Ontogênese De Órgãos Reprodutores

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Ontogênese De Órgãos Reprodutores MINISTÉRIO DO MEIO AMBIENTE INSTITUTO DE PESQUISAS JARDIM BOTÂNICO DO RIO DE JANEIRO DIRETORIA DE PESQUISAS ONTOGÊNESE DE ÓRGÃOS REPRODUTORES Rio de Janeiro, junho de 2017. ONTOGÊNESE DE ÓRGÃOS REPRODUTORES COORDENADOR Karen L. G. De Toni (Instituto de Pesquisas Jardim Botânico do Rio de Janeiro) LOCAIS DE EXECUÇÃO DO PROJETO Instituto de Pesquisas do Jardim Botânico do Rio de Janeiro - Laboratório de Botânica Estrutural. Avenida Pacheco Leão, nº 915. Rio de Janeiro / RJ. APRESENTAÇÃO O estudo da “Ontogênese de Órgãos Reprodutores” é um projeto amplo, dentro do grupo de pesquisa “A anatomia vegetal no conhecimento da diversidade biologia: buscando compreender as estratégias de sobrevivência vegetais” do Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. Este projeto visa analisar, dentro de estudos de caso, a ontogênese dos órgãos reprodutores de plantas vasculares. A ontogenia é o estudo do desenvolvimento dos seres vivos, tanto animais como vegetais, a partir da célula ovo até sua maturação (Font Quer 1985). A análise da formação de parte ou de todo o organismo é analisada (Raven et al. 1999), sendo resultado da ação de fatores externos em resposta a fatores internos do organismo, que serão transmitidos para seus descendentes (Turrill 1942), evidenciando a rápida e curta recapitulação da história evolutiva da espécie (Haeckel 1866). Atualmente, a ontogenia é também utilizada com o intuito de esclarecer problemas filogenéticos e taxonômicos entre os taxa. De acordo com Nelson (1973; 1978) a ontogenia é o único recurso direto para determinação de caracteres filogenéticos. Porém, com o avanço da biologia molecular, nos últimos anos, têm-se utilizado, além da ontogenia dos órgãos vegetativos e reprodutivos, dados macromoleculares (DNA e/ou RNA) para caracterizar os taxa. O estudo da ontogenia floral tem destacado diversos aspectos do desenvolvimento, simetria, heterocronia, ordem de iniciação dos órgãos e casos de fusão, proliferação, supressão ou perda de órgãos (Tucker 1987, 1989, 1992, 1997, 1999). A ontogênese da flor envolve desde a iniciação do primórdio floral até sua antese, revelando, por exemplo, na família Leguminosae, diversos caracteres no estádio inicial de organogênese, de grande importância para a compreensão da evolução nesta família e, conseqüentemente, para a sistemática (Tucker 1984, 1989, 1992, 1997, 1999, Tucker & Douglas 1994). Partindo deste princípio, Mansano et al. (2002) descrevem e comparam a ontogênese floral de Lecointea Ducke, Zollernia Maximil. & Nees, Exostyles Schott ex Ontogênese dos Órgãos Reprodutores Spreng. e Harleyodendron R.S.Cowan (Leguminosae – Papilionoideae – Swartzieae), e concluem que as espécies analisadas, embora incluídas na tribo Swartzieae, mais se aproximam da tribo Sophoreae, devido à ausência do meristema em anel dos taxa analisados, estrutura característica de Swartzieae. A descrição dos caracteres ontogenéticos baseia-se na morfologia do meristema floral e dos primórdios dos órgãos, no momento em que essa morfologia se expressa. Também faz parte do estudo do desenvolvimento floral a formação dos esporângios e eventos da esporogênese e gametogênese, sendo os últimos também incluídos na área da embriologia (Maheshwari 1950). As obras clássicas de Johansen (1950), Maheshwari (1950), Davis (1966), Johri (1984) e Johri et al. (1992) confirmam a importância do uso dos caracteres embriológicos na sistemática das angiospermas. Os caracteres embriológicos, aliados ao conjunto de informações de outras áreas de estudo, subsidiam uma melhor avaliação das relações entre os taxa (Herr 1984). Segundo esse autor, os caracteres mais utilizados e freqüentemente aplicados à sistemática são: − antera: número de esporângios, origem das camadas parietais, delimitação dos esporos por divisão simultânea ou sucessiva e forma da tétrade de esporos; − andrófito (grão de pólen): número e posição de suas aberturas, ornamentação da exina, localização da célula generativa e número de células no grão de pólen; − rudimento seminal: características de seu desenvolvimento e maturação, número e origem dos tegumentos, origem do nucelo, vascularização, presença ou ausência de endotélio, hipóstase, epistase, arilo, obturador, carúncula e estrofíolo; − ginosporogênese: número de células arquesporiais e seu posicionamento no nucelo, alterações no esporo para a realização da meiose (movimento nuclear, grau de vacuolação, alteração de tamanho e forma) e posição do ginósporo funcional; − ginófito: padrão de desenvolvimento (monospórico, bispórico e tetraspórico), diferenciação do aparelho antipodal, estrutura das antípodas, sinérgides e oosfera, e posição dos núcleos polares; − fecundação: entrada do tubo polínico, intervalo entre a polinização e a fecundação; − endosperma: padrão de desenvolvimento, posição dos núcleos durante estádios precoces e orientação das paredes celulares, origem do haustório (quando presente), tipo e quantidade de reserva, persistência na semente madura; − embriogênese: tipo de desenvolvimento, forma do embrião maduro, suspensor; − testa da semente: número de camadas celulares e sua derivação (a partir de um ou dois tegumentos), padrão de diferenciação celular. 