Non-chlorophyllous and crypto-chlorophyllous fern spores differ in their mobilisation of fatty acids during priming.

During fern spore germination, lipid hydrolysis primarily provides the energy to activate their metabolism. In this research, fatty acids (linoleic, oleic, palmitic and stearic) were quantified in the spores exposed or not to priming (hydration-dehydration treatments). Five fern species were investigated, two from xerophilous shrubland and three from a cloud forest. We hypothesised that during the priming hydration phase, the fatty acids profile would change in concentration, depending on the spore type (non-chlorophyllous and crypto-chlorophyllous). The fatty acid concentration was determined by gas chromatograph-mass spectrometer. Chlorophyll in spores was vizualised by epifluorescence microscopy and quantified by high-resolution liquid chromatography with a DAD-UV/Vis detector. Considering all five species and all the treatments, the oleic acid was the most catabolised. After priming, we identified two patterns in the fatty acid metabolism: (1) in non-chlorophyllous species, oleic, palmitic, and linoleic acids were catabolised during imbibition and (2) in crypto-chlorophyllous species, these fatty acids increased in concentration. These patterns suggest that crypto-chlorophyllous spores with homoiochlorophylly (chlorophyll retained after drying) might not require the assembly of new photosynthetic apparatus during dark imbibition. Thus, these spores might require less energy from pre-existing lipids and less fatty acids as 'building blocks' for cell membranes than non-chlorophyllous spores, which require de novo synthesis and structuring of the photosynthetic apparatus.

Can fern spores develop hydration memory in response to priming?

Fern spores and seeds initiate germination with fast water uptake, followed by a stationary phase with no appreciable water uptake and biochemical and metabolic processes that precede germination. After that, seed, germination is avoided by dehydration, as part of the priming treatments. After dehydration, seeds maintain their metabolic advances (hydration memory). As a result, rehydrated seeds germinate rapidly. We hypothesized that, as seeds, fern spores may be capable of developing hydration memory. To assess priming, spores of six fern species were exposed to: four or eight days of hydration in water (hydro-priming) or in a soil matrix (matrix-priming); or 1 month of hydration in the soil of the collection site (natural-priming). At the end of the treatments, the spores were dehydrated in the dark and germinated under laboratory conditions. Germination was evaluated using lag-time, germination rate and germination percentage. Priming treatments shortened lag time and/or increased germination rate or germination percentage in relation to the controls. Matrix-priming (8 days) reduced the spore germination percentage in three species. Our results provide evidence that fern spores possess a hydration memory that probably evolved in the soil bank and suggests that hydration-dehydration cycles within the natural soil might provide advantages for successful germination.

Gametophyte morphology of Acrostichum aureum and A. danaeifolium (Pteridaceae) / Morfología del gametofito de Acrostichum aureum y Acrostichum danaeifolium (Pteridaceae)

Abstract Acrostichum is a pantropical genus and has four species, two of which occur in the Neotropics, A. aureum and A. danaeifolium. In Mexico, A. danaeifolium grows further in land wet soils and is much more common than A. aureum, which is typically found in brackish or saline habitats near the coast, and is restricted to coastal saline mangrove communities. The purpose of this paper was to describe and compare the morphogenesis of the sexual phase of A. aureum and A. danaeifolium for systematic purposes. For this, spores of each species were sown in Petri dishes with agar, previously enriched with sterilized Thompson's medium. To avoid contamination and dehydration, the dishes were kept in transparent plastic bags under laboratory conditions. For the micro-morphological observation with SEM, the gametophyte development phases were fixed in FAA with 0.8 % sucrose for 24 h. Photomicrographs of spores, development stages of gametophytes and young sporophytes were observed with scanning electron microscope. Our results showed that the spores of both species are triletes, globose and positive photoblastic. Germination is Vittaria-type; the germinate filaments are short and uniseriate (5 to 7 cells), and prothallial development is Ceratopteris-type. The adult gametophytes of both species have asymmetrical wings. Adult gametophytes in culture are cordiform-spatulate. Antheridia have a broad basal cell, an annular cell, and an asymmetric opercular cell. Archegonia have short necks and four triangular cells at the mouth of the neck. The first leaf of the sporophyte is lobed, with dichotomous veins and anomocytic stomata. The gemmae are formed in adult gametophytes in both species. The development of the gametophyte of A. aureum, A. danaeifolium and A. speciosum share many similarities such as the development of a lateral meristem, asymmetric nature of the mature prothallus, lack of hairs on the prothallus, and undivided asymmetrical opercular antheridia morphology. The genus Acrostichum is the sister group of Ceratopteris, another genus of aquatic ferns; they differ in the antheridium morphology, Acrostichum has an asymmetric opercular cell and Ceratopteris shows an undivided cap cell, but the notable difference is the sporophyte morphology. Rev. Biol. Trop. 66(1): 178-188. Epub 2018 March 01.

Resumen Acrostichum es pantropical y tiene cuatro especies dos de las cuáles ocurren en el Neotrópico, A. aureum y A. danaeifolium. El género se encuentra típicamente en hábitats salobres o salinos cerca de la costa. También puede crecer más lejos en tierra en suelos húmedos. Acrostichum aureum está restringido a las comunidades de manglares salinos costeros. A. danaeifolium es mucho más común que A. aureum en México. El propósito de este trabajo es describir y comparar la morfogénesis de la fase sexual de A. aureum y A. danaeifolium. Las esporas de cada especie fueron sembradas en cajas de Petri, en agar previamente enriquecido con medio de Thompson esterilizado. Para evitar la contaminación y la deshidratación, las cajas se mantuvieron dentro de bolsas de plástico transparentes en condiciones de laboratorio. Para la observación micro-morfológica con el MEB, las fases de desarrollo del gametofito se fijaron en FAA con sacarosa al 0.8 % durante 24 h. Se observaron las esporas, fases de desarrollo de gametofitos y esporofitos jóvenes con un microscopio electrónico de barrido. Las esporas de ambas especies son triletes, globosas y fotoblásticas positivas. La germinación es tipo-Vittaria, los filamentos germinativos son cortos y uniseriados (5 a 7 células) y el desarrollo protálico es de tipo-Ceratopteris. Los gametofitos de ambas especies tienen alas asimétricas. Los gametofitos adultos en cultivo son cordiforme-espatulados. Los anteridios tienen una célula basal amplia, una célula anular y una célula opercular asimétrica. Los arquegonios tienen cuellos cortos y cuatro células triangulares en la boca del cuello. La primera hoja del esporofito es lobulada, con venación dicotómica y estomas anomocíticos. Las yemas se forman en gametofitos adultos en ambas especies. El desarrollo del gametofito de A. aureum, A. danaeifolium y A. speciosum comparten características tales como el desarrollo de un meristemo lateral, el protalo maduro de naturaleza asimétrica, ausencia de pelos en el protalo y anteridio con célula opercular asimétrica no dividida. Acrostichum es el grupo hermano de Ceratopteris, otro género de helecho acuático, que difieren en la morfología del anteridio, Acrostichum tiene una célula opercular asimétrica y Ceratopteris muestran una célula opercular no dividida.

Desarrollo de los gametofitos de especies mexicanas de helechos de la familia Gleicheniaceae / Gametophyte development in Mexican species of Gleicheniaceae ferns

Resumen: En México, la familia Gleicheniaceae está representada por Dicranopteris flexuosa, Diplopterygium bancroftii, Gleichenella pectinata, Sticherus bifidus, S. brevipubis, S. palmatus y S. underwoodianus; existen pocos trabajos sobre el estudio de los gametofitos de algunas especies de esta familia, por lo que el objetivo es describir, comparar y aportar información sobre su germinación, desarrollo de los gametofitos y finalmente determinar si los caracteres de los prótalos son útiles en la delimitación taxonómica del grupo. Para ello se recolectaron ejemplares y esporas de cada taxón en el campo, las esporas se sembraron en cajas de Petri que contenían agar y medio nutritivo de Thompson, se cultivaron en germinadoras bajo condiciones controladas de luz (12hrs luz/oscuridad), humedad (50 %) y temperatura (18 °C noche y 25 °C día); se realizaron observaciones de material fresco y se tomaron fotomicrografías utilizando un microscopio óptico y un microscopio electrónico de barrido. Como resultado se distinguieron dos tipos de germinación Gleichenia y Cyathea. En cuanto al desarrollo protálico, se observaron tres tipos, Marattia, Osmunda y Drynaria. Debido a las variaciones en el tipo de germinación que no habían sido reportadas para la familia en la literatura, así como por la heterogeneidad presente en el patrón de desarrollo protálico y la presencia de más de cuatro células en los gametangios, es importante ampliar el estudio de otras especies para determinar el valor taxonómico de las características morfológicas de los gametofitos, así como para determinar si estas variaciones se presentan en otras especies de la familia.

