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Identificação E Caracterização Dos Genes Da Família MADS-Box Em Passiflora Organensis

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Identificação E Caracterização Dos Genes Da Família MADS-Box Em Passiflora Organensis UNIVERSIDADE ESTADUAL DE CAMPINAS INSTITUTO DE BIOLOGIA DEPARTAMENTO DE BIOLOGIA VEGETAL HELENA AUGUSTO GIOPPATO Identificação e caracterização dos genes da família MADS-box em Passiflora organensis ORIENTADOR: MARCELO CARNIER DORNELAS CAMPINAS 2019 HELENA AUGUSTO GIOPPATO Identificação e caracterização dos genes da família MADS-box em Passiflora organensis Dissertação apresentada ao Instituto de Biologia da Universidade Estadual de Campinas como parte dos requisitos exigidos para a obtenção do título de Mestra em Biologia Vegetal. Orientador: Marcelo Carnier Dornelas Este trabalho corresponde à versão final dissertação defendida pela aluna Helena Augusto Gioppato e orientada pelo Prof. Dr. Marcelo Carnier Dornelas CAMPINAS 2019 FICHA CATALOGRÁFICA BANCA EXAMINADORA Prof. Dr. Marcelo Carnier Dornelas Prof. Dr. Fábio Pinheiro Prof. Dr. Jean Carlos Cardoso A Ata da defesa com as respectivas assinaturas dos membros encontra-se no SIGA/Sistema de Fluxo de Dissertação/Tese e na Secretaria do Programa da Unidade. AGRADECIMENTOS À Universidade Estadual de Campinas - UNICAMP, aos seus professores, alunos e funcionários, que tornam possível a expansão da formação acadêmica para a formação pessoal. Ao Instituto de Biologia da Unicamp pelos cursos em Ciências Biológicas e Mestrado em Biologia Vegetal, que me trouxeram até esta dissertação. Ao CNPq por ter financiado este projeto de pesquisa. Ao Centro de Energia Nuclear na Agricultura – CENA/ ESALQ – Universidade de São Paulo, por todo apoio desde o cultivo das plantas e análises de biologia molecular até as amizades que lá encontrei, uma delas a doutoranda Tatiana de Souza Moraes, com quem aprendi e ainda aprendo muito sobre biologia e sobre a vida. Ao orientador Prof. Dr. Marcelo Carnier Dornelas que desde a iniciação científica me orienta, me ensina e me incentiva a ir além, sempre confiando muito em meu trabalho, por toda a dedicação em me orientar este tempo todo e em especial neste projeto. Ao Prof. Dr. Diego Ismael Rocha, por todos os ensinamentos durante minha iniciação científica, os quais com toda certeza foram fundamentais para que chegasse até aqui. Às Profas. Dras. Sandra Maria Carmello-Guerreiro e Adriana Martinelli Pinheiro, convidadas para as bancas examinadoras de etapas anteriores à defesa, por todas as sugestões e contribuições para o melhor desenvolvimento do trabalho. Aos Profs. Drs. Fábio Pinheiro e Jean Carlos Cardoso, convidados para a banca examinadora, por aceitarem o convite e contribuírem com o trabalho. Ao meu pai Silvio por ter me apresentado desde pequena o universo da ciência e pelo incentivo em permanecer na área, sempre contribuindo para a minha formação pessoal e intelectual, à minha mãe Georgete pelo amor incondicional e por todo apoio e incentivo que me dá para seguir em frente com minhas escolhas, e aos meus irmãos Gustavo e Fernando pela amizade e lições de vida que temos juntos. Às amizades que me acompanham desde a graduação pela companhia e apoio constantes, que sempre fizeram a diferença no meu dia a dia. Agradeço especialmente a minha grande amiga Carla Maneira da Silva por estar sempre presente para me apoiar e compartilhar as alegrias e mazelas da vida acadêmica. Às amigas, Jusceley, Tatiane, Mariana e Bruna, companheiras de bancada e campo que alegraram meus dias no laboratório, cada uma a sua maneira. Aos amigos do Departamento de Biologia Vegetal, especialmente à Natália Volpi, Juan e Franklin Magnum, que comigo compartilharam o conhecimento e a amizade, e me auxiliaram sempre que necessário, por toda a ajuda, pela companhia e aprendizados que tornaram o trabalho ainda mais prazeroso. RESUMO Uma importante questão da biologia vegetal é como as novidades morfológicas foram produzidas e como elas afetaram o processo evolutivo. Para se estudar essa questão, foi necessário lançar mão de estudos na área da biologia do desenvolvimento. Estes estudos levaram à descrição de diversos genes reguladores que controlam muitos processos de desenvolvimento em todos os organismos. Alguns destes genes fazem parte de uma família multigênica chamada MADS-box, que entre as angiospermas, é responsável por regular vários processos de seu desenvolvimento, dentre eles o desenvolvimento floral. Duplicações ou perdas parálogos de MADS-box podem alterar o padrão de desenvolvimento das estruturas florais. O gênero Passiflora é um dos grupos de angiospermas que possui indivíduos que produzem estruturas complexas cujas origens ainda não foram elucidadas, como os filamentos da corona. Sendo assim, e considerando também o seqüenciamento recente do genoma de uma espécie de Passiflora, Passiflora organensis, o presente trabalho identificou e caracterizou 72 genes da família MADS-box