Centromere drive may propel the evolution of chromosome and genome size in plants.

BACKGROUND: Genome size is influenced by natural selection and genetic drift acting on variations from polyploidy and repetitive DNA sequences. We hypothesized that centromere drive, where centromeres compete for inclusion in the functional gamete during meiosis, may also affect genome and chromosome size. This competition occurs in asymmetric meiosis, where only one of the four meiotic products becomes a gamete. If centromere drive influences chromosome size evolution, it may also impact post-polyploid diploidization, where a polyploid genome is restructured to function more like a diploid through chromosomal rearrangements, including fusions. We tested if plant lineages with asymmetric meiosis exhibit faster chromosome size evolution compared to those with only symmetric meiosis, which lack centromere drive as all four meiotic products become gametes. We also examined if positive selection on centromeric histone H3 (CENH3), a protein that can suppress centromere drive, is more frequent in these asymmetric lineages. METHODS: We analyzed plant groups with different meiotic modes: asymmetric in gymnosperms and angiosperms, and symmetric in bryophytes, lycophytes, and ferns. We selected species based on available CENH3 gene sequences and chromosome size data. Using Ornstein-Uhlenbeck evolutionary models and phylogenetic regressions, we assessed the rates of chromosome size evolution and the frequency of positive selection on CENH3 in these clades. RESULTS: Our analyses showed that clades with asymmetric meiosis have a higher frequency of positive selection on CENH3 and increased rates of chromosome size evolution compared to symmetric clades. CONCLUSIONS: Our findings support the hypothesis that centromere drive accelerates chromosome and genome size evolution, potenatially also influencing the process of post-polyploid diploidization. We propose a model which in a single famework helps explain the stability of chromosome size in symmetric lineages (bryophytes, lycophytes, and ferns) and its variability in asymmetric lineages (gymnosperms and angiosperms), providing a foundation for future research in plant genome evolution.

Morphological and functional evolution of gametophytes in epilithic Hymenasplenium murakami-hatanakae (Aspleniaceae): The fifth family capable of producing the independent gametophytes.

The fern independent gametophytes that can maintain populations by vegetative reproduction without conspecific sporophytes have been considered an unusual phenomenon found in some epiphytic or epilithic species of Hymenophyllaceae, Pteridaceae, Lomariopsidaceae, and Polypodiaceae. By chance, the discovery of mysterious strap-like gametophytes on Izu-Oshima Island, Japan, has led to the hypothesis that Hymenasplenium murakami-hatanakae, a fern species belonging to Aspleniaceae, can also form independent gametophytes. Our investigation revealed gametophyte populations of H. murakami-hatanakae on three islands in the Izu Islands. Based on chloroplast DNA analysis of the gametophyte and sporophyte populations, the gametophytes were found to be maintained by vegetative reproduction without a new supply of spores from sporophytes. A comparison of the surrounding vegetation at the collection sites showed that environmental factors such as light and humidity may influence the maintenance of gametophyte populations. These results clearly show that H. murakami-hatanakae is one of the ferns capable of forming independent gametophytes. This is the first report of independent gametophytes from the suborder Aspleniineae (eupolypod II). The discovery of the independent gametophyte within a phylogenetic lineage previously thought not to form independent gametophytes will provide important insights into the morphological and functional evolution of gametophytes in ferns.

Polyploid goldback and silverback ferns (Pentagramma) occupy a wider, colder, and wetter bioclimatic niche than diploid counterparts.

PREMISE: The western North American fern genus Pentagramma (Pteridaceae) is characterized by complex patterns of ploidy variation, an understanding of which is critical to comprehending both the evolutionary processes within the genus and its current diversity. METHODS: We undertook a cytogeographic study across the range of the genus, using a combination of chromosome counts and flow cytometry to infer ploidy level. Bioclimatic variables and elevation were used to compare niches. RESULTS: We found that diploids and tetraploids are common and widespread, and triploids are rare and sporadic; in contrast with genome size inferences in earlier studies, no hexaploids were found. Diploids and tetraploids show different geographic ranges: only tetraploids were found in the northernmost portion of the range (Washington, Oregon, and British Columbia) and only diploids were found in the Sierra Nevada of California. Diploid, triploid, and tetraploid cytotypes were found to co-occur in relatively few localities: in the southern (San Diego County, California) and desert Southwest (Arizona) parts of the range, and along the Pacific Coast of California. CONCLUSIONS: Tetraploids occupy a wider bioclimatic niche than diploids both within P. triangularis and at the genus-wide scale. It is unknown whether the wider niche of tetraploids is due to their expansion upon the diploid niche, if diploids have contracted their niche due to competition or changing abiotic conditions, or if this wider niche occupancy is due to multiple origins of tetraploids.

