ABSTRACT
Spores morphology and synangia in neotropical fern species of Marattia (Marattiaceae). The Marattiaceae are represented by a small family of four to six genera that bear esporogenous structures of two types: sorus with free eusporangia in Angiopteris and Archangiopteris, and indurated synangium in Christensenia, Danaea and Marattia. Marattia is a pantropical genus of about eight to ten species in the paleotropic and seven to eight species in the neotropic. In order to describe the spores and sinangia morphology, this study analyzed the shape of the receptacles, and the position of the synangia, and evaluated the spores with SEM, of seven neotropical species of the genus Marattia: M. alata, M. cicutifolia, M. excavata, M. interposita, M. laevis, M. laxa y M. weinmanniifolia from several collections. The receptacles were fully developed in M. cicutifolia and M. laevis, and scarcely overelevated in the rest of the species. The synangium was ellipsoidal and had intramarginal to supramedial position in the laminae. The spores of Marattia were elliptic. Among the taxa, only monolete spores were found, with no trilete, aborted or deformed spores. The laesura was linear and reached about two of the total length of the spore. The perispore appears as a continuous thin layer deposited on the exospore according to its ornamentation in M. cicutifolia and M. laevis. It is smooth in M. alata, rugate in M. excavata and pustulate-rugate in two species: M. interposita and M. laxa. The exospore is echinate in M. cicutifolia and M. laevis and pustulate in the other species. In M. weinmannifolia spores produced by the same sinangium may have different ornamentation types. We concluded that, while the presence of ellipsoidal and superficial synangia and monolete spores aperture were generic traits, the micro and macro-ornamentation types of the perispores and exospores vary at specific level. Besides, macro-ornamentation can be bulliform (pustulate), a combination of bulliform and muriform types (pustulate-rugate), muriform (rugate-retate) and stelliform (echinate); finally, granular micro-ornamentation can be seen frecuently in perispores. Rev. Biol. Trop. 59 (4): 1833-1844. Epub 2011 December 01.
La familia Marattiaceae, que incluye de cuatro a seis géneros, presenta estructuras esporógenas de dos tipos: sinangios en Christensenia, Danaea y Marattia; y soros con eusporangios libres en Angiopteris y Archangiopteris. Marattia es un género pantropical con unas ocho-diez especies en el paleotrópico y siete-ocho en el neotrópico. Mediante MEB se estudiaron las esporas de las siete especies neotropicales de Marattia: M. alata, M. cicutifolia, M.excavata, M. interposita, M. laevis, M. laxa y M. weinmanniifolia, basado en la forma de los receptáculos y la posición de los sinangios en material proveniente de distintas colecciones. Los receptáculos son hiperdesarrollados en M. cicutifolia y M. laevis, y apenas sobreelevados en el resto de las especies. Los sinangios son elipsoidales y la posición en la lámina es intramarginal a medial. Las esporas son elípticas, siempre monoletes y no se encuentran esporas triletes, abortadas o deformadas. La lesura tiene una longitud aproximadamente igual a la mitad del diámetro de la espora. El perisporio aparece como una capa delgada continua que se deposita siguiendo los procesos del exosporio, en M. cicutifolia y M. laevis. Es liso en M. alata, rugado en M. excavata y pustulado-rugado en dos especies: M. interposita y M. laxa. El exosporio es equinado en M. cicutifolia y M. laevis, y pustulado en las otras especies. En M. weinmanniifolia, las esporas producidas por el mismo sinangio tienen diferentes tipos de ornamentación. La presencia de sinangios elipsoidales superficiales y de esporas monoletes, son rasgos genéricos; mientras que los tipos de micro-ornamentación y macroornamentación en exosporios y perisporios son caracteres diagnósticos específicos. La macro-ornamentación puede ser buliforme (pustulada), una combinación de buliforme y muriforme (pustulada-rugada), muriforme (rugada-retiada) y esteliforme (equinada); es frecuente la presencia de micro-ornamentación granular en el perisporio.
Subject(s)
Ferns/cytology , Spores/cytology , Costa Rica , Ferns/classification , Spores/ultrastructureABSTRACT
The spore morphology and wall ultrastructure of Sphaeropteris gardneri (Hook.) R.M. Tryon from Brazil were analyzed with LM, SEM and TEM. The spores are trilete with an ornamentation formed of short low ridges with spines in their margins. The exospore is 2.5μm thick, two- layered in section and single or branched channels are present. The perispore is 1.2μm thick and two-layered. The inner layer has three strata: the inner stratum is formed of a network of branched and fused threads, the middle stratum has threads with a radial orientation and in the outer stratum thin, dark fibres are immersed in a less dense contrasted matrix. The outer layer of the perispore is the one that forms the echinate-ridges and is constituted of threads arranged in a compact way. Globules of different sizes are observed on the surface. The differences found in the perispore ornamentation and ultrastructure in Alsophila, which was previously studied, and those of Sphaeropteris, show a tendency to wall complexity.
