÃO DE BIBLIOTECA - ESALQ/USP Kavamura, Vanessa Nessner Bactérias associadas às cactáceas da Caatinga: promoção de crescimento de plantas sob estresse hídrico / Vanessa Nessner Kavamura.- - Piracicaba, 2012. 244 p: il. Tese (Doutorado) - - Escola Superior de Agricultura “Luiz de Queiroz”, 2012. 1. Bactérias 2. Caatinga 3. Cactáceas 4. Crescimento vegetal 5. Estresse hídrico 6. Mandacaru 7. Rizosfera 8. Semiárido I. Título CDD 633.39 K21b “Permitida a cópia total ou parcial deste documento, desde que citada a fonte – O autor” 3 DEDICO Aos meus pais, Teruo e Maria Catarina, por todo o amor, apoio e incentivo incondicionais, em todos os momentos de minha vida. Eu amo vocês! OFEREÇO Aos meus queridos avós, Anton (ota), Anna (oma) Tsuguio (ditian) e Kashiko (batian), que muito contribuíram para a minha formação pessoal. Saudades eternas... vocês sempre estarão em meu coração! 4 5 AGRADECIMENTOS É muito difícil expressar em palavras a gratidão que sinto por todos que contribuíram direta ou indiretamente para a realização deste trabalho. Entretanto, tentarei fazê-lo da melhor forma possível, listando aqueles que contribuíram imensamente, não somente para a conclusão de alguns anos de estudo, mas também para a minha formação. À Deus e à Meishu-Sama, pela vida, agradeço humildemente; À Universidade de São Paulo e à Capes pela concessão da bolsa de doutorado; Ao meu orientador, Dr. Itamar Soares de Melo, pela oportunidade, confiança e orientação, muito obrigada! À Embrapa Meio Ambiente, pela infraestrutura oferecida para realização da pesquisa; Ao prof. Dr. Fernando Dini Andreote, pela confiança ao me indicar ao Itamar e pela paciência nos ensinamentos; Ao pesquisador, Dr. Rodrigo Mendes, pelas correções do trabalho e pelas valiosas dicas; Ao Dr. Rodrigo Gouvêa Taketani, obrigada pelo auxílio aos assuntos referentes à biologia molecular e também pela amizade! Ao Dr. Tiago Domingues Zucchi, obrigada pelas questões relacionadas à taxonomia; Aos amigos de trabalho e de lazer: Luciana Aparecida Ávila (Lu), Mírian Lobo Sáber (Mí), Natália Franco Taketani, Suikinai Nobre Santos (Suiki), Wallace Rafael de Souza (Wall), muito obrigada pelas horas de descontração e boas risadas durante as viagens, passeios e comilança !!! Aos colegas e amigos do LMA com os quais convivo todos os dias e que tornam a ida ao laboratório mais especial: Ana Gabriele Barbosa Casteliani, Andiale Pinto dos Santos, Carlos Eduardo Oliveira da Silva, Cassiano Forner, José Abrahão Haddad Galvão, Leonardo José da Silva (Léo), Lúcio Bertoldo Costa, Michelli de Souza dos Santos, Milena Duarte Lançoni, Rafael Eduardo Silva, Regiane Iost (Figura) e Zayame Vegette Pinto. Aos novos colegas: Clederson Ferreira e Lucas Dantas Lopes, obrigada pelas discussões referentes às análises utilizadas. Aos colegas e amigos do LMA que deixaram saudades: Alexandre Visconti (Tiozinho), Jayme Pieroni Junior, Lívia Mendes, Luis Alexandre Sereda (Alê), Maria Augusta de Camargo Ferraz Machado (Guta), Osvaldo Luiz Ferreira Junior e Rachel Temperani Amaral Machado. Aos funcionários do Laboratório de Microbiologia Ambiental (LMA) da Embrapa - Meio Ambiente: Anamaria Ferreira Mayer Dentzien, Elke Simoni Dias Vilela (pelas questões relacionadas à química), João Luiz da Silva (pelo grande auxílio nas coletas), Márcia Maria Parma (pelas análises do MIDI e sequenciamento), Rosely dos Santos Nascimento (por todas as dicas), Tatiana Rigamonte Fernandes, meu sincero agradecimento por todo o auxílio e paciência! Aos funcionários do P10 da Embrapa - Meio Ambiente, muito obrigada pelo auxílio nas atividades de campo, em especial ao Henrique Barros Vieira pelos espaços concedidos aos experimentos e à Célia Batista da Silva de Lima por todo o carinho! 6 Aos demais funcionários da Embrapa: pesquisadores, analistas, assistentes, pessoal da manutenção, motoristas e pessoal da limpeza, minhas lembranças! Ao Tomio, por todo o companheirismo em todos os momentos de minha vida, por me aturar nas horas estressantes, por me auxiliar em alguns experimentos, enfim, por tudo, muito obrigada! 7 “Só existem dois dias no ano que nada pode ser feito. Um se chama ontem e o outro se chama amanhã, portanto hoje é o dia certo para amar, acreditar, fazer e principalmente viver.” Dalai Lama 8 9 SUMÁRIO RESUMO..................................................................................................................................13 ABSTRACT..............................................................................................................................15 1 INTRODUÇÃO ..................................................................................................................... 