Living jewels: iterative evolution of iridescent blue leaves from helicoidal cell walls.

BACKGROUND AND AIMS: Structural colour is responsible for the remarkable metallic blue colour seen in the leaves of several plants. Species belonging to only ten genera have been investigated to date, revealing four photonic structures responsible for structurally coloured leaves. One of these is the helicoidal cell wall, known to create structural colour in the leaf cells of five taxa. Here we investigate a broad selection of land plants to understand the phylogenetic distribution of this photonic structure in leaves. METHODS: We identified helicoidal structures in the leaf epidermal cells of 19 species using transmission electron microscopy. Pitch measurements of the helicoids were compared with the reflectance spectra of circularly polarized light from the cells to confirm the structure-colour relationship. RESULTS: By incorporating species examined with a polarizing filter, our results increase the number of taxa with photonic helicoidal cell walls to species belonging to at least 35 genera. These include 19 monocot genera, from the orders Asparagales (Orchidaceae) and Poales (Cyperaceae, Eriocaulaceae, Rapateaceae) and 16 fern genera, from the orders Marattiales (Marattiaceae), Schizaeales (Anemiaceae) and Polypodiales (Blechnaceae, Dryopteridaceae, Lomariopsidaceae, Polypodiaceae, Pteridaceae, Tectariaceae). CONCLUSIONS: Our investigation adds considerably to the recorded diversity of plants with structurally coloured leaves. The iterative evolution of photonic helicoidal walls has resulted in a broad phylogenetic distribution, centred on ferns and monocots. We speculate that the primary function of the helicoidal wall is to provide strength and support, so structural colour could have evolved as a potentially beneficial chance function of this structure.

'Out of Africa' origin of the pantropical staghorn fern genus Platycerium (Polypodiaceae) supported by plastid phylogenomics and biogeographical analysis.

BACKGROUND AND AIMS: The staghorn fern genus Platycerium is one of the most commonly grown ornamental ferns, and it evolved to occupy a typical pantropical intercontinental disjunction. However, species-level relationships in the genus have not been well resolved, and the spatiotemporal evolutionary history of the genus also needs to be explored. METHODS: Plastomes of all the 18 Platycerium species were newly sequenced. Using plastome data, we reconstructed the phylogenetic relationships among Polypodiaceae members with a focus on Platycerium species, and further conducted molecular dating and biogeographical analyses of the genus. KEY RESULTS: The present analyses yielded a robustly supported phylogenetic hypothesis of Platycerium. Molecular dating results showed that Platycerium split from its sister genus Hovenkampia ~35.2 million years ago (Ma) near the Eocene-Oligocene boundary and began to diverge ~26.3 Ma during the late Oligocene, while multiple speciation events within Platycerium occurred during the middle to late Miocene. Biogeographical analysis suggested that Platycerium originated in tropical Africa and then dispersed eastward to southeast Asia-Australasia and westward to neotropical areas. CONCLUSIONS: Our analyses using a plastid phylogenomic approach improved our understanding of the species-level relationships within Platycerium. The global climate changes of both the Late Oligocene Warming and the cooling following the mid-Miocene Climate Optimum may have promoted the speciation of Platycerium, and transoceanic long-distance dispersal is the most plausible explanation for the pantropical distribution of the genus today. Our study investigating the biogeographical history of Platycerium provides a case study not only for the formation of the pantropical intercontinental disjunction of this fern genus but also the 'out of Africa' origin of plant lineages.

The traditional utilization, biological activity and chemical composition of edible fern species.

ETHNOPHARMACOLOGICAL RELEVANCE: Ferns form an important part of the human diet. Young fern fiddleheads are mostly consumed as vegetables, while the rhizomes are often extracted for starch. These edible ferns are also often employed in traditional medicine, where all parts of the plant are used, mostly to prepare extracts. These extracts are applied either externally as lotions and baths or internally as potions, decoctions and teas. Ailments traditionally treated with ferns include coughs, colds, fevers, pain, burns and wounds, asthma, rheumatism, diarrhoea, or skin diseases (eczema, rashes, itching, leprosy). AIM OF THE REVIEW: This review aims to compile the worldwide knowledge on the traditional medicinal uses of edible fern species correlating to reported biological activities and isolated bioactive compounds. MATERIALS AND METHODS: The articles and books published on edible fern species were searched through the online databases Web of Science, Pubmed and Google Scholar, with critical evaluation of the hits. The time period up to the end of 2022 was included. RESULTS: First, the edible fern species were identified based on the literature data. A total of 90 fern species were identified that are eaten around the world and are also used in traditional medicine. Ailments treated are often associated with inflammation or bacterial infection. However, only the most common and well-known fern species, were investigated for their biological activity. The most studied species are Blechnum orientale L., Cibotium barometz (L.) J. Sm., Diplazium esculentum (Retz.) Sw., Marsilea minuta L., Osmunda japonica Thunb., Polypodium vulgare L., and Stenochlaena palustris (Burm.) Bedd. Most of the fern extracts have been studied for their antioxidant, anti-inflammatory and antimicrobial activities. Not surprisingly, antioxidant capacity has been the most studied, with results reported for 28 edible fern species. Ferns have been found to be very rich sources of flavonoids, polyphenols, polyunsaturated fatty acids, carotenoids, terpenoids and steroids and most of these compounds are remarkable free radical scavengers responsible for the outstanding antioxidant capacity of fern extracts. As far as clinical trials are concerned, extracts from only three edible fern species have been evaluated. CONCLUSIONS: The extracts of edible fern species exert antioxidant anti-inflammatory and related biological activities, which is consistent with their traditional medicinal use in the treatment of wounds, burns, colds, coughs, skin diseases and intestinal diseases. However, studies to prove pharmacological activities are scarce, and require chemical-biological standardization. Furthermore, correct botanical classification needs to be included in publications to simplify data acquisition. Finally, more in-depth phytochemical studies, allowing the linking of traditional use to pharmacological relevance are needed to be done in a standardized way.

Taxonomic updates of Cheilanthes Sw. (Pteridaceae) in Peru, with a conservation assessment for four endemics / Actualización taxonómica de Cheilanthes Sw. (Pteridaceae) en Perú, con una evaluación de conservación para cuatro endemismos

Resumen El género Cheilanthes en su concepto más reciente incluye tres clados distribuidos en Sudamérica, África y Australasia. Trabajos previos para el Perú registraron 23 especies, sin embargo, los avances en sistemática molecular han resultado en la segregación de ocho de ellas en al menos tres géneros. En este trabajo, se reconocen 15 especies en la flora peruana en base al estudio de 459 ejemplares de herbario y campo, y la evaluación de caracteres morfológicos usando 131 especímenes. Las afinidades entre los linajes propuestos en este trabajo son consistentes con la reciente hipótesis filogenética, donde las especies de Cheilanthes en el Perú serían integrantes de dos clados, el Sudamericano y el Australásico-Sudamericano, y los caracteres de importancia taxonómica serian: la forma de las escamas rizomáticas y tipos de indumento de la fronda (escamas, microescamas, escuámulas y pelos). Encontramos que el género Cheilanthes estaría presente en 20 departamentos del Perú, la mayoría de las especies con amplia distribución en las regiones centro-sur Andinas, cuatro especies con el estatus de endemismo. Mientras que C. cantangensis, C. lonchophylla y C. obducta están pobremente representadas en la región norte. Las dos primeras especies son consideradas endémicas y son categorizadas como En Peligro (EN), y otras dos como de Preocupación Menor (LC). Además, se presenta una clave taxonómica.

Abstract In its most recent concept, the genus Cheilanthes encompasses three clades distributed across South America, Africa, and Australasia. Previous studies in Peru recorded 23 species; however, advancements in molecular systematic have led to the segregation of eight of them into at least three genera. In this study, 15 species are recognized in the Peruvian flora based on the examination of 459 herbarium and field specimens, and the evaluation of morphological characters using 131 specimens. The affinities among the lineages proposed in this study are consistent with recent phylogenetic hypotheses, wherein Cheilanthes species in Peru are members of two clades, the South American and the Australasian-South American, with taxonomically important characters being the shape of rhizomatous scales and types of frond indumentum (scales, microscales, scuamules, and hairs). We found that the genus Cheilanthes is present in 20 departments of Peru, with most species having wide distribution in the central-southern Andean regions, four species having endemic status. Meanwhile, C. cantangensis, C. lonchophylla, and C. obducta are poorly represented in the northern region. The first two species are considered endemic and categorized as Endangered (EN), while the other two are classified as Least Concern (LC). Additionally, a taxonomic key is provided.

Ethnomedicinal and traditional uses of the Ferns of Khyber Pakhtunkhwa, Pakistan / Usos etnomedicinais e tradicionais das samambaias de Khyber Pakhtunkhwa, Paquistão

Abstract Ferns are often used by indigenous people in Khyber Pakhtunkhwa, Pakistan. This study was designed to collect the ethnomedicinal and traditional knowledge of these locals about this group of vascular plants. Forty taxa belong to nineteen genera and ten families are used in the treatment of different diseases. The Pteridaceae was the most representative family with twelve taxa (30%), followed by Athyriaceae and Dryopteridaceae with six taxa each (30%), and Thelypteridaceae with five taxa (12.5%). Regarding the genera, Adiantum, Asplenium and Dryopteris ranked first with four taxa each (30%), followed by Aleuritopteris, Diplazium, Pteris and Equisetum with three taxa each (30%), followed by Athyrium, Oeosporangium, Polystichum and Pseudophegopteris with two taxa each (20%). These taxa were commonly used in the treatment of respiratory disorders i.e. asthma, bronchitis, emphysema, pneumonia; intestinal ulcer, stomach, urinary ailments and skin disorders by the methods of decoction and infusion. Traditional knowledge about ethnomedicinal plants is a valuable and essential source for the discovery of allopathic, herbal and homeopathic medicines.

Resumo As samambaias são frequentemente usadas pelos indígenas em Khyber Pakhtunkhwa, Paquistão. Este estudo foi desenhado para coletar o conhecimento etnomedicinal e tradicional desses moradores sobre este grupo de plantas vasculares. Quarenta táxons pertencem a dezenove gêneros, e dez famílias são utilizadas no tratamento de diferentes doenças. Pteridaceae foi a família mais representativa com doze táxons (30%), seguida por Athyriaceae e Dryopteridaceae com seis táxons cada (30%), e Thelypteridaceae com cinco táxons (12,5%). Em relação aos gêneros, Adiantum, Asplenium e Dryopteris ficaram em primeiro lugar com quatro táxons cada (30%), seguidos por Aleuritopteris, Diplazium, Pteris e Equisetum com três táxons cada (30%), e Athyrium, Oeosporangium, Polystichum e Pseudophegopteris com dois táxons cada (20%). Estes táxons foram comumente usados ​​no tratamento de distúrbios respiratórios, isto é, asma, bronquite, enfisema, pneumonia; úlcera intestinal, estômago, doenças urinárias e doenças da pele pelos métodos de decocção e infusão. O conhecimento tradicional sobre plantas etnomedicinais é uma fonte valiosa e essencial para a descoberta de medicamentos alopáticos, fitoterápicos e homeopáticos.

