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.

Cloud forests figure as one of the most diverse ecosystems, accounting not only for a high number of plant species but also with a great variety of interactions among them. A common interaction in these forests is the one between vascular epiphytes and their hosts. However, few studies have used the network approach to analyze them. Here, we analyze the horizontal and vertical structure of the vascular epiphyte - host network in a cloud forest in central Mexico. We quantified the number of epiphyte stands on each host both total and per-stratum. Complete network, group, and species metrics were estimated at both levels of analysis. The host - epiphyte networks had relatively low network size but were highly connected; moderately nested, with low specialization, and modularity; but higher vulnerability than generality, and high niche overlap. The community was composed by a high number of generalist species. To our knowledge this is the first study in which network analyses are conducted with standardized data and including all host and epiphyte species in the community. The analyses suggest that the networks are robust, and that functional redundancy might be probable, two advantageous characteristics in a very fragmented and threatened cloud forest.

OBJECTIVE: Hybridization is recognized as an important mechanism in fern speciation, with many allopolyploids known among congeners, as well as evidence of ancient genome duplications. Several contemporary instances of deep (intergeneric) hybridization have been noted, invariably resulting in sterile progeny. We chose the christelloid lineage of the family Thelypteridaceae, recognized for its high frequency of both intra- and intergeneric hybrids, to investigate recent hybrid speciation between deeply diverged lineages. We also seek to understand the ecological and evolutionary outcomes of resulting lineages across the landscape.
METHODS: By phasing captured reads within a phylogenomic data set of GoFlag 408 nuclear loci using HybPhaser, we investigated candidate hybrids to identify parental lineages. We estimated divergence ages by inferring a dated phylogeny using fossil calibrations with treePL. We investigated ecological niche conservatism between one confirmed intergeneric allotetraploid and its diploid progenitors using the centroid, overlap, unfilling, and expansion (COUE) framework.
RESULTS: We provide evidence for at least six instances of intergeneric hybrid speciation within the christelloid clade and estimate up to 45 million years of divergence between progenitors. The niche quantification analysis showed moderate niche overlap between an allopolyploid species and its progenitors, with significant divergence from the niche of one progenitor and conservatism to the other.
CONCLUSIONS: The examples provided here highlight the overlooked role that allopolyploidization following intergeneric hybridization may play in fern diversification and range and niche expansions. Applying this approach to other fern taxa may reveal a similar pattern of deep hybridization resulting in highly successful novel lineages.

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.

The latest research shows that ferns and lycophytes have distinct evolutionary lineages. The codon usage patterns of lycophytes and ferns have not yet been documented. To investigate the gene expression profiles across various plant lineages with respect to codon usage, analyze the disparities and determinants of gene evolution in primitive plant species, and identify appropriate exogenous gene expression platforms, the whole-genome sequences of four distinct species were retrieved from the NCBI database. The findings indicated that Ceratopteris richardii, Adiantum capillus-veneris, and Selaginella moellendorffii exhibited an elevated A/U content in their codon base composition and a tendency to end with A/U. Additionally, S. capillus-veneris had more C/G in its codons and a tendency to end with C/G. The ENC values derived from both ENC-plot and ENC-ratio analyses deviated significantly from the standard curves, suggesting that the codon usage preferences of these four species were primarily influenced by genetic mutations and natural selection, with natural selection exerting a more prominent influence. This finding was further supported by PR2-Plot, neutrality plot analysis, and COA. A combination of RSCU and ENC values was used as a reference criterion to rank the codons and further identify the optimal codons. The study identified 24 high-frequency codons in C. richardii, A. capillus-veneris, and Diphasiastrum complanatum, with no shared optimal codons among the four species. Arabidopsis thaliana and Ginkgo biloba exhibited similar codon preferences to the three species, except for S. moellendorffii. This research offers a theoretical framework at the genomic codon level for investigating the phylogenetic relationships between lycophytes and ferns, shedding light on gene codon optimization and its implications for genetic engineering in breeding.

Most land plants alternate between generations of sexual gametophytes and asexual sporophytes. Unlike seed plants, fern gametophytes are free living and grow independently of their sporophytes. In homosporous ferns such as Ceratopteris, gametophytes derived from genetically identical spores exhibit sexual dimorphism, developing as either males or hermaphrodites. Males lack meristems and promote cell differentiation into sperm-producing antheridia. In contrast, hermaphrodites initiate multicellular meristems that stay undifferentiated, sustain cell division and prothallus expansion, and drive the formation of egg-producing archegonia. Once initiating the meristem, hermaphrodites secrete the pheromone antheridiogen, which triggers neighboring slower-growing gametophytes to develop as males, while the hermaphrodites themselves remain insensitive to antheridiogen. This strategy promotes outcrossing and prevents all individuals in the colony from becoming males. This study reveals that an evolutionarily conserved GRAS-domain transcriptional regulator (CrHAM), directly repressed by Ceratopteris microRNA171 (CrmiR171), promotes meristem development in Ceratopteris gametophytes and determines the male-to-hermaphrodite ratio in the colony. CrHAM preferentially accumulates within the meristems of hermaphrodites but is excluded from differentiated antheridia. CrHAM sustains meristem proliferation and cell division through conserved hormone pathways. In the meantime, CrHAM inhibits the antheridiogen-induced conversion of hermaphrodites to males by suppressing the male program expression and preventing meristem cells from differentiating into sperm-producing antheridia. This finding establishes a connection between meristem indeterminacy and sex determination in ferns, suggesting both conserved and diversified roles of meristem regulators in land plants.

The Azolla filiculoides plants were challenged with different arsenic (As) concentration under low pH stress conditions. The growth rate and doubling time of the plants were severely affected by higher As treatments at pH 5.00 when compared with stress pH 4.75 treatments. Hence, pH 5.00 was considered for further studies. In 10-30 μM As treated cultures, after 6 days, the relative growth rate (RGR) of Azolla plants was significantly reduced and in higher concentration of As, the RGR was negatively regulated. The root trait parameters were also significantly affected by increasing concentrations of As. Further, photosynthetic performance indicators also show significant decline with increasing As stress. Overall, the plants treated with 40 and 50 μM of As displayed stress phenotypes like negative RGR, reduced doubling time and root growth, browning of leaves and root withering. The total proline, H2O2, POD and Catalase activities were significantly affected by As treatments. Meantime, 30 μM of As treated cultures displayed 15 μg/g/Fw As accumulation and moderate growth rate. Thus, the Azolla plants are suitable for the phytoremediation of As (up to 30 μM concentration) in the aquatic environment under low pH conditions (5.00). Furthermore, the transcriptome studies on revealed that the importance of positively regulated transporters like ACR3, AceTr family, ABC transporter super family in As (10 μM) stress tolerance, uptake and accumulation. The transporters like CPA1, sugar transporters, PiT were highly down-regulated. Further, expression analysis showed that the MATE1, CIP31, HAC1 and ACR3 were highly altered during the As stress conditions.

There are great concerns for the accumulation in the environment of small dimension plastics, such as micro- and nanoplastics. Due to their small size, which facilitates their uptake by organisms, nanoplastics are of particular concern. The toxic effects of nanoplastics on plants are already reported in the literature, however nothing is known, to date, about the possible effects of climate change, in particular of increasing temperatures, on their toxicity for plants. To address this issue, plants of the water fern Azolla filiculoides were grown at optimal (25 °C) or high (35 °C) temperature, with or without polystyrene nanoplastics, and the effects of these stressors were assessed using a multidisciplinary approach. Green fluorescent polystyrene nanoplastics were used to track their possible uptake by A. filiculoides. The development and physiology of our model plant was adversely affected by both nanoplastics and high temperatures. Overall, histological, morphological, and photosynthetic parameters worsened under co-treatment, in accordance with the increased uptake of nanoplastics under higher temperature, as observed by fluorescence images. Based on our findings, the concern regarding the potential for increased toxicity of pollutants, specifically nanoplastics, at high temperatures is well-founded and warrants attention as a potential negative consequence of climate change. Additionally, there is cause for concern regarding the increase in nanoplastic uptake at high temperatures, particularly if this phenomenon extends to food and feed crops, which could lead to greater entry into the food chain.

OBJECTIVE: Bracken (Pteridium, Dennstaedtiaceae) is a cosmopolitan genus of aggressive disturbance colonizers that are toxic to agricultural livestock. The taxonomy of Pteridium has been treated in multiple schemes, ranging from one to six species worldwide, with numerous subspecies and varieties. Recent work has focused on the worldwide distribution and systematics of the bracken fern, but South America has been poorly represented. We present the first continent-wide sampling and analysis of Pteridium esculentum, a Southern Hemisphere diploid species.
METHODS: Within South America, P. esculentum has several morphotypes, distinguished into subspecies by variation in indument and lamina architecture. We used double digest restriction site-associated DNA sequencing (ddRADSeq) to assess the phylogenetic relationships of P. esculentum subspecies.
RESULTS: We found a striking genetic homogeneity in the species, being able to support only two morphotypes from molecular data: P. e. arachnoideum and P. e. campestre. We had high confidence for shallow and deep phylogenetic relationships, but less support for relationships among crown groups.
CONCLUSIONS: We describe an east-west geographic pattern that would explain the relationships between populations; and, in contrast to previous studies, we detected differences with P. esculentum from Australia. These results will lay the foundations for studying variations in this species\' behavior as a weed, as well as its impact on the production of agricultural livestock in South America.

Monitoring of pesticide concentration distribution across farm fields is crucial to ensure precise and efficient application while preventing overuse or untreated areas. Inspired by nature\'s wettability patterns, we developed a biomimetic fern leaf pesticide collection patch using laser-induced graphene (LIG) alongside an external electrochemical LIG biosensor. This \"collect-and-sense\" system allows for rapid pesticide spray monitoring in the farm field. The LIG is synthesized and patterned on polyimide through a high-throughput gantry-based CO2 laser process, making it amenable to scalable manufacturing. The resulting LIG-based leaf exhibits a remarkable water collection capacity, harvesting spray mist/fog at a rate approximately 11 times greater than a natural ostrich fern leaf when the collection is normalized to surface area. The developed three-electrode LIG pesticide biosensor, featuring a working electrode functionalized with electrodeposited platinum nanoparticles (PtNPs) and the enzyme glycine oxidase, displayed a linear range of 10-260 μM, a detection limit of 1.15 μM, and a sensitivity of 5.64 nA μM-1 for the widely used herbicide glyphosate. Also, a portable potentiostat with a user-friendly interface was developed for remote operation, achieving an accuracy of up to 97%, when compared to a standard commercial benchtop potentiostat. The LIG \"collect-and-sense\" system can consistently collect and monitor glyphosate spray after 24-48 hours of spraying, a time that corresponds to the restricted-entry interval required to enter most farm fields after pesticide spraying. Hence, this innovative \"collect-and-sense\" system not only advances precision agriculture by enabling monitoring and mapping of pesticide distribution but also holds the potential to significantly reduce environmental impact, enhance crop management practices, and contribute to the sustainable and efficient use of agrochemicals in modern agriculture.