Abstract: In Mexico, the Gleicheniaceae family is represented by different species such as Dicranopteris flexuosa, Diplopterygium bancroftii, Gleichenella pectinata, Sticherus bifidus, S. brevipubis, S. palmatus and S. underwoodianus. Currently, few studies have described the gametophytes of some species in this family, and our objective was to contribute to the knowledge, and to describe and compare different aspects of their germination, gametophyte development, and to determine if the prothallus characters are useful for taxonomic delimitations in the group. For this purpose, specimens and spores of each taxon were collected in the field, spores were sown in Petri dishes containing agar and Thompson nutrient medium, and grown in a plant growing chamber under controlled conditions of light (12 hr light/darkness), (50 %) humidity, and temperature (18 °C night, 25 °C day). Additionally, observations of fresh materials were made and photomicrographs were taken using both optical and scanning electron microscopes. Our observations allowed distinguishing two types of germination Gleichenia and Cyathea; and three types of prothallial development Marattia, Osmunda and Drynaria. Gametangia presented more than three cells, and this is considered a primitive feature by other authors. As some variations in the germination type were observed and have not previously been reported in the literature for this family, and because of the heterogenity in the patterns of the prothallial cell development, and gametangia of more than four cells, it is important to broaden the study to other species, in order to determine the taxonomic value of the morphological characters of the gametophyte, as well as to determine if these variations are present in other species of the family. Rev. Biol. Trop. 65 (3): 939-952. Epub 2017 September 01.

Observations of the spore, gametophyte and young sporophyte of Pteridium caudatum (L.) Maxon using scanning electron microscopy.

This paper describes the development of the sexual phase of the invasive fern, Pteridium caudatum, from spore germination to young sporophyte formation. Spores samples for gametophyte cultures were taken from various sporophytes and then sown on mineral agar with Thompson's media. Gametophytes were maintained under fluorescent light on a 12h light, 12h dark cycle at 24-25°C. Developmental phases were fixed in FAA-sucrose solution and processed for observation with the scanning electron microscope. Spores are trilete and germination takes place on the second day after sowing; germination is of the Vittaria-type. Adiantum-type prothallial development was observed. The differentiation of a two-dimensional thallus begins 5 days after germination maturation of adult gametophytes occurs about 30 days after sowing. Adult gametophytes are heart-shaped, bisexual and glabrous. Antheridia are formed by three cells: basal, annular and opercular cell with a pore. Archegonia have a neck of 4-cells. The young sporophyte becomes visible within 8 weeks after spores are sown. The taxonomic significance of the gametophyte morphology is discussed.

Gametophyte morphology of Platycerium andinum Baker and Platycerium wandae Racif.

This paper describes the morphology of the sexual phase and spores of Platycerium andinum and Platycerium wandae. Spores were sown in Thompson's media and the cultures were kept at 24-25 degrees C, with 12h light/darkness photoperiod. Developmental phases were fixed in FAA and processed for observation with the scanning electron microscope. Spores of both species are monolete; Vittaria-type germination and Aspidium-type prothallial development were observed. In the phase of development, the gametophytes develop unicellular secretory and as they mature, develop bifurcated or branched pluricellular trichomes, both in the cushion and near the meristematic zone. Adult gametophytes in culture are cordiform-spatulate to cordiform-reniform, most are unisexual and a few are bisexual. Gametangia belong to the leptosporangiate fern type. Archegonial morphology is uniform, with an elongate, thin neck curved toward the base of the gametophyte. Antheridia have a basal cell, an annular cell and an undivided opercular cell. Three hundred days after the spores were sown, sporophytes still had not developed. In both species, some spores germinate inside the sporangial capsule (intra-sporangial germination). We provide new information on morphogenesis in the genus Platycerium.

