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Estudios Embriológicos En Luma Apiculata Introducción

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Estudios Embriológicos En Luma Apiculata Introducción ISSN 0373-580 X Bol. Soc. Argent. Bot. 38 (3-4): 265 - 275. 2003 ESTUDIOS EMBRIOLóGICOS EN LUMA APICULATA (MYRTACEAE) A. FAIGÓN SOVERNA1, S. FRONTERA1y B. G. GALATI1 Summary: Embryological studies on Luma apiculata (Myrtaceae). The development of the sporangia, sporogenesis, and gametogenesis of Luma apiculata was studied. The anther wall comprises epider¬ mis, endothecium, two middle layers, and tapetum. The tapetum is of the secretory type and has binucleate cells. A tapetai membrane with Ubisch bodies is formed and persists until the opening of the anther. The microspore tetrads are tetrahedral, the pollen grains are tricolporate and are shed at 2-celled stage. The stigma is constituted by 1-celled papillae and the style is solid. The ovule is anacampylotropus, crassinucellate and bitegmic. Both integuments form a zig-zag micropyle. Megaspore tetrads are linear and the chalazal one develops into a Polygonum type of embryo sac. No abnormalities suggestive of any type of sterility or impediments to the sexual reproduction of L. apiculata have been observed during sporogenesis and gametogenesis. Key words: pollen, Ubisch bodies, megagametophyte. Resumen: En este trabajo se estudia la esporogénesis y la gametogénesis de Luma apiculata. La pared de la antera comprende epidermis, endotecio, dos capas medias y tapete. El tapete es de tipo secretor y sus células son binucleadas. Una membrana tapetai con cuerpos de Ubisch persiste hasta la apertura de la antera. Las tétrades de micrósporas son tetraédricas. Ei grano de polen es tricolporado, y se dispersa en estado bicelulár. El estigma está formado por papilas unicelulares y el estilo es de tipo sólido. Los óvulos son crasinucelados, anacampilótropos, bitegumentados, con micrópilo en zig-zag. La tétrade de megásporas tiene un arreglo lineal. La megáspora calazal da origen a un S E. tipo Polygonum. No se detectó ninguna anormalidad en los procesos de esporogénesis y gametogénesis que pueda sugerir algún tipo de esterilidad o impedimento para la reproducción sexual de L. apiculata. Palabras clave: polen, cuerpos de Ubisch, megagametófito. INTRODUCCIóN Algunos estudios hanrevelado anormalidades como: sacos embrionarios incompletos o no organizados, Luma apiculata (DC.) Burret es una especie óvulos estériles sin saco embrionario, células del arbórea a subarbórea, nativa del sur argentino que megagametófito con polaridad invertida y núcleos suele encontrarse formando bosques puros bastante polares supernumerarios (Roy, 1962; Prakash, 1973) extensos. Vulgarmente conocida como “arrayán” atrae El conocimiento de la biología reproductiva de especialmente la atenciónpor elcolor canela y lalisu¬ las especies vegetales es de gran utilidad e impor¬ ra de su corteza. tancia en la implementación de planes de manejo y Los trabajos embriológicos realizados hasta el conservación. Por esta razón, y debido a lo poco momento en la familia Myrtaceae son escasos y res¬ • que se conoce sobre este aspecto en L. apiculata, tringidos en su mayoría al género Eucalyptus y a se decidió encarar en esta especie un estudio em¬ otras especies paleotropicales (Johri et al. 1992). briológico que abarque los aspectos relativos a la pre- y post-fertilización. En esta primera etapa se 1 Laboratorio de Biología Reproductiva en Plantas ha realizado el análisis detallado de la Vasculares, Depto. Biodiversidad y Biología Experimental, esporogénesis, la gametogénesis y la ontogenia FCEyN, UBA, [email protected] de las estructuras esporofíticas relacionadas. Que- 265 Bol. Soc. Argent. Bot. 38 (3-4) 2003 da para una segunda etapa el estudio de la forma- ultrafinos para su observación al MET. A tal fin se ción de la Semilla y el fruto. contrastaronconacetato de uranilo y citrato deplomo (Reynolds) (O’Brien&Me Cully, 1981). MATERIALES Y MéTODOS RESULTADOS Materialestudiado:ARGENTINA.Prov.Neuquén: Depto. Los Lagos, Villa La Angostura, Galati 380 Microsporangio (BAFC). La antera de L. apiculata es tetrasporangiada y Estudios al microscopio óptico (MO) condehiscencialongitudinal.Eltejido conectivo está bien desarrollado yposee células tánicas e idioblastos Los materiales fueron fijados en F.A.A. (formol, con drusas (Fig. 1 F). La pared de la antera joven alcohol y ácido acético). Capullos en distintos esta- consta de afuera hacia adentro de: epidermis, dios de desarrollo y flores en antesis se endotecio, 1 a 2 capas medias y tapete de tipo secre- deshidratararon en una serie ascendente de alcoho- tor (Fig. 1 A, B; 2 A-C). El origen de la pared les (alcohol etílico-xilol) y se incluyeronenparafina, corresponde al tipo básico (Davis, 1966), es decir, Se realizaron cortes micrométricos de 5 a 10 pm de una capa media tiene origen común con el tapete y espesor con micrótomo rotativo tipo Minot, los cua- otra con el endotecio. les fueron teñidos según la doble coloración En los primordios