nesta espécie, bem como possíveis expansões ou retrações da família, e a partir destes dados, inferir possíveis explicações para a morfologia floral distinta encontradas dentro do gênero Passiflora. Oito genes foram relacionados ao desenvolvimento de estames e carpelos. Análises da expressão diferencial destes genes revelaram um papel potencial na diversificação das estruturas florais em Passilfora. ABSTRACT An important issue in plant biology is how the morphological novelties were produced and how they affected the evolutionary process. In order to study these subjects, it was necessary to use studies in the field of developmental biology. These studies have led to the description of several regulatory genes that control many developmental processes in all organisms. Some of these genes are part of a multigenic family called MADS-box, which among the angiosperms, is responsible for regulating various processes of its development, among them the floral development. Doubling or loss paralogs of MADS-box may alter the pattern of development of floral structures. The genus Passiflora is one of the groups of angiosperms that has individuals that produce complex structures whose origins have not yet been elucidated, such as the corona filaments. The present work identified and characterized 72 genes of the MADS-box family in this species, as well as possible expansions or retractions of the family, and from this data, to infer possible explanations for the distinct floral morphology found within the genus Passiflora. Eight genes were related to the development of stamens and carpels. Analyzes of the differential expression of these genes revealed a potential role in the diversification of floral structures in Passiflora. LISTA DE ABREVIATURAS E SIGLAS Genes MADS-box de Arabidopsis thaliana: AP1 APPETALA 1 RT-qPCR e RT-PCR: AP2 APETALA 2 Ct Cycle Threshold (ciclo no AP3 APETALA 3 qual a reação atinge o limiar AG AGAMOUS da fase exponencial) CAL CAULIFLOWER FW Primer no sentido ‘forward’ FUL FRUITFULL RV Primer no sentido ‘reverse’ PI PISTILLATA Tm Temperatura de anelamento STK SEEDSTICK SHP SHATTERPROOF Genes de referência: SEP SEPALLATA CAC Clathrin adaptor complex; TM6 TOMATO 6 ADAPTOR PROTEIN-2 GOA GORDITA SAND SAND family protein; TT16 TRANSPARENT TESTA 16 MONENSIN SENSITIVITY1 Genes MADS-box de Passiflora organensis: PoAP1 APPETALA 1 PoAP2 APETALA 2 PoAP3 APETALA 3 PoAG AGAMOUS PoPIci PISTILLATA com íntrons PoPIsi PISTILLATA sem íntrons PoSTK SEEDSTICK PoSHP SHATTERPROOF PoSEP SEPALLATA PoTM6 TOMATO 6 PoGOA GORDITA PoTT16 TRANSPARENT TESTA 16 SUMÁRIO RESUMO ABSTRACT LISTA DE ABREVIATURAS E SIGLAS INTRODUÇÃO GERAL ......................................................................................................... 11 OBJETIVOS GERAIS ............................................................................................................. 18 OBJETIVOS ESPECÍFICOS ................................................................................................... 19 CAPÍTULO I: Identificação dos genes ortólogos aos genes da família MADS-box em Passiflora organensis .............................................................................................................. 20 Resumo ................................................................................................................................. 20 Abstract ................................................................................................................................. 20 Introdução ............................................................................................................................. 21 Objetivos ............................................................................................................................... 25 Materiais e métodos .............................................................................................................. 25 Material genômico ............................................................................................................ 25 Identificação dos genes da família MADS-box em Passiflora organensis ...................... 25 Caracterização estrutural dos genes MADS-box de Passiflora organensis...................... 26 Resultados e discussão .......................................................................................................... 27 Identificação e filogenia dos genes MADS-box de Passiflora organensis....................... 27 Estrutura gênica dos genes MADS-box de Passiflora organensis ................................... 30 CAPÍTULO II: Identificação dos genes de classe B da família MADS-box em Passiflora organensis ................................................................................................................................ 32 Resumo ................................................................................................................................
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