2 Ontogênese dos Órgãos Reprodutores Alguns trabalhos exemplificam a utilização da ontogenia dos órgãos reprodutores com ênfase à sistemática. Entre estes, os estudos realizados na família Podostemaceae que, apesar de apresentar caracteres morfológicos, anatômicos e ecológicos exclusivos, possui grande afinidade com as famílias Hydrostachyaceae, Saxifragaceae e Crassulaceae (Hutchinson 1973, Engler 1930, Rendle 1952, Mauritzon 1939). Após a investigação embriológica da família Podostemaceae, com o objetivo de acrescentar dados à discussão filogenética (Battaglia 1971, Nagendram 1974, Arekal & Nagendram 1975), uma estreita relação com a família Crassulaceae foi constatada, devido à ampla similaridade de caracteres encontrados, principalmente com relação à ontogênese do rudimento seminal (nucelo) e do ginófito (Herr 1984). A ontogênese dos órgãos reprodutores auxilia, também, no posicionamento de taxa, como no caso do gênero Nyctanthes L., família Oleaceae. Atualmente a maioria dos taxonomistas inclui o gênero Nyctanthes na família Oleaceae. Porém Airy Shaw (1952) baseando-se na natureza quadrangular de seu caule, transferiu Nyctanthes para a família Verbenaceae, posição mantida por Stant (1952) e Hutchinson (1973). Kapil & Vani (1966) analisaram as características embriológicas e morfológicas de Nyctanthes e as comparou com as de alguns gêneros de Oleaceae e Verbenaceae. Estes autores concluíram que Nyctanthes assemelha-se mais a Oleaceae do que a Verbenaceae, principalmente com relação à quantidade de estames, morfologia do grão de pólen, ornamentação da exina, número de células arquesporiais, padrão de formação do ginófito, antípodas efêmeras, ausência de haustório no endosperma e tipo de fruto. De modo geral, considerando os diversos enfoques direcionados à ontogenia, há, ainda, a necessidade de estudos ontogenéticos em muitos grupos vegetais, visando o enriquecimento da sistemática e da filogenia. Diversos caracteres morfológicos dos carpelos, rudimentos seminais e particularidades dos eventos da ginosporogênese são amplamente utilizados na sistemática. Bocquet (1959) e Bocquet & Bersier (1960) foram os primeiros a ressaltarem a importância da ontogênese do rudimento seminal. Entretanto, foram os estudos de Bouman (1971, 1974, 1984), sobre a ontogênese dos rudimentos seminais e de seus tegumentos, que inspiraram hipóteses sobre a filogenia das angiospermas. O processo de redução nucelar pode ser observado em muitas famílias de angiospermas simpétalas. O nucelo tende a tornar-se reduzido e a apresentar somente uma célula arquesporial, coberta por algumas células epidérmicas (Bhandari et al. 1976). Este processo de redução do nucelo é registrado para Loranthaceae (Cocucci & Venturelli 1982) e Rubiaceae (Fagerlind 1937, Andronova 1977, Galati 1991, Mariath & Cocucci 1997, De Toni & Mariath 2004), entre outras famílias. Estudos ontogenéticos do rudimento seminal de diversas espécies da família Rubiaceae sugerem que, além da redução do nucelo, outras características são importantes para a filogenia da família, como o desaparecimento do tegumento externo, inclusão do nucelo na calaza, achatamento da superfície da epiderme nucelar, redução e alongamento de suas células, e diminuição no número 3 Ontogênese dos Órgãos Reprodutores de células arquesporiais. A partir desses caracteres, diagramas evolutivos foram elaborados, pelos autores anteriormente citados, para a família. Ao comparar os diagramas resultantes das análises ontogenéticas com os resultados de estudos moleculares, observa-se uma uniformidade de resultados, ou seja, os resultados morfológicos corroboram com os moleculares, e confirmam a proposta de Bremer (1996), que indica a alteração das subfamílias de Rubiaceae, discordando de Robbrecht (1988). Além dos exemplos citados, outros caracteres relacionados aos carpelos podem ser considerados na sistemática e filogenia das espécies, como, por exemplo, a redução no número de carpelos, o estabelecimento da posição do ovário (ínfera, súpera ou semi-ínfera), a ginosporogênese e a ginogametogênese. Com relação ao androceu, sua ontogênese tem-se mostrado importante na delimitação de subclasses e ordens das angiospermas (Cronquist 1988). Os primórdios estaminais em flores com androceu polistêmone podem surgir em seqüência centrípeta ou centrífuga. Representantes das subclasses Magnoliidae, com flores polistêmones, possuem
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