Stomatal behaviour and water relations in ferns and lycophytes across habits and habitats.

Stomatal anatomy and behaviour are key to managing gas exchange fluxes, which require coordination with the plant vascular system to adequately supply leaves with water. Stomatal response times and regulation of water loss are generally understudied in ferns, especially across habits (i.e. epiphytic and terrestrial) and habitats (i.e. wet mesic and dry xeric environments). Our objectives were to (i) determine if hydraulic and anatomical traits that control water use are correlated with their habitats (i.e. xeric, mesic) and habits (i.e. epiphytic, terrestrial) for ferns and lycophytes across taxa, and (ii) explore how those traits and others like average leaf water residence time correlate with stomatal function using a subset of closely related species. Epiphytic species had lower vein densities than terrestrial species, while xeric species had higher vein densities than mesic species. Xeric ferns also had smaller stomata than mesic ferns but had similar stomatal densities. Further, in a subset of mesic and xeric ferns, the xeric ferns had higher maximum stomatal conductance and water content, as well as shorter average stomatal opening responses to light intensity, but stomatal closing times did not differ. Finally, shorter stomatal opening and closing responses were correlated with shorter water residence time. Our study highlights anatomical and physiological differences between ferns and lycophytes, which may partially explain habitat preference based on their optimization of light and water.

Conservation applications of niche modeling: Native and naturalized ferns may compete for limited Hawaiian dryland habitat.

Premise: Competition from naturalized species and habitat loss are common threats to native biodiversity and may act synergistically to increase competition for decreasing habitat availability. We use Hawaiian dryland ferns as a model for the interactions between land-use change and competition from naturalized species in determining habitat availability. Methods: We used fine-resolution climatic variables and carefully curated occurrence data from herbaria and community science repositories to estimate the distributions of Hawaiian dryland ferns. We quantified the degree to which naturalized ferns tend to occupy areas suitable for native species and mapped the remaining available habitat given land-use change. Results: Of all native species, Doryopteris angelica had the lowest percentage of occurrences of naturalized species in its suitable area while D. decora had the highest. However, all Doryopteris spp. had a higher percentage overlap, while Pellaea ternifolia had a lower percentage overlap, than expected by chance. Doryopteris decora and D. decipiens had the lowest proportions (<20%) of suitable area covering native habitat. Discussion: Areas characterized by shared environmental preferences of native and naturalized ferns may decrease due to human development and fallowed agricultural lands. Our study demonstrates the value of place-based application of a recently developed correlative ecological niche modeling approach for conservation risk assessment in a rapidly changing and urbanized island ecosystem.

SEM and LM insights: Spore morphology and its taxonomic significance in Sino Himalayan, Malesian, and European elements of Pteridaceae.

The Pteridaceae family, known for its taxonomic complexity, presents challenges in identification due to high variability among its species. This study investigates the spore morphology employing both SEM and LM techniques in 10 Pteridaceae taxa phytogeographicaly Sino-Himalayan, Malesian, and European elements in Pakistan. The taxa include Adiantum capillus-veneris, A. incisum, A. venustum, Aleuritopteris bicolor, Oeosporangium nitidulum, O. pteridioides, Onychium cryptogrammoides, O. vermae, Pteris cretica, and P. vittata. The objective is to assess their taxonomic relevance and develop a spore-based taxonomic key. Findings indicate differences in spore shape, sizes, exospore thickness, and in surface ornamentation highlighting the potential for taxonomic differentiation. Spores are trilete, and notable differences are observed in the dimension of spores in both distal and proximal sides. Equatorial dimensions vary between 35 and 50 µm, while the polar diameter ranges from 29 to 50 µm. SEM revealed different spore ornamentation types that show several useful characteristics establishing valuable taxonomic variations. The studied Adiantum taxa feature a perispore with tubercules and a micro-granulose surface. The spores of examined Oeosporangium and Aleuritopteris taxa shows cristate sculptures with variable ornamentations. Both species of Onychium have tuberculate-pleated tubercles with sinuous folds on both distal and proximal sides. The surface ornamentation among examined Pteris taxa show variability. PCA analysis indicated that spore quantitative data identified distinct groups, underscoring taxonomic significance. Nevertheless, there was variation observed in surface ornamentation and spore shape, indicating the potential for discrimination among taxa. RESEARCH HIGHLIGHTS: Spore morphology of 10 Pteridaceae taxa has been investigated through LM and SEM. Investigated species shows differences in spore shape, sizes, exospore thickness, and in surface ornamentation. Ornamentation on the perispore provides several valuable characteristics, establishing useful taxonomic distinctions. Spore morphological analysis is effective at the generic level, with minor distinctions discernible at the species level.

The size diversity of the Pteridaceae family chloroplast genome is caused by overlong intergenic spacers.

BACKGROUND: While the size of chloroplast genomes (cpDNAs) is often influenced by the expansion and contraction of inverted repeat regions and the enrichment of repeats, it is the intergenic spacers (IGSs) that appear to play a pivotal role in determining the size of Pteridaceae cpDNAs. This provides an opportunity to delve into the evolution of chloroplast genomic structures of the Pteridaceae family. This study added five Pteridaceae species, comparing them with 36 published counterparts. RESULTS: Poor alignment in the non-coding regions of the Pteridaceae family was observed, and this was attributed to the widespread presence of overlong IGSs in Pteridaceae cpDNAs. These overlong IGSs were identified as a major factor influencing variations in cpDNA size. In comparison to non-expanded IGSs, overlong IGSs exhibited significantly higher GC content and were rich in repetitive sequences. Species divergence time estimations suggest that these overlong IGSs may have already existed during the early radiation of the Pteridaceae family. CONCLUSIONS: This study reveals new insights into the genetic variation, evolutionary history, and dynamic changes in the cpDNA structure of the Pteridaceae family, providing a fundamental resource for further exploring its evolutionary research.

Three new pterosins from Pteris semipinnata.

Three new pterosins, named as semipterosin A (1), B (2) and C (3), together with 11 known pterosins (4-14), were isolated from the aerial parts of Pteris semipinnata. Their structures were elucidated by HRESI-MS, NMR spectral data, CD and literature comparisons. Three new pterosins were assessed for their anti-inflammatory activity. Compounds 1-3 inhibited the NF-kB induction by 40.7%, 61.9% and 34.0%, respectively. This is the first report of the isolation of compounds 6-14 from this plant.

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.

Adaptive evolution and co-evolution of chloroplast genomes in Pteridaceae species occupying different habitats: overlapping residues are always highly mutated.

BACKGROUND: The evolution of protein residues depends on the mutation rates of their encoding nucleotides, but it may also be affected by co-evolution with other residues. Chloroplasts function as environmental sensors, transforming fluctuating environmental signals into different physiological responses. We reasoned that habitat diversity may affect their rate and mode of evolution, which might be evidenced in the chloroplast genome. The Pteridaceae family of ferns occupy an unusually broad range of ecological niches, which provides an ideal system for analysis. RESULTS: We conducted adaptive evolution and intra-molecular co-evolution analyses of Pteridaceae chloroplast DNAs (cpDNAs). The results indicate that the residues undergoing adaptive evolution and co-evolution were mostly independent, with only a few residues being simultaneously involved in both processes, and these overlapping residues tend to exhibit high mutations. Additionally, our data showed that Pteridaceae chloroplast genes are under purifying selection. Regardless of whether we grouped species by lineage (which corresponded with ecological niches), we determined that positively selected residues mainly target photosynthetic genes. CONCLUSIONS: Our work provides evidence for the adaptive evolution of Pteridaceae cpDNAs, especially photosynthetic genes, to different habitats and sheds light on the adaptive evolution and co-evolution of proteins.

Structure characterization of novel heteropolysaccharides from Pteridium revolutum with antioxidant and antiglycated activities.

This study aims to analysis the structures of polysaccharides isolated from Pteridium revolutum and their antioxidant and antiglycated activities. Three novel water-soluble heteropolysaccharides, named PRP0, PRP1, and PRP2, were isolated from P. revolutum. The average molecular weight was determined by high performance gel permeation chromatography analysis as 1.04 × 106, 8.39 × 105, and 7.37 × 105 Da, respectively. Their structures were characterized using physicochemical and spectroscopic methods. The antioxidant and antiglycated activities were assayed in vitro. PRP0, PRP1, and PRP2 consist of l-Ara, l-Rha, d-Man, d-Xyl, d-Fuc, d-Gal, and d-Glc in different proportions. PRP1 mainly has a backbone of (1 â†’ 3,6)-linked d-Man and (1 â†’ 3)-linked d-Gal on main chain. PRP2 is mainly composed of (1 â†’ 2,4)-linked d-Man and (1 â†’ 3)-linked d-Gal on main chain. All polysaccharides have strong scavenging power on 2,2-difenil-1-picril-hidrazil and hydroxyl radicals and significantly antiglycated activity in Bovine serum albumin-Glucose model, which showing that the polysaccharides have potential application value on the functional food.

The complete chloroplast genome sequence of the water fern Ceratopteris thalictroides (Pteridaceae).

This work determined and analyzed the complete chloroplast genome sequence of Ceratopteris thalictroides (Linnaeus) Brongniart 1822 (Pteridaceae). The results indicate that the total chloroplast genome size of C. thalictroides is 149,399 bp in length, and the genome contains a large single-copy (LSC) region of 83,580 bp, a small single-copy (SSC) region of 21,241 bp, and a pair of inverted repeat (IR) regions of 22,289 bp. The GC content of C. thalictroides is 36.7%. The genome encodes a total of 131 unique genes, including 82 protein-coding genes, 38 tRNA genes, and 8 rRNA genes. The phylogenetic analysis results strongly suggest that C. thalictroides is closely related to C. cornuta.

Structure Characterization, Antioxidant and Immunomodulatory Activities of Polysaccharide from Pteridium aquilinum (L.) Kuhn.

Pteridium aquilinum (L.) Kuhn (Pteridaceae family) has been widely used as a food and medicine in China and Korea. Previous studies indicate that P. aquilinum contains a variety of bioactive chemical components such as flavonoids, phenols, terpenoids, saponins, polysaccharides, and so on. In the present study, a novel polysaccharide (named as PAP-3) with average molecular weight of 2.14 × 105 Da was obtained from P. aquilinum. The structure was studied through physicochemical and spectroscopic analysis. The results indicated that PAP-3 consists of arabinose, rhamnose, fucose, galactose, mannose, and xylose in a molar ratio of 1.58:1.00:3.26:4.57:4.81:3.33. The polysaccharide is mainly composed of (1→2)-linked xylose and (1→3,6)-linked mannose on the main chain, with (1→2)-linked xylose, (1→6)-linked mannose, and (1→6)- and (1→3,6)-linked galactose as side chains. Galactose, fucose, and xylose are located at the end of the side chains. The in vitro immunomodulatory and antioxidant activities were assayed. PAP-3 has strong free-radical scavenging activity on DPPH and ABTS radicals and significant immunomodulatory activity on RAW264.7 cells. These data provide useful information for further study on the polysaccharides of P. aquilinum and their applications in the food and medical industries.

Cytochalasans from the endophytic fungus Diaporthe ueckerae associated with the fern Pteris vittata.

Six previously undescribed cytochalasans, ueckerchalasins A-E and 4'-hydroxycytochalasin J3, together with eight known congeners, were isolated from solid cultures of the endophytic fungus Diaporthe ueckerae SC-J0123 which was originally isolated from the leaves of Pteris vittata L. Their structures were elucidated by extensive spectroscopic analysis, single-crystal X-ray diffraction, and theoretical simulations of ECD spectra and 13C NMR shifts. Ueckerchalasins A-C have a carbon-carbon bridge between C-14 and C-20, forming a rare 5/6/6/7-fused heterocyclic core. Ueckerchalasins C and D displayed selective activity against human carcinoma HeLa and HepG2 cells. Ueckerchalasins C was also active against Staphylococcus aureus and methicillin-resistant S. aureus (MRSA).

Once more unto the breach, dear friends: Resolving the origins and relationships of the Pellaea wrightiana hybrid complex.

PREMISE: The taxonomic status of Wright's cliff brake fern, Pellaea wrightiana, has been in dispute ever since it was first described by Hooker in 1858. Previously published evidence suggested that this "taxon" may represent a polyploid complex rather than a single discrete species, a hypothesis tested here using a multifaceted analytical approach. METHODS: Data derived from cytogenetics, spore analyses, leaf morphometrics, enzyme electrophoresis, and phylogenetic analyses of plastid and nuclear DNA sequences are used to elucidate the origin, relationships, and taxonomic circumscription of P. wrightiana. RESULTS: Plants traditionally assigned to this taxon represent three distinct polyploids. The most widespread, P. wrightiana, is a fertile allotetraploid that arose through hybridization between two divergent diploid species, P. truncata and P. ternifolia. Sterile triploids commonly identified as P. wrightiana, were found to be backcross hybrids between this fertile tetraploid and diploid P. truncata. Relatively common across Arizona and New Mexico, they are here assigned to P. ×wagneri hyb. nov. In addition, occasional sterile tetraploid plants assigned to P. wrightiana are shown here to be hybrids between the fertile allotetraploid and the tetraploid P. ternifolia subsp. arizonica. These tetraploid hybrids originated independently in two regions of parental sympatry (southern Arizona and west Texas) and are here assigned to P. ×gooddingii hyb. nov. CONCLUSIONS: Weaving together data from a diversity of taxonomic approaches, we show that plants identified as P. wrightiana represent three morphologically distinguishable polyploids that have arisen through repeated hybridization events involving the divergent sexual taxa P. ternifolia and P. truncata.

Ptercresions A-C: three new terpenes with hepatoprotective activity from Pteris cretica L.

Three new terpenes were isolated from Pteris cretica L., namely ptercresions A (1), B (2) and C (3), together with two known terpenes and one terpene glycoside respectively identified as callisalignene D (4), berberiside A (5), creoside I (6). The structures of isolates were determined by HR-ESI-MS, ECD, NMR and acid hydrolysate analysis. The in vitro hepatoprotective activities of compounds 1-6 were studied against paracetamol-induced L-02 hepatocyte injury. The results revealed that compounds 2-4 showed moderate anti hepatocyte injury activity.

Spores morphology of the family Pteridaceae from Shandong Province, China.

In this article, we explored systematically the spore morphology of Pteridaceae by observation of the species distributed in Shandong Province using scanning electron microscopy (SEM). The results showed that the spore morphology of all the species in the family is tetrahedral and trilete. The characters of spore ornamentation are intraspecies stable, but significantly different among species and genera. Spore morphology is significant in exploring the phylogenetic relationships of Pteridaceae as well as in generic and specific delimitations.

The complete chloroplast genome of a distinctive fern, Coniogramme intermedia Hieron. (Pteridaceae).

Coniogramme intermedia Hieron. is a morphologically distinctive species in the genus. It is identified by lanceolate pinnules with serrated margins, free veins, hydathodes extending into teeth, and laminae abaxially hairy. It is mainly distributed in the tropical and subtropical regions of Asia. Herein, we report the first complete chloroplast genome sequence of C. intermedia. Also, it is the opening one of the genus Coniogramme Fée. The chloroplast genome sequence is 153,561 bp in length. The genome has a typical quadripartite structure, including a large single-copy (LSC) region of 82,817 bp, a small single-copy (SSC) region of 21,236 bp, and two inverted repeat (IR) regions of 24,754bp each. The total GC content is 45.0%. The complete plastome sequence contains 114 genes, including, 81 protein-coding, 29 tRNA, and four rRNA genes. The phylogenetic analysis of Pteridaceae based on the complete chloroplast genomes was also presented in this study.

When it only takes one to tango: assessing the impact of apomixis in the fern genus Pteris.

PREMISE: Apomixis (asexual reproduction by seed, spore, or egg) has evolved repeatedly across the tree of life. Studies of animals and angiosperms show that apomictic lineages are often evolutionarily short-lived and frequently exhibit different distributions than their sexual relatives. However, apomixis is rare in these groups. Less is known about the role of apomixis in the evolution and biogeography of ferns, in which ~10% of species are apomictic. Apomixis is especially common in the fern genus Pteris (34-39% of species); however, because of the limited taxonomic and geographic sampling of previous studies, the true frequency of apomixis and its associations with geography and phylogeny in this lineage remain unclear. METHODS: We used spore analyses of herbarium specimens to determine reproductive mode for 127 previously unsampled Pteris species. Then we leveraged biogeographic and phylogenetic analyses to estimate the global distribution and evolution of apomixis in Pteris. RESULTS: Among all Pteris species examined, we found that 21% are exclusively apomictic, 71% are exclusively sexual, and 8% have conflicting reports. Apomixis is unevenly distributed across the range of the genus, with the Paleotropics exhibiting the highest frequency, and has evolved numerous times across the Pteris phylogeny, with predominantly East Asian and South Asian clades containing the most apomictic species. CONCLUSIONS: Apomixis arises frequently in Pteris, but apomictic species do not appear to diversify. Species that encompass both apomictic and sexual populations have wider ranges than exclusively sexual or apomictic species, which suggests that sexual and apomictic ferns could occupy separate ecological niches.

Arsenic Toxicity-Induced Physiological and Metabolic Changes in the Shoots of Pteris cretica and Spinacia oleracea.

Arsenic is a ubiquitous toxic element that can be accumulated into plant parts. The present study investigated the response of Pteris cretica and Spinacia oleracea to As treatment through the analysis of selected physiological and metabolic parameters. Plants were grown in pots in As(V) spiked soil (20 and 100 mg/kg). Plants' physiological condition was estimated through the determination of elements, gas-exchange parameters, chlorophyll fluorescence, water potential, photosynthetic pigments, and free amino acid content. The results confirmed differing As accumulation in plants, as well as in shoots and roots, which indicated that P. cretica is an As-hyperaccumulator and that S. oleracea is an As-root excluder. Variations in physiological and metabolic parameters were observed among As treatments. Overall, the results revealed a significant effect of 100 mg/kg As treatment on the analysed parameters. In both plants, this treatment affected growth, N, Mg, S, Mn, and Zn content, as well as net photosynthetic rate, chlorophyll fluorescence, and total free amino acid content. In conclusion, the results reflect the similarity between P. cretica and S. oleracea in some aspects of plants' response to As treatment, while physiological and metabolic parameter changes related to As treatments indicate the higher sensitivity of S. oleracea.

Taxonomy and spore morphology of selected taxa of Cheilanthoideae and Pteridoideae (Pteridaceae) from Pakistan.

Taxonomy and spore morphology of 12 taxa of Cheilanthoideae and Pteridoideae (Pteridaceae, Polypodiales) from Pakistan is illustrated with scanning electron microscopy images based upon the specimens collected from various localities. A total of six genera belong to 12 taxa viz. Actiniopteris radiata, Aleuritopteris albomarginata, A. ancepes, Notholaena himalaica, Oeosporangium nitidulum, O. pteridioides subsp. acrosticum, Onychium cryptogrammoides subsp. cryptogrammoides, O. vermae, Pteris cretica subsp. cretica, P. cretica subsp. laeta, P. vittata subsp. emodi, and P. vittata subsp. vittata were reported. Spore morphology of the taxa was trilete, triangular in proximal and distal view, ellipsoidal and hemicircular in equatorial view, polar proximal and distal surface with cristate, granulose, reticulate, perforate and tuberculate ornamentation.

Variation in frequency of plastid RNA editing within Adiantum implies rapid evolution in fern plastomes.

PREMISE: Recent studies of plant RNA editing have demonstrated that the number of editing sites can vary widely among large taxonomic groups (orders, families). Yet, very little is known about intrageneric variation in frequency of plant RNA editing, and no study has been conducted in ferns. METHODS: We determined plastid RNA-editing counts for two species of Adiantum (Pteridaceae), A. shastense and A. aleuticum, by implementing a pipeline that integrated read-mapping and SNP-calling software to identify RNA-editing sites. We then compared the edits found in A. aleuticum and A. shastense with previously published edits from A. capillus-veneris by generating alignments for each plastid gene. RESULTS: We found direct evidence for 505 plastid RNA-editing sites in A. aleuticum and 509 in A. shastense, compared with 350 sites in A. capillus-veneris. We observed striking variation in the number and location of the RNA-editing sites among the three species, with reverse (U-to-C) editing sites showing a higher degree of conservation than forward (C-to-U) sites. Additionally, sites involving start and stop codons were highly conserved. CONCLUSIONS: Variation in the frequency of RNA editing within Adiantum implies that RNA-editing sites can be rapidly gained or lost throughout evolution. However, varying degrees of conservation between both C-to-U and U-to-C sites and sites in start or stop codons, versus other codons, hints at the likely independent origin of both types of edits and a potential selective advantage conferred by RNA editing.

Taxonomy and spore morphology of genus Adiantum (Vittarioideae; Pteridaceae) from Pakistan.

Taxonomy and spore morphology of genus Adiantum (Vittarioideae; Pteridaceae) from Pakistan is illustrated with scanning electron microscopy images based upon the specimens examined in herbaria of Pakistan, United Kingdom, and United States. A total of five species viz. Adiantum capillus-veneris, A. incisum subsp. incisum, A. pedatum subsp. pedatum, A. tibeticum, and A. venustum were reported with the habitat ecology of the species from Pakistan. Spore morphology of the taxa was trilete, triangular in proximal and distal view, ellipsoidal and hemicircular in equatorial view, polar proximal and distal surface was with tuberculate and granulose ornamentation.