33 2 OBJETIVOS E QUESTÕES BIOLÓGICAS ........................................................................ 35 3 REVISÃO BIBLIOGRÁFICA .............................................................................................. 37 3.1 Região semiárida do nordeste brasileiro: bioma Caatinga ................................................. 37 3.2 Micro-organismos ............................................................................................................... 42 3.2.1 De solo e rizosfera ........................................................................................................... 42 3.2.2 Extremófilos: ambientes áridos e semiáridos .................................................................. 43 3.2.3 Ferramentas moleculares de análise ................................................................................ 45 3.2.4 Rizobactérias promotoras de crescimento de plantas (RPCP)......................................... 46 3.2.4.1 Mecanismos diretos de ação ......................................................................................... 47 3.2.4.2 Mecanismos indiretos de ação ...................................................................................... 51 3.3 Estresse ambiental .............................................................................................................. 52 3.3.1 Respostas fisiológicas dos organismos ao estresse.......................................................... 53 3.3.1.1 Produção de biofilme .................................................................................................... 53 3.3.1.2 Produção de exopolissacarídeos (EPS) ........................................................................ 54 3.3.1.3 Produção de osmólitos intracelulares ........................................................................... 55 3.3.2 Estresse hídrico: tolerância x resistência ......................................................................... 55 3.3.2.1 O papel dos micro-organismos na tolerância vegetal ao estresse hídrico .................... 56 Referências ............................................................................................................................... 58 4 COMUNIDADE BACTERIANA DE SOLO E RIZOSFERA DE CEREUS JAMACARU DURANTE O PERÍODO CHUVOSO E DE SECA ............................................................... 74 Resumo......................................................................................................................................74 Abstract.....................................................................................................................................75 4.1 Introdução ........................................................................................................................... 76 4.2 Desenvolvimento ................................................................................................................ 77 4.2.1 Material e Métodos .......................................................................................................... 77 4.2.1.1 Área de estudo e coleta das amostras de solo e rizosfera de Cereus jamacaru durante o período chuvoso e de seca ........................................................................................................ 77 4.2.1.2 Extração de DNA metagenômico de solo e rizosfera ................................................... 81 10 4.2.1.3 Estrutura da comunidade bacteriana por meio da técnica de polimorfismo dos fragmentos terminais de restrição (Terminal Restriction Fragment Length Polymorphism – T- RFLP).................... ................................................................................................................... 81 4.2.1.4 Análise da comunidade bacteriana por meio do sequenciamento parcial do gene 16S rRNA em larga escala .............................................................................................................. 84 4.2.2 Resultados e Discussão ................................................................................................... 87 4.2.2.1 Análise de T-RFLP do gene 16S rRNA de Bacteria ..................................................
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