Developmental gametophyte morphology of seven species of Thelypteris subg. Cyclosorus (Thelypteridaceae).

The morphogenesis of the sexual phase of seven species of Thelypteris subg. Cyclosorus (Thelypteridaceae) is described and compared. Spores of all species are monolete, ellipsoid and have positive photoblastism. They have Vittaria-type germination, germinal filaments are short and uniseriate, and Aspidium-type development. Adult gametophytes are spatulate-cordiform, and have wide wings with numerous glandular, marginal and superficial hairs. Antheridia have a narrow or widened basal cell, an undivided annular cell, and an opercular cell; antherozoids are liberated through three mechanisms. Archegonia have short necks and four triangular cells on the neck's mouth. The sporophyte's first leaf is lobed, with open dichotomous veins, glandular hairs similar to those of the gametophyte, and anomocytic stomata. Differences observed between the studied species are: spore size; ornamentation of the perispore; germination and emergence periods; size of hairs; developmental period of gametangia; and antherozoid liberation mechanisms through total detachment of the operculum, partial detachment of the operculum, and development of an apical pore through tearing of the operculum. These taxonomically valuable characters, combined with the sporophyte characteristics, can be useful tools in the identification of the species studied to the subg. Cyclosorus.

Analysis of fern spore banks from the soil of three vegetation types in the central region of Mexico.

The vertical structure of fern spore banks was studied in a xerophilous shrubland, montane rain forest, and pine-oak forest in Hidalgo, Mexico, using the emergence method. Soil samples were collected in April 1999 at depths of 0-10, 10-20, and 20-30 cm. Viable spores decreased significantly with depth in all vegetation types, and the highest number of prothallia and sporophytes was found in the uppermost layer. The montane rain forest and the xerophilous shrubland had the largest and the richest banks, respectively. Twenty-three fern taxa were registered in the aboveground vegetation, 12 in the soil banks, and 43.5% were in both. Aboveground and in the soil bank, the xerophilous shrubland, the montane rain forest, and the pine-oak forest had, 17 and 7, 1 and 6, and 7 and 3 taxa, respectively. These were distributed differentially in relation to depth. The Sørensen index indicated a similarity of 61.5% between the xerophilous shrubland and the montane rain forest, and the Czeckanovsky index indicated 19.75%. The presence of viable spores in the soil of all vegetation types confirmed the existence of natural spore banks. Long-distance dispersal was an important factor determining the specific composition of the xerophilous shrubland and the pine-oak forest.

Fase sexual de los helechos Odontosoria schlechtendalii y Odontosoria scandens (Dennstaedtiaceae) / Sexual phase of the ferns Odontosoria schlechtendalii and Odontosoria scandens (Dennstaedtiaceae)

The results of morphogenesis studies of the sexual phase of Odontosoria schlechtendalii and O. scandens are presented in this paper. In O. schlechtendalii and O. scandens the spores are triletes, non-chlorophyllous, and germinación is of the Vittaria-type, devoid of perine, the exine is smooth and sometimes coarsely ridged to reticulate. The development pattern is of the Adiantum-type. The adult gametophyte is cordate-spathulate, with probable presence of anteridiogen in O. schlechtendalii, both species are glabrous. Sex organs are of the common type of the leptosporangiate ferns. The first leaves appeared after 56-92 days of culture, with petiole and plate divide narrow, trichomes bifurcate and stomate anomocytic.