florales pequeños, las células Safranina-Fast Green (D’Ambrogio, 1986) y monta- que componen los 4 estratos de la pared tienen m dos luego en resina sintética. tamaño y aspecto semejantes; luego durante el Cortes micrométricos de pimpollos jóvenes se desarrollo van sufriendo transformaciones que le desparafinaron, se colorearonconuna solución acuo- confieren a cada estrato características particulares. samuydiluida de azul de algodón(0,05%) y se obser¬ Las células tapétales sonlasprimeras enaumentar varon euun microscopio de epifluorescencia para la de tamaño; son binucleadas y poseen un citoplasma detección de calosa(O’Brien&McCully, 1981;Rost, microvacuolado(Fig. 2 A). Cuando las células madres 1992). Se comprobó lapresencia de oxalato de calcio de los micrósporos (CMm) se rodean de calosa, el usando una solución acuosa de acetato cúprico a citoplasma de las células tapétales comienza a saturación(D’Ambrogio, 1986). consumirse en forma gradual, susnúcleos a obliterarse y al momento de la reducción meiótica las paredes Estudios al microscopio electrónico de barrido tangencial externa y radiales ya se han degradado (MEB) completamente (Fig. 1 B-C; 2 B, C, E). Comienza entonces a formarse una membrana tapetai con Delmaterial fijado enF.A.A. se separaron anteras; cuerpos de Ubisch (C U),la cual formauna suerte de se deshidrataron en una serie ascendente de bolsa que encierra a los granos depolen(Fig. 2F,G). acetonas, luego se secaron por medio de la técnica Las orbículas o CUson esféricos a sub-esféricos y en depunto crítico.Finalmente se montaronymetalizaron corte se observan sólidos, sinun corazón central y con la con oro-paladio para su observación al MEB. mismaelectrodensidadquelaexina(Fig.3 A;4C).Porotra parte reaccionan con los colorantes de la misma manera Estudios al microscopio electrónico de transmisión 9ue Ia esporopolenina de los granos de polen. (MET) Tanto las células de la epidermis como las del endotecio y de las capas medias crecen radial y Anteras maduras fueron fijadas en glutaraldehído tangéncialmente amedida quela antera vamadurando 3% en buffer fosfato pH 7,6 y refijadas en tetróxido (Fig. 2 B, C,E). Las células epidérmicas desarrollan de osmio 1,5% enelmismobuffer y a igualpH.Luego una gruesa cutícula estriada en los últimos estadios fueron deshidratadas en una serie ascendente de del desarrollo (Fig. 2F,G). alcoholes para su posterior inclusión en resinas ep¬ Durante la microgametogénesis las células del oxy. Se realizaron cortes de 1 pm de espesor con endotecio forman en sus paredes tangenciales ultramicrótomo los cuales fueron coloreados conazul internas y radiales engrasamientos lignificados en de toluidina para su observación al MO y cortes forma de bandas de naturaleza secundaria que se 266 A. Faigón Soverna et al., estudios embriológicos en Luma apiculata (Myrtaceae) r - v . 4 ' m /ÿ f - -r * i4 4>c. / B C r 7 M —i* ’ *2;*1*1 t V - m i I i ¡ •• - i «¡jala*».-t I/?, . s JE - •ÿ * <s-, mIB. ara ? VÍy'i *P5f«p aMÿ: - * m ÉÜ I I , MSKJBSVJKÍJé D E í - ’ .;T p*m ** r-r¿i > .vM Ü ... — /- m / w # Ljf e. 1 *1 í*; 17/V !--.‘ *%ÿ V.. nr 5. m , FA **jpgí *dp €. L ST • “- |¥ % 1/1 ' »’*» *• « 1 í># V:A *f kI »ÿaL_\ .4':‘ - \¿S~" J¿s» 4» !ÿ«r -Jé-ce Ra* H Fig. 1. A- H: Fotomicrografías al microscopio óptico. A: Lóculo del microspórangio con tejido arquespórico én sección transversal; B: Célula madre de los micrósporos con pared de calosa (c) reffingente; C: Reducción meiótica en célula madre de la micróspora; D: Tétrades tetraédricas de micrósporos rodeadas por calosa; E: Micrósporos recién liberados de la tétrade con restos decalosa; F: Antera madura en sección transversal , tejido conectivo con drusas (d) y células tánicas; G: Célula madre de la megáspora (cmm) situada profundamente en la núcela; H: Tétrade lineal de.megásporas, calosa en paredes transversales que separan a las megásporas (flechas). Escalas:A-E; G-H = 13 mm; F= 500 qm. 267 Bol. Soc. Argent. Bot. 38 (3-4) 2003 / 'y' 'Jg? 'él © (i O % Y/ ,0/ m. fí 'A x QmÊM e. © 15 ® A Ib ñiQmÊÊÿ?Ê QNj I RJ \ c n ;•S§s3- oWm\ \B» j fe-’ 0\%2 23 © 2 í© 12<k>, D B 7# 0/ / ®/W . e <ÿ S)/ •• # ® : © i.vV; .• el le ,-«5 u * i© ,-• © iv.f'í E r\ ,ot= t í ©/,TT ai 2 •3.: t 7 Li / 0íí i y t -7 i l. Q> * ©f, 1 t F O G Fig. 2. A-D: Esquemas de la microesporogénesis. E-G: Microgametogenesis. A: Detalle de corte transversal de antera joven con tejido arquespórico, células tapétales uninucleadas; B: Célula madre de la micróspora con pared de calosa, células tapétales bi-nucleadas; C: Reducción meiótica en célula madre de los micrósporos, las paredes radiales de las células tapétales se han degradado; D: Tétrades tetraédricas de micrósporos rodeadas por calosa; E: Micrósporos libres; F: Micrósporos vacuolados y con exina en formación, pared tangencial interna de las células tapétales con cuerpos de Ubisch, los estratos parietales se han consumido y la cutícula de la epidermis presenta estriaciones; G: Granos de polen bicelulares, endotecio con engrasamientos fibrosos o en banda.
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