A morfologia dos esporos e a ultraestrutura da parede de Sphaeropteris gardneri (Hook.) R.M. Tryon, Brasil, foram analisadas com MO, MEV e MET. Os esporos são trilete com uma ornamentação formada por cristas curtas e baixas e com espinhos em suas margens. O exosporo possui 2,5μm de espessura, duas camadas em secção e estão presentes canais simples ou ramificados. A camada interna possui três estratos: o estrato interno é formado por uma rede de filamentos ramificados e fusionados, o estrato médio tem fios com uma orientação radial e no estrato externo fino, fibras escuras estão imersas em uma matrix menos densa. A outra camada do perisporo é a que forma as cristas equinatas e é constituída de filamentos dispostos em um arranjo compacto. Glóbulos de diferentes tamanhos são observados na superfície. As diferenças encontradas na ornamentação do perisporo e na ultraestrutura do Alsophila estudado previamente e aqueles de Sphaeropteris mostram uma tendência à complexidade da parede.
Subject(s)
Ferns/cytology , Spores/ultrastructure , Microscopy, Electron/methodsABSTRACT
The fern Blechnum sprucei grows in Mesoamerica (Costa Rica) and South America, from Colombia to Bolivia, SE and centre of Brazil, Paraguay and Argentina. It is a distinctive, somewhat vulnerable, mostly orophilous species. Fresh and dry herbarium material was used for this study. Herbarium material for anatomical studies comes from CTES, BA, LP, MA, SI and UC (Holmgren et al.1990). Selected representative specimens are additionally cited after taxonomic treatment of the species. Dry material was restored with aqueous 4:1 butil cellosolve. Pinnae were cleared with aqueous 6% NaOH, then coloured with aqueous 1 % TBO (Gurr 1966). Hand made transverse sections of young and adult stipes, and costae were done in fresh and restored herbarium material. Venation and epidermal patterns were analyzed in basal, apical and medium pinnae, but only the latter were illustrated. The size and density of stomata were measured in medium pinnae from all studied samples, values shown are the average of 25 measures per sample; sizes are expressed as minimum, media and maximum length x width, in µm, and density as minimum, media and maximum number of stomata / mm². Spores were studied with SEM, mounted on metal stubs with double sided tape, covered with gold under vacuum and photographed with a Jeol /EO JSM 6360 (15 KV) SEM. Spores were also studied with light microscope, mounted in DePex (DePex mounting medium, Gurr, BDH Laboratory Supplies, Poole BH15 1TD, UK) and measured using an ocular micrometer. Measurements are based on a minimum sample of 100 spores taken from different specimens. Sizes are expressed as the longest equatorial diameter/ polar diameter, in µm. Gametophytes were studied from material collected in the subtropical forest of Tucumán Province, Argentina. Spore samples for cultures were taken from single sporophytes kept dry at room temperature since the date plants were collected. Gametophytes were grown under fluorescent light. Multispore cultures were established on mineral agar. Percentage of germination was recorded for a random sample of 50 spores from each of the two plates, every three days until there was no further increasing. Gametophytes were stained with chloral hydrate acetocarmine. The species has large sporophytes, suberect, scaly rhizomes, and dimorphic fronds with short, scaly stipes and lanceolate to elliptic sterile laminae. The rachises can grow indefinitely as radicant axis that vegetatively multi-plicate the plants. Pinnae are lanceolate, herbaceous, with crenate and papillose margins, superficially scaly and hairy, peciolulate, with free, visible veins regularly once furcated near the costa, ending in large, active hydathodes. The broadly elliptic fertile laminae bear distant pinnae, with vegetative tissue reduced to the portion that supports the indusium and the continuous coenosorus; terminal indefinite rachis, not proliferous, may be present. Rev. Biol. Trop. 56 (4): 2027-2040. Epub 2008 December 12.
El helecho Blechnum sprucei crece en Mesoamérica (Costa Rica) y Sudamérica, desde Colombia a Bolivia, SE y centro de Brasil, Paraguay y Argentina. Es una especie característica, algo vulnerable y orófila. Se estudiaron caracteres exomorfológicos de especimenes provenientes de distintos puntos de su gran área de distribución. Adicionalmente, se llevó a cabo un detallado análisis de los modelos epidérmicos jóvenes y maduros, del indumento de la lámina y los ejes, y de la organización vascular en los estipes y costas. Se han estudiado por primera vez la morfología esporal, el desarrollo de los gametófitos, que resultaron cordados y pelosos, y su expresión sexual. Presenta esporófitos grandes, suberectos, con rizomas escamosos y frondas dimórficas, con estipes cortos y escamosos y láminas estériles de lanceoladas a elípticas. El raquis puede crecer indefinidamente como un eje radicante que multiplica las plantas vegetativamente. Las pinnas son lanceoladas, herbáceas, con márgenes crenados y papilosos, y la superficie escamosa y pelosa. Son pecioluladas, con venas libres y visibles, regularmente furcadas cerca de la costa, terminando en hidatodos grandes y activos. Las láminas fértiles son anchamente elípticas y portan pinnas distales; presentan tejido vegetativo reducido a la porción que soporta el indusio y el cenosoro continuo. Pueden presentar raquis indefinidos, pero no prolíferos.