Morfogénesis de los gametofitos de especies mexicanas de Pleopeltis (Polypodiaceae, subfamilia Pleopeltoideae) / Gametophyte morphogenesis of Mexican species of Pleopeltis (Polypodiaceae, subfamily Pleopeltoideae)

The development and morphology of the gametophytes of seven species of ferns from genus Pleopeltis are described and compared. The spore germination is Vittaria-type in P. astrolepis, P. crassinervata, P. macrocarpa, P. polylepis and P. revoluta. For P. angusta and P. mexicana it was proposed a new germination pattern is Pleopeltis-type. The prothallial development is Drynaria-type in P. astrolepis, P. crassinervata, P. macrocarpa, P. polylepis and P. revoluta and Ceratopteris-type for P. angusta and P. mexicana. The gametangia are typical of the leptosporangiate ferns, sporophytes after six and a half months in culture did not appeared.

[Gametophyte morphogenesis of Mexican species of Pleopeltis (Polypodiaceae, subfamily Pleopeltoideae)]. / Morfogénesis de los gametofitos de especies mexicanas de Pleopeltis (Polypodiaceae, subfamilia Pleopeltoideae).

The development and morphology of the gametophytes of seven species of ferns from genus Pleopeltis are described and compared. The spore germination is Vittaria-type in P. astrolepis, P. crassinervata, P. macrocarpa, P. polylepis and P. revoluta. For P. angusta and P. mexicana it was proposed a new germination pattern is Pleopeltis-type. The prothallial development is Drynaria-type in P. astrolepis, P. crassinervata, P. macrocarpa, P. polylepis and P. revoluta and Ceratopteris-type for P. angusta and P. mexicana. The gametangia are typical of the leptosporangiate ferns, sporophytes after six and a half months in culture did not appeared.

[Sexual phase of the ferns Odontosoria schlechtendalii and Odontosoria scandens (Dennstaedtiaceae)]. / Fase sexual de los helechos Odontosoria schlechtendalii y Odontosoria scandens (Dennstaedtiaceae).

The results of morphogenesis studies of the sexual phase of Odontosoria schlechtendalii and O. scandens are presented in this paper. In O. schlechtendalii and O. scandens the spores are triletes, non-chlorophyllous, and germinación is of the Vittaria-type, devoid of perine, the exine is smooth and sometimes coarsely ridged to reticulate. The development pattern is of the Adiantum-type. The adult gametophyte is cordate-spathulate, with probable presence of anteridiogen in O. schlechtendalii, both species are glabrous. Sex organs are of the common type of the leptosporangiate ferns. The first leaves appeared after 56-92 days of culture, with petiole and plate divide narrow, trichomes bifurcate and stomate anomocytic.

Morfología vegetal neotropical / Neotropical plant morphology

An analysis on plant morphology and the sources that are important to the morphologic interpretations is done. An additional analysis is presented on all published papers in this subject by the Revista de Biología Tropical since its foundation, as well as its contribution to the plant morphology development in the neotropics

[Neotropical plant morphology]. / Morfología vegetal neotropical.

An analysis on plant morphology and the sources that are important to the morphologic interpretations is done. An additional analysis is presented on all published papers in this subject by the Revista de Biología Tropical since its foundation, as well as its contribution to the plant morphology development in the neotropics.

Morfogénesis de la fase sexual de seis especies mexicanas del género Dryopteris (Dryopteridaceae), Parte II / Morphogenesis of the sexual phase of six Mexican species of the Dryopteris (Dryopteridaceae), Part II

The development of the sexual phase of six Mexican species of Dryopteris is described and compared. Spores of all studied species are monolete, ellipsoid and have a rugose surface; the perine is folded, brown to dark brown, with a tubercled outline. Germination pattern is of the Vittaria-type and the development pattern of the prothallia is of the Aspidium-type. Gametangia are of the common type for the leptosporangiate advanced ferns. First leaves of the sporophytes appear 258-265 after sowing and apparently in Dryopteris pseudo-filix-mas the sporophyte have an apogamic origin (80 days). To make a comparative analysis of gametophytic characteristics in the twelve Mexican species and conclude of germination is of the Vittaria-type and development pattern prothallial is of the Aspidium-type, and unicelular trichomes on margin and superficial gametophytic to yield irregular aspect are characteristics to yield unit and characteristic to genera to conform Dryopteridaceae family (sensu Moran 1995) with the exception of Didymochlaena genus.