Ethnomedicinal and traditional uses of the Ferns of Khyber Pakhtunkhwa, Pakistan

Abstract Ferns are often used by indigenous people in Khyber Pakhtunkhwa, Pakistan. This study was designed to collect the ethnomedicinal and traditional knowledge of these locals about this group of vascular plants. Forty taxa belong to nineteen genera and ten families are used in the treatment of different diseases. The Pteridaceae was the most representative family with twelve taxa (30%), followed by Athyriaceae and Dryopteridaceae with six taxa each (30%), and Thelypteridaceae with five taxa (12.5%). Regarding the genera, Adiantum, Asplenium and Dryopteris ranked first with four taxa each (30%), followed by Aleuritopteris, Diplazium, Pteris and Equisetum with three taxa each (30%), followed by Athyrium, Oeosporangium, Polystichum and Pseudophegopteris with two taxa each (20%). These taxa were commonly used in the treatment of respiratory disorders i.e. asthma, bronchitis, emphysema, pneumonia; intestinal ulcer, stomach, urinary ailments and skin disorders by the methods of decoction and infusion. Traditional knowledge about ethnomedicinal plants is a valuable and essential source for the discovery of allopathic, herbal and homeopathic medicines.

Resumo As samambaias são frequentemente usadas pelos indígenas em Khyber Pakhtunkhwa, Paquistão. Este estudo foi desenhado para coletar o conhecimento etnomedicinal e tradicional desses moradores sobre este grupo de plantas vasculares. Quarenta táxons pertencem a dezenove gêneros, e dez famílias são utilizadas no tratamento de diferentes doenças. Pteridaceae foi a família mais representativa com doze táxons (30%), seguida por Athyriaceae e Dryopteridaceae com seis táxons cada (30%), e Thelypteridaceae com cinco táxons (12,5%). Em relação aos gêneros, Adiantum, Asplenium e Dryopteris ficaram em primeiro lugar com quatro táxons cada (30%), seguidos por Aleuritopteris, Diplazium, Pteris e Equisetum com três táxons cada (30%), e Athyrium, Oeosporangium, Polystichum e Pseudophegopteris com dois táxons cada (20%). Estes táxons foram comumente usados no tratamento de distúrbios respiratórios, isto é, asma, bronquite, enfisema, pneumonia; úlcera intestinal, estômago, doenças urinárias e doenças da pele pelos métodos de decocção e infusão. O conhecimento tradicional sobre plantas etnomedicinais é uma fonte valiosa e essencial para a descoberta de medicamentos alopáticos, fitoterápicos e homeopáticos.

Determination of ptaquiloside in ten species of Pteris in northwestern Argentina.

The toxic effect of ferns of the genus of Pteris in bovines is caused by ptaquiloside, the main carcinogenic toxin. In this study, ten species of Pteris fern in different phenologic stages and plant conditions were collected in northwest Argentina. The phytochemical analysis showed the presence of Pt in the recent collected samples (adults and young plants) but not in the herbarium specimens. The results show a great variation of Pt concentration that depends on the phenologic stage, plant condition, and collection site. Pt was measured in 6-4326 µg/g concentration, with a mean concentration of 644 µg/g. No Pt was detected in eight species of Pteris collected from herbarium samples; such results may be a false negative. It is important to notice that analysis of herbarium samples for Pt may not be a reliable method to determine its presence. It is important to further understand the potential toxicity caused by these ferns because of their effect on animals, public health, and the environment.

Interploidy gene flow via a 'pentaploid bridge' and ploidy reduction in Cystopteris fragilis fern complex (Cystopteridaceae: Polypodiales).

KEY MESSAGE: Our results indicate the existence of interploidy gene flow in Cystopteris fragilis, resulting in sexual triploid and diploid gametophytes from pentaploid parents. Similar evolutionary dynamics might operate in other fern complexes and need further investigation. Polyploidization and hybridization are a key evolutionary processes in ferns. Here, we outline an interploidy gene flow pathway operating in the polyploid Cystopteris fragilis complex. The conditions necessary for the existence of this pathway were tested. A total of 365 C. fragilis individuals were collected covering representatives of all three predominant ploidy levels (tetraploid, pentaploid, and hexaploid), cultivated, had their ploidy level estimated by flow cytometry, and their spores collected. The spores, as well as gametophytes and sporophytes established from them, were analysed by flow cytometry. Spore abortion rate was also estimated. In tetraploids, we observed the formation of unreduced (tetraploid) spores (ca 2%). Collected pentaploid individuals indicate ongoing hybridization between ploidy levels. Pentaploids formed up to 52% viable spores, ca 79% of them reduced, i.e. diploid and triploid. Reduced spores formed viable gametophytes, and, in the case of triploids, filial hexaploid sporophytes, showing evidence of sexual reproduction. Some tetraploid sporophytes reproduce apomictically (based on uniform ploidy of their metagenesis up to filial sporophytes). Triploid and diploid gametophytes from pentaploid parents are able to mate among themselves, or with "normal" reduced gametophytes from the sexual tetraploid sporophytes (the dominant ploidy level in the sporophytes in this populations), to produce tetraploid, pentaploid, and hexaploid sporophytes, allowing for geneflow from the pentaploids to both the tetraploid and hexaploid populations. Similar evolutionary dynamics might operate in other fern complexes and need further investigation.

