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Clint Chapple - One of the best experts on this subject based on the ideXlab platform.
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independent recruitment of an o methyltransferase for syringyl lignin biosynthesis in Selaginella moellendorffii
The Plant Cell, 2011Co-Authors: Jingke Weng, John Ralph, Takuya Akiyama, Clint ChappleAbstract:Syringyl lignin, an important component of the secondary cell wall, has traditionally been considered to be a hallmark of angiosperms because ferns and gymnosperms in general lack lignin of this type. Interestingly, syringyl lignin was also detected in Selaginella, a genus that represents an extant lineage of the most basal of the vascular plants, the lycophytes. In angiosperms, syringyl lignin biosynthesis requires the activity of ferulate 5-hydroxylase (F5H), a cytochrome P450-dependent monooxygenase, and caffeic acid/5-hydroxyferulic acid O-methyltransferase (COMT). Together, these two enzymes divert metabolic flux from the biosynthesis of guaiacyl lignin, a lignin type common to all vascular plants, toward syringyl lignin. Selaginella has independently evolved an alternative lignin biosynthetic pathway in which syringyl subunits are directly derived from the precursors of p-hydroxyphenyl lignin, through the action of a dual specificity phenylpropanoid meta-hydroxylase, Sm F5H. Here, we report the characterization of an O-methyltransferase from Selaginella moellendorffii, COMT, the coding sequence of which is clustered together with F5H at the adjacent genomic locus. COMT is a bifunctional phenylpropanoid O-methyltransferase that can methylate phenylpropanoid meta-hydroxyls at both the 3- and 5-position and function in concert with F5H in syringyl lignin biosynthesis in S. moellendorffii. Phylogenetic analysis reveals that Sm COMT, like F5H, evolved independently from its angiosperm counterparts.
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independent origins of syringyl lignin in vascular plants
Proceedings of the National Academy of Sciences of the United States of America, 2008Co-Authors: Jingke Weng, Jake Stout, Xu Li, Clint ChappleAbstract:Lycophytes arose in the early Silurian (≈400 Mya) and represent a major lineage of vascular plants that has evolved in parallel with the ferns, gymnosperms, and angiosperms. A hallmark of vascular plants is the presence of the phenolic lignin heteropolymer in xylem and other sclerified cell types. Although syringyl lignin is often considered to be restricted in angiosperms, it has been detected in lycophytes as well. Here we report the characterization of a cytochrome P450-dependent monooxygenase from the lycophyte Selaginella moellendorffii. Gene expression data, cross-species complementation experiments, and in vitro enzyme assays indicate that this P450 is a ferulic acid/coniferaldehyde/coniferyl alcohol 5-hydroxylase (F5H), and is capable of diverting guaiacyl-substituted intermediates into syringyl lignin biosynthesis. Phylogenetic analysis indicates that the Selaginella F5H represents a new family of plant P450s and suggests that it has evolved independently of angiosperm F5Hs.
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Parallels in lignin biosynthesis: A study in Selaginella moellendorffii reveals convergence across 400 million years of evolution.
Communicative & Integrative Biology, 2008Co-Authors: Jingke Weng, Jo Ann Banks, Clint ChappleAbstract:A hallmark of vascular plants is the development of a complex water-conducting system, which is physically reinforced by the heterogeneous aromatic polymer lignin. Syringyl lignin, a major building block of lignin, is often thought to be uniquely characteristic of angiosperms; however, it was demonstrated over fifty years ago that that syringyl lignin is found in another group of plants, known as the lycophytes, the ancestors of which diverged from all the other vascular plant lineages 400 million years ago.1 To determine the biochemical basis for this common biosynthetic ability, we isolated and characterized cytochrome P450-dependent monooxygenases (P450s) from the lycophyte Selaginella moellendorffii and compared them to the enzyme that is required for syringyl lignin synthesis in angiosperms. Our results showed that one of these P450s encodes an enzyme that is functionally analogous to but phylogenetically independent from its angiosperm counterpart. Here, we discuss the evolution of lignin biosynthes...
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Functional analysis and comparative genomics of expressed sequence tags from the lycophyte Selaginella moellendorffii
BMC Genomics, 2005Co-Authors: Jingke Weng, Milos Tanurdzic, Clint ChappleAbstract:Background The lycophyte Selaginella moellendorffii is a member of one of the oldest lineages of vascular plants on Earth. Fossil records show that the lycophyte clade arose 400 million years ago, 150–200 million years earlier than angiosperms, a group of plants that includes the well-studied flowering plant Arabidopsis thaliana. S. moellendorffii has a genome size of approximately 100 Mbp, as small or smaller than that of A. thaliana. S. moellendorffii has the potential to provide significant comparative information to better understand the evolution of vascular plants.
Jingke Weng - One of the best experts on this subject based on the ideXlab platform.
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independent recruitment of an o methyltransferase for syringyl lignin biosynthesis in Selaginella moellendorffii
The Plant Cell, 2011Co-Authors: