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Modelo De Artigo Para Publicao Na Revista Principia Do CORE Metadata, citation and similar papers at core.ac.uk Provided by Universidade Federal do Maranhão (UFMA): Portal de Periódicos Revista Tropica: Ciências Agrárias e Biológicas, v.7,n.1- 2013 Centro de Ciências Agrarias e Ambientais-Universidade Federal do Maranhão, MA. ISSN 1982-4831 Embriogênese adventícia em Eugenia uniflora L. (Myrtaceae). André Luiz Silva1, Bruno Cardoso Lopes2, Lygia Doleres Ribeiro de Santiago Fernandes2 Resumo – O óvulo de Eugenia uniflora é hemicampilótropo, unitegmentado, crassinucelado, com obturador tegumentar. O óvulo é vascularizado por um feixe que penetra no funículo e sofre bifurcação formando dois ramos, um deles estende-se até a calaza enquanto o outro segue até as proximidades da micrópila. O nucelo tem o ápice estendido formando uma caliptro nucelor estreita e pontiaguda. O gametófito maduro de Eugenia uniflora enquadra-se no tipo Polygonum. A embriogênese tem início no período compreendido entre 48 e 72 horas pós-polinização. A partir do terceiro dia foram observados dois tipos de desenvolvimento seminal em diferentes óvulos. Em certas sementes o processo é convencional, com a presença de um embrião globular, endosperma nuclear volumoso e nucelo com células vacuoladas e citoplasma denso no quarto dia pós-polinização. No outro tipo de desenvolvimento, foram encontradas estruturas volumosas de contornos circulares, pluricelulares, com protoderme e circundadas pelo endosperma em sementes de sete dias pós-polinização. Estas estruturas foram interpretadas como embriões de origem nucelor. O desenvolvimento destes embriões adventícios é muito veloz e dependente da fecundação o que sugere a ocorrência de competição por recursos maternos. Esta hipótese, caso confirmada, poderia explicar a baixa relação semente/óvulo em Eugenia uniflora. Adventitious embryogenesis in Eugenia uniflora L. (Myrtaceae). Abstract-The ovule of Eugenia uniflora is hemicampylotropous, unitegmic, crassinucellate with obturator tegumentary obturador. Vascularization of the ovule is formed by a bundle which penetrates the funiculus and suffers split forming two branches, one of which extends to the chalaza while the other goes to the vicinity of the micropyle. The nucellus has the extended apex forming a nucellar calyptra narrow and pointed. The mature gametophyte of Eugenia uniflora is the Polygonum type. The Embryogenesis begins in the period between 48 and 72 hours post-pollination. From the third day was observed two kinds of developing seminal in different ovules. In certain seeds the process is conventional, with a globular embryo, endosperm nuclear bulky, nucellus with vacuolated cells and dense cytoplasm on the fourth day after pollination. In other development type, were found bulky structures of circular contours, multicellular, with protoderm and surrounded by the endosperm in seeds of seven days post-pollination. These structures were interpreted as nucellar embryos. The development of these adventitious embryos is very fast and dependent fertilization suggesting the occurrence of competition for maternal resources. This hypothesis, if confirmed, could explain the low ratio seed / ovule in Eugenia uniflora. INTRODUÇÃO O estudo acerca da biologia muitas espécies não aparentadas reprodutiva de Eugenia uniflora (Mohana et al. , 2001). Características evidenciou um crescimento diferencial hereditárias ocasionadas por forças entre as sementes de um mesmo fruto e, seletivas ao longo da evolução podem na maturidade, uma baixa razão ser responsáveis pela baixa razão S/O semente/óvulo (S/O) (Silva & Pinheiro, em muitas espécies (Casper & Wiens, 2009). Este fato é característico de 1981, Charlesworth, 1989). Esse várias espécies de Eugenia (Wyk & fenômeno pode também ser resultado de Lowrey, 1988, Beardsell et al ., 1993a, uma competição seletiva entre os b; Proença & Gibbs, 1994; Lughadha & embriões em desenvolvimento por Proença, 1996) assim como de outros recursos maternos (Aratthi et al ., 1996). gêneros de Myrtaceae (Arathi et al., A redução do número de sementes 1996; Schimidt-Adam et al. , 2000) e de por fruto pode ser uma vantagem para 1Centro de Ciências Agrárias e Ambientais – Universidade Federal do Maranhão. E-mail para correspondência: [email protected] 2Departamento de Botânica – Museu Nacional – Universidade Federal do Rio de Janeiro Revista Trópica – Ciências Agrárias e Biológicas V. 7, N.1, pág. 97, 2013 SILVA André Luiz Gomes da et al espécies que investem em recursos para Syzygium. Tiwari (1926 apud Pijl, estabelecimento das plântulas em 1934), analisando Eugenia cumini (L.) ambientes com pouca concentração de Druce. (= S. cumini (L.) Skeels) nutrientes no solo (Casper & Wiens l.c verificou pela primeira vez a ocorrência .; Sedgley, 1989; Charlesworth l.c .; de poliembrionia no grupo. Dalling & Hubbell, 2002). Por outro Posteriormente, Pijl (1934) registrou lado, caso haja a perda de uma semente poliembrionia em Eugenia malaccensis nestas condições, por aborto ou L., (= S. malaccense (L.) Merr. & L.M. predação, isso pode significar uma Perry), E. aquea Burm. (= S. aqueum perda considerável do investimento (Burm.) F. Alston) e E. jambos L. (= S. energético feito em uma flor (Mitchel, jambos (L.) Alston), nesta última com 1977; Ramirez 1992). Não há para início do desenvolvimento anterior à Eugenia, no entanto, trabalhos fertilização. Johnson (1936, apud anatômicos referentes ao Johansen, 1950) registrou “dois a vinte desenvolvimento comparado das embriões de origem presumivelmente sementes de um mesmo fruto, de modo nucelor” em E. hookeri. a permitir o registro da etapa em que se Outros trabalhos sobre tecnologia de dá a divergência no desenvolvimento, sementes e biologia reprodutiva também definindo quais sementes chegarão à relatam sementes poliembriônicas, em maturidade e quais terão seu algumas espécies de Myrtaceae (Silva et desenvolvimento abortado. Roy al., 2003; Lughadha & Proença, 1996). (1953,1955,1960,1961,1962a, b) Neste artigo analisou-se o descreveu as características do óvulo e desenvolvimento das sementes de do saco embrionário e registrou Eugenia uniflora, com o intuito de sucintamente variações nas mesmas em fornecer subsídios para a compreensão Eugenia jambos L., E. bracteata Roxb., do fenômeno da baixa produção de E. malaccensis L., E. fruticosa L. e E. sementes e verificar a possibilidade da myrtifolia Roxb, respectivamente. Em ocorrência de embriogênese adventícia E. bracteata e E. malaccensis , registrou durante as duas primeiras semanas pós- a existência de sacos embrionários polinização. anormais, os quais considerou como possivelmente apospóricos. Descreveu MATERIAL E MÉTODOS os óvulos de E. bracteata como bitegumentados e os de E. malaccensis Foram realizadas polinizações como unitegumentados. Entretanto, em xenogâmicas manuais em flores razão de alterações propostas por ensacadas um dia antes do início de sua Schmid (1972) atualmente quatro destas antese na restinga de Grumari, Rio de espécies estão incluídas em Syzygium Janeiro – RJ, Brasil e em plantas como S. jambos (L.) Alston, S. cultivadas no Horto Botânico do Museu malaccense (L.) Merr. & L.M. Perry. S. Nacional/UFRJ, São Cristóvão, Rio de fruticosum (Roxb.) DC., S. myrtifolium Janeiro – RJ, Brasil. As flores foram (Roxb.) DC. Miq.. Eugenia bracteata marcadas e ovários e frutos foram Rich. foi transferida para o gênero coletados em vários estágios do Myrcia como M. bracteata (Rich.) DC. desenvolvimento, desde a pré-antese até 14 dias após a polinização. Testes de A literatura registra, ainda a apomixia foram realizados retirando-se ocorrência de poliembrionia e de o estilete de algumas flores previamente embriões nucelores em algumas ensacadas na fase de botão (Richards, espécies também identificadas à época 1986) e marcadas para posterior das publicações como pertencentes à observação e coleta em plantas Eugenia e atualmente incluídas em Revista Trópica – Ciências Agrárias e Biológicas V. 7, N.1, pág. 98, 2013 SILVA André Luiz Gomes da et al cultivadas no Horto Botânico do Museu cujos óvulos diferem por serem citados Nacional/UFRJ. como anátropos e bitegumentados Alguns frutos coletados foram (Davis, 1966 apud Schmid, 1972). No dissecados em diferentes fases para a entanto, como não tivemos acesso ao observação do desenvolvimento trabalho de Davis (loc. cit.), não seminal. Os ovários e frutos restantes dispomos de informações sobre quais foram fixados em formaldeído 4% e espécies foram analisadas, havendo glutaraldeído 2,5% em tampão fosfato inclusive a possibilidade de pertencerem de sódio 50mM pH 7,2, desidratados em atualmente a outros gêneros, face às série alcoólica e emblocados em grandes revisões realizadas na Historesina (Leica®) (Feder & O'Brien, taxonomia do grupo neste período. 1968). Secções de 1-3 μm de espessura Syzygium, cujas quatro espécies foram obtidas com navalha de vidro em estudadas apresentam óvulos micrótomo rotativo Spencer e coradas unitegumentados (Roy, 1953, com Azul de Toluidina. Secções 1960,1961,1962) e pertenciam a seriadas dos óvulos foram observadas e Eugenia s.l. à época da publicação do fotografadas em microscópio Orthoplan trabalho de Davis (Davis, 1966 apud – Leitz. Schmid, 1972), é a única exceção conhecida para toda a família. Diante da RESULTADOS E DISCUSSÃO escassez de estudos embriológicos em Myrtaceae, Tobe & Raven (1983) já O óvulo de Eugenia uniflora é consideravam a informação hemicampilótropo, unitegumentado, embriológica disponível baseada em crassinucelodo, com obturador informações inadequadas e que tegumentar (figura 1A, B e 2). Estas provavelmente a unitegmia
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