Phylogenomic evolutionary insights in the fern family Gleicheniaceae.

The pantropical fern family Gleicheniaceae comprises approximately 157 species. Seven genera are currently recognized in the family, although their monophyly is still uncertain due to low sampling in phylogenetic studies. We examined the monophyly of the genera through extended sampling, using the first phylogenomic inference of the family including data from both nuclear and plastid genomes. Seventy-six samples were sequenced (70 Gleicheniaceae species and six outgroups) using high throughput sequencing, including all seven currently recognized genera. Plastid and nuclear data were recovered and assembled; the nuclear data was phased to reduce paralogy as well as hybrid noise in the final recovered topology. Maximum likelihood trees were built for each locus, and a concatenated dataset was built for both datasets. A species tree based on a multispecies coalescent model was generated, and divergence time analyses performed. We here present the first genomic phylogenetic inferences concerning Gleicheniaceae, confirming the monophyly of most genera except Sticherus, which we recovered as paraphyletic. Although most of the extant genera of Gleicheniaceae originated during the Mesozoic, several genera show Neogene and even Quaternary diversifications, and our results suggest that reticulation and polyploidy may have played significant roles during this diversification. However, some genera, such as Rouxopteris and Stromatopteris, appear to represent evolutionary relicts.

Phytochemical Characterization and In Vitro and In Silico Biological Studies from Ferns of Genus Blechnum (Blechnaceae, Polypodiales).

The genus Blechnum represents one of the most ecologically and therapeutically important groups of ferns that grow in tropical, subtropical and temperate regions. In this work, the chemical fingerprint of lyophilized extracts of Blechnum chilense, B. hastatum, B. magellanicum and B. penna-marina species, the determination of their antioxidant activity through ORAC, FRAP and DPPH assays and inhibition of cholinesterase enzymes (AChE and BChE), and an in silico analysis of selected majority compounds on cholinesterase enzymes were identified. Nineteen compounds were recorded for B. chilense, nine in B. hastatum, seventeen in B. magellanicum and seventeen in B. penna-marina by liquid chromatography coupled with quadrupole-time-of-flight mass spectrometry (UHPLC-ESI-QTOF-MS). The content of phenolic compounds, flavonoids, antioxidant activity and enzyme inhibition were variable among species, with best results for B. penna-marina. Molecular docking evidenced low toxicities, significant pharmacokinetic properties, and significant binding affinities of the tested compounds for the AChE and BChE enzymes. These fern species show high diversity of bioactive compounds and represent a promising resource in phytotherapy, especially for their optimal levels of phenolic compounds that support their antioxidant activity.

Global patterns and climatic determinants of phylogenetic structure of regional fern floras.

Knowledge of relationships between phylogenetic structure of a biological assemblage and ecological factors that drive the variation of phylogenetic structure among regions is crucial for understanding the causes of variation in taxonomic composition and richness among regions, but this knowledge is lacking for the global flora of ferns. Here, we fill this critical knowledge gap. We divided the globe into 392 geographic units on land, collated species lists of ferns for each geographic unit, and used different phylogenetic metrics (tip- vs basal-weighted) reflecting different evolutionary depths to quantify phylogenetic structure. We then related taxonomic and phylogenetic structure metrics to six climatic variables for ferns as a whole and for two groups of ferns (old clades vs polypods) reflecting different evolutionary histories across the globe and within each continental region. We found that when old clades and polypods were considered separately, temperature-related variables explained more variation in these metrics than did precipitation-related variables in both groups. When analyses were conducted for continental regions separately, this pattern holds in most cases. Climate extremes have a stronger relationship with phylogenetic structure of ferns than does climate seasonality. Climatic variables explained more variation in phylogenetic structure at deeper evolutionary depths.

Locked up Inside the Vessels: Rare Earth Elements Are Transferred and Stored in the Conductive Tissues of the Accumulating Fern Dryopteris erythrosora.

Rare earth elements (REEs) are strategic metals strongly involved in low-carbon energy conversion. However, these emerging contaminants are increasingly disseminated into ecosystems, raising concern regarding their toxicity. REE-accumulating plants are crucial subjects to better understand REE transfer to the trophic chain but are also promising phytoremediation tools. In this analysis, we deciphered REE accumulation sites in the REE-accumulating fern Dryopteris erythrosora by synchrotron X-ray µfluorescence (µXRF). This technique allows a high-resolution and in situ analysis of fresh samples or frozen-hydrated cross sections of different organs of the plant. In the sporophyte, REEs were translocated from the roots to the fronds by the xylem sap and were stored within the xylem conductive system. The comparison of REE distribution and accumulation levels in the healthy and necrotic parts of the frond shed light on the differential mobility between light and heavy REEs. Furthermore, the comparison emphasized that necrotized areas were not the main REE-accumulating sites. Finally, the absence of cell-to-cell mobility of REEs in the gametophyte suggested the absence of REE-compatible transporters in photosynthetic tissues. These results provide valuable knowledge on the physiology of REE-accumulating ferns to understand the REE cycle in biological systems and the expansion of phytotechnologies for REE-enriched or REE-contaminated soils.

Comparative assessment of nutritional composition, polyphenol profile and antioxidative properties of wild edible ferns from northeastern China.

Ferns are one of the prevalent species of wild edible plants but one of the least explored terrestrial plants. Therefore, the aim of this study was to assess the nutrient composition, polyphenol profile and antioxidative properties of four wild edible ferns commonly utilized in northeastern China. We studied the content of ash, polysaccharide, protein, fat and mineral elements of the samples. Furthermore, the samples were found to have good total phenolic and total flavonoid contents and some level of antioxidant capacity as determined by 1,1-diphenyl-2-picrylhydrazyl (DPPH), 2,2'-azino-bis(3-ethylbenzothiazoline) 6-sufonic acid (ABTS) and ferric reducing antioxidant power (FRAP). They also exhibited different specific accumulation of polyphenol profiles, estimated by liquid chromatography/mass spectrometry (LC/MS). Significance analysis revealed a significant correlation between individual phenolic compounds and the antioxidant activity of the ferns. The results of the study suggest that wild edible ferns are rich in nutritional value and have potential as a natural source of antioxidants.

Phylogeny, character evolution, and biogeography of the fern genus Bolbitis (Dryopteridaceae).

Bolbitis is a pantropical fern genus of Dryopteridaceae with ca. 80 species mainly in tropical Asia. Earlier studies confirmed the monophyly of Bolbitis when Mickelia is excluded and identified three major clades in Bolbitis. However, earlier studies are based on relatively small sampling and the majority of Asian species are not sampled. In this study, DNA sequences of three plastid markers of 169 accessions representing ca. 68 (85 % of total) species of Bolbitis in nine out of the 10 series recognized by Hennipman (1977), and 54 accessions representing the five remaining bolbitidoid genera are used to infer a global phylogeny with a focus on Asian species. The major results include: (1) Bolbitis is strongly supported as monophyletic; (2) species of Bolbitis are resolved into four major clades and their relationships are: the Malagasy/Mascarene clade is sister to the rest, followed by the African clade which is sister to the American clade + the Asian clade; (3) six well-supported subclades are identified in the most speciose Asian clade; (4) the free-veined Egenolfia is embedded in Bolbitis and is paraphyletic in relation to species with anastomosing venation; (5) three series sensu Hennipman (1977), B. ser. Alienae, B. ser. Egenolfianae, and B. ser. Heteroclitae, are paraphyletic or polyphyletic; (6) evolution of six morphological characters is analyzed and free venation is found to have evolved from anastomosing venation and reversed to free venation in Bolbitis; and (7) biogeographical implications are drawn and it is shown that a single recent dispersal from Asia resulted in continental disjunction of closely related ferns of Bolbitis between Africa and America.

Revisiting ancient polyploidy in leptosporangiate ferns.

Ferns, and particularly homosporous ferns, have long been assumed to have experienced recurrent whole-genome duplication (WGD) events because of their substantially large genome sizes, surprisingly high chromosome numbers, and high degrees of polyploidy among many extant members. As the number of sequenced fern genomes is limited, recent studies have employed transcriptome data to find evidence for WGDs in ferns. However, they have reached conflicting results concerning the occurrence of ancient polyploidy, for instance, in the lineage of leptosporangiate ferns. Because identifying WGDs in a phylogenetic context is the foremost step in studying the contribution of ancient polyploidy to evolution, we here revisited earlier identified WGDs in leptosporangiate ferns, mainly the core leptosporangiate ferns, by building KS -age distributions and applying substitution rate corrections and by conducting statistical gene tree-species tree reconciliation analyses. Our integrative analyses not only identified four ancient WGDs in the sampled core leptosporangiate ferns but also identified false positives and false negatives for WGDs that recent studies have reported earlier. In conclusion, we underscore the significance of substitution rate corrections and uncertainties in gene tree-species tree reconciliations in calling WGD events and advance an exemplar workflow to overcome such often-overlooked issues.

Early land plants: Plentiful but neglected nutritional resources for herbivores?

Plants and herbivores have been engaged in a co-evolutionary arms race for millions of years, during which plants evolved various defenses and other traits to cope with herbivores, whereas herbivores evolved traits to overcome the plants' resistance strategies. Herbivores may also avoid certain plants merely because these lack suitable nutrients for their development. Interestingly, the number of herbivores that attack individual early land plants like mosses and ferns is quite low. Among others, poor nutrient quality has been hypothesized to explain the apparent low herbivory pressure on such plants but still waits for scientific evidences. Here, the nutritive suitability of representative mosses and liverworts (bryophytes) and ferns (pteridophytes) for herbivores was investigated using feeding assays combined with quantifications of nutrients (proteins, amino acids, and sugars). Growth and survival of two polyphagous herbivores, a caterpillar and a snail, were monitored when fed on 15 species of bryophytes and pteridophytes, as well as on maize (Zea mays, angiosperm) used as an external indicative nutritional resource. Overall, our results show that the poor performance of the herbivores on the studied early land plants is not correlated with nutritional quality. The growth and performance of snails and caterpillars fed with these plants were highly variable and independent of nutrient content. These findings arguably dismiss the poor nutrient quality hypothesis as the cause of herbivory deficit in bryophytes and pteridophytes. They suggest the possible presence of early resistance traits that have persisted all through the long evolutionary history of plant-herbivore interactions.

Exploring wild Aspleniaceae ferns as safety sources of polyphenols: The case of Asplenium trichomanes L. and Ceterach officinarum Willd.

The forest ecosystem is a source of material resources used since ancient times by mankind. Ferns are part of different oriental systems of traditional medicine due to the phytochemical variety of their fronds, which have allowed their traditional use to be validated through ethnopharmacological studies. In Europe, different cultures have used the same fern with a wide variety of applications due to its presence in most European forests. In recent years, studies on the phytocharacterization and biological activity of the fronds of the main European ferns have been published. In this study, the presence of polyphenolic phytochemicals has been evaluated by high-performance liquid chromatography-tandem mass spectrometry (HPLC-MS/MS) in the fronds of two wild ferns together with in vitro activities in non-tumoral and human tumoral cell lines. The polyphenols were extracted from Asplenium trichomanes L. and Ceterach officinarum Willd. by cold maceration using methanol. The main phytochemicals of polyphenolic origin in the extracts of A. trichomanes and C. officinarum determined by HPLC-MS/MS were the flavonol hyperoside and the phenolic acid chlorogenic acid, respectively. This different polyphenolic nature of both extracts contributes to the divergence of the behavior experienced in the biological activities tested, but none of the extracts showed a cytotoxic or phototoxic profile in the different tested cell lines. However, the cytoprotective values in front of the H2O2 oxidative stress induced in the 3T3 and HaCaT cell lines position these extracts as possible candidates for future health applications.