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Universidade Federal De Pernambuco Centro De Ciências Biológicas Programa De Pós-Graduação Em Biologia Vegetal

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Universidade Federal De Pernambuco Centro De Ciências Biológicas Programa De Pós-Graduação Em Biologia Vegetal Universidade Federal de Pernambuco Centro de Ciências Biológicas Programa de Pós-Graduação em Biologia Vegetal CITOGENÉTICA, ORIGEM E EVOLUÇÃO DE Nothoscordum gracile (Ailton) Stearn (ALLIACEAE) E ESPÉCIES AFINS DA SECÇÃO Inodorum Recife Fevereiro 2008 Universidade Federal de Pernambuco Centro de Ciências Biológicas Programa de Pós-Graduação em Biologia Vegetal CITOGENÉTICA, ORIGEM E EVOLUÇÃO DE Nothoscordum gracile (Ailton) Stearn (ALLIACEAE) E ESPÉCIES AFINS DA SECÇÃO Inodorum Dissertação apresentada por Luiz Gustavo Rodrigues Souza como requisito para obtenção do grau de Mestre em Biologia Vegetal (área de concentração em Florística e Sistemática) pelo curso de Pós- Graduação em Biologia Vegetal da UFPE Orientador: Profº Marcelo Guerra (Depto. Botânica, UFPE) Recife Fevereiro 2008 Souza,, Luiz Gustavo Rodrigues Citogenética, origem e evolução de Nothoscordum gracile (Aílton) Stearn (Alliaceae) e espécies afins da secção Inodorum. / Luiz Gustavo Rodrigues Souza. – Recife: O Autor, 2008. 84 fls. .: il. Dissertação (Mestrado em Biologia Vegetal) – UFPE. CCB 1. Biologia vegetal 2. Evolução 3. Nothoscordum I.Título 575 CDU (2ª. Ed.) UFPE 575 CDD (22ª. Ed.) CCB – 2008 –082 “A distância de quem amamos traz saudade, jamais esquecimento” À minha família e amigos Dedico AGRADECIMENTOS Agradeço a todos aqueles que direta ou indiretamente contribuíram para a realização desse trabalho, em especial: Ao CNAPq pela bolsa sem a qual eu não conseguiria prosseguir nos estudos. Ao Profº. Marcelo Guerra pela excelente orientação, paciência nos momentos difíceis, broncas e sugestões (que me foram muito úteis) quando necessário e confiança em mim. À Andrea Pedrosa por todo o auxílio teórico e prático e por tentar constantemente fazer de mim uma pessoa menos “bagunceira”. À Ana Emília por toda a ajuda e conselhos tanto dos tramites legais da UFPE quanto da citogenética em si. À Cícero Carlos pela amizade, incentivo e companheirismo. Aos amigos do laboratório pela amizade e colaborações dadas à realização desse trabalho, Ana Paula, Aretuza, Eliene, Fagna, Fernando, Gabi, Joana, Lili, Rogério, Sandra, Silvana e Thiago Ribeiro. Às cachorras do laboratório: Artur, Darwin (André), Pipe (Felipe) e Thiago Thel (Cachorra mor) pela amizade e cumplicidade. Aos meus pais Regina e Valdir, irmã Poliana e todos os familiares que sempre acreditaram em mim e incentivaram a minha vinda para Pernambuco. Pelo amor e alegria nos curtos momentos em que eu pude estar em casa. À toda a turma do pensionato, em especial Dulce por todo o carinho. À todos os professores do Departamento de Botânica pela ótimas disciplinas ministradas e por expandir meus conhecimentos sobre o “Mundo Vegetal”. Ao funcionário da Pós-Graduação em Biologia Vegetal, Hidelbrando pelo apoio prestado. Aos vigilantes do Centro de Ciências Biológicas, pelo apoio prestado, o qual possibilitou minha permanência no laboratório, com segurança, nos muitos horários fora do expediente normal. Por fim, aos “novos” amigos dos laboratórios vizinhos com os quais espero ter oportunidade (em especial tempo) de compartilhar momentos felizes: Beti, Jéferson, Katarina, Diogo et al. (MTV); Bruno e Laura (Fisiologia/Ecologia). SUMÁRIO Pág. LISTA DE FIGURAS LISTA DE TABELAS RESUMO ABSTRACT 1. INTRODUÇÃO 1 2. FUNDAMENTAÇÃO TEÓRICA 2 2.1. Citotaxonomia 2 2.2. Heterocromatina: bandeamento C e fluorocromos 3 2.3. Hibridização in situ fluorescente (FISH) 6 2.4. Hibridização genômica in situ (GISH) 7 2.5. Translocações robertsonianas 8 2.6. O gênero Nothoscordum 10 2.7. Citogenética da família Alliaceae 11 2.8. Citogenética da subfamília Gilliesioideae 12 2.9. Citogenética do gênero Nothoscordum 13 2.10. Origem e evolução de Nothoscordum gracile 14 3. REFERÊNCIAS BIBLIOGRÁFICAS 16 4. MANUSCRITO A SER SUBMETIDO AO PERIÓDICO GENETICS AND 24 MOLECULAR BIOLOGY Título, autores 24 Resumo 25 Introdução 26 Material e métodos 28 Resultados 30 Discussão 31 Referências bibliográficas 33 5. MANUSCRITO A SER SUBMETIDO AO PERIÓDICO ANNALS OF 41 BOTANY Título, autores 41 Resumo 42 Introdução 43 Material e métodos 45 Resultados 47 Discussão 51 Literatura citada 55 6. CONCLUSÕES 65 7. ANEXOS 66 7.1. Normas do periódico Genetics and Molecular Biology 66 7.2. Normas do periódico Annals of Botany 71 LISTA DE FIGURAS MANUSCRITO A SER SUBMETIDO AO PERIÓDICO GENETICS AND MOLECULAR BIOLOGY Pág. Figura 1 Distribuição da heterocromatina e sítios de DNAr em 39 Nothoscordum arenarium. (a) Metáfase mostrando as bandas C (seta indica banda menor). (b) Metáfase corada com o fluorocromo CMA. (c) Metáfase após a FISH mostrando pequenas bandas brilhantes com DAPI co-localizados com centrômeros (cabeças de setas cheias), com sítios de DNAr 5S (cabeças de setas vazadas) e 45S (setas). (d) Mesma célula de b, hibridizada in situ com DNAr 5S (vermelho) e 45S (verde). (e) Núcleo interfásico corado com CMA/DAPI mostrando cromocentros menores associados ao nucléolo. (f) Idiograma com bandas CMA (amarelo), sítios de DNAr 5S (vermelho) e 45S (verde). NC, numeração cromossômica; T, tamanho cromossômico; RB, razão entre braços. Barra em (e) equivale a 10 µm. Figura 2 Metáfase de um indivíduo com uma translocação recíproca, 40 corada com CMA/DAPI (a) e hibridizada in situ (b) com DNAr 5S (vermelho) e 45S (verde). Cabeças de setas indicam possíveis pontos de quebra envolvidos na translocação. (c) Idiograma com bandas CMA (amarelo), sítios de DNAr 5S (vermelho) e 45S (verde). NC, numeração cromossômica; T, tamanho cromossômico; RB, razão entre braços. Linha nos cromossomos II e V circunscreve pares heteromórficos. (e) Barra em (b) equivale a 10 µm. MANUSCRITO A SER SUBMETIDO AO PERIÓDICO ANNALS OF BOTANY Figura 1 Metáfases de algumas espécies de Nothoscordum coradas com 61 CMA (a, c, e) e hibridizadas in situ com sonda de DNAr 5S e 45S. (b, d, f) e diacinese de N. macrostemon (n = 5) corada com CMA (g). (a, b) N. gracile 2n = 19 (NOT 1716). (c, d) Nothoscordum sp. 2n = 15 (NOT 1707). (e, f, g) N. macorstemon 2n = 10 (NOT 1710). Sondas marcadas em verde correspondem a sítios de DNAr 45S e em vermelho a sítios de 5S. Detalhes de cromossomos mostram pequenas bandas CMA+ (a, c, e) e sítios de DNAr 45S (b, d, f). Cabeças de setas indicam bancas CMA+. Barra em g equivale a 10 µm. Figura 2 Metáfases de algumas espécies de Nothoscordum coradas com 62 CMA (c, d), hibridizadas in situ com sonda de DNAr 5S e 45S (b, c, e) e análises meióticas de N. macrostemon (f, g) e N. nudicaule (h). (a, b) N. nudicaule 2n = 10 (NOT 104). (c) N. nudicaule 2n = 18 (NOT 1714). (d, e) N. nudicaule 2n = 19 (NOT 1717). (f) anáfase II de N. macrostemon 2n = 19 (NOT 1712). (g) N. nudicaule 2n = 19 (NOT 1717). (h) N. macrostemon 2n = 19 (NOT 1712). Em g e h, I indica univalente, II bivalente, III trivalente, IV tetravalente e VII heptavalente. Sondas marcadas em verde correspondem a sítios de DNAr 45S e em vermelho a sítios de 5S. As cabeças de seta indicam bancas CMA+. Em b “*” indica cromossomos associados pela RON. Barra em h equivale a 10 µm. Figura 3 Principais configurações meióticas observadas em tetraplóides 63 da secção Inodorum. (a, b, c, d, e, f) Nothoscordum nudicaule NOT 1717. (g, h, i, j, k, l) N. macrostemon NOT 1712. (a, g) univalentes de metacêntricos. (b) univalente de acrocêntricos. (c, h) bivalentes de metacêntricos. (d, i) bivalente de acrocêntricos. (e, j) heterotrivalentes. (f, k) tetravalentes de metacêntricos. (l) heptavalnte. Cabeças de setas indicam bancas CMA+ intersticiais. Figura 4 Idiogramas do complementos cromossômicos das amostras 64 analisadas. (a, b, c) citótipos de N. gracile NOT 1706 e 1718 (a), NOT 1716 (b) e NOT 1711 (c). (d) Nothoscordum sp. (NOT 1707). (e, f) N. macrostemon NOT 1710 (e) e NOT 1712 (f). (g, h, i) N. nudicaule NOT 104 (g), NOT 1714 e 1717 (h) e NOT 1717 (i). São apresentadas bandas CMA+ (amarelo) e sítios de DNAr 5S (vermelho) e 45S (verde). Todos os sítios de DNAr 45S são co-localizados com bandas CMA+. O ordenamento foi do maior para o menor braço curto nos metacêntricos e do maior para o menor braço longo nos acrocêntricos. Barra em (a) equivale a 10 µm. LISTA DE TABELAS 2. FUNDAMENTAÇÃO TEÓRICA Pág. Tabela 1 Números cromossômicos descritos para espécies do gênero 23 Nothoscordum. 4. MANUSCRITO A SER SUBMETIDO AO PERIÓDICO GENETICS AND MOLECULAR BIOLOGY Pág. Tabela 1 Locais de coleta, número de indivíduos analisados, números 37 cromossômicos e variação cariotípica observada em N. arenarium. 5. MANUSCRITO A SER SUBMETIDO AO PERIÓDICO ANNALS OF BOTANY Pág. Tabela 1 Local de coleta, números cromossômicos, fórmula cariotípica, 58 número de células analisadas e tamanho cromossômico médio de metacêntricos e acrocêntricos das amostras analisadas de Nothoscordum. RESUMO O gênero Nothoscordum (Alliaceae) é formado por cerca de 20 espécies nativas da América do Sul, estando dividido em duas secções, Nothoscordum e Inodorum. N. gracile (secção Inodorum) é a única espécie invasora e apresenta um cariótipo incomum (2n = 19, 13M + 6A). Para tentar entender a origem do cariótipo de N. gracile foram analisadas espécies afins da secção Inodorum, empregando a técnica de coloração com fluorocromos (CMA/DAPI) e FISH (DNAr 5S e 45S). As espécies apresentaram uma baixa quantidade de heterocromatina CMA+, distribuída no braço curto de todos os acrocêntricos e em uma ou duas bandas intersticiais de alguns destes cromossomos. Entre os diplóides, N. nudicaule foi caracterizado pela ausência de bandas CMA+ intersticiais enquanto N. macrostemon e N. arenarium apresentavam bandas intersticiais em ambos os pares de acrocêntricos. Com base nesses padrões e no heteromorfismo de bandas CMA+ encontrado em N. gracile, essa espécie pode ter se originado por alopoliploidia, possivelmente derivada do cruzamento de N. macrostemon e N. nudicaule. Análises meióticas, bem como as medições cromossômicas indicaram uma grande variabilidade cariotípica nas formas tetraplóides da secção Inodorum. Contrariamente, os citótipos diplóides apresentaram uma baixa variabilidade cariotípica. Isso foi demonstrado em uma análise populacional de N. arenarium. Esses resultados sugerem que translocações robertsonianas e alopoliploidia são os principais mecanismos envolvidos na evolução cariotípica das espécies da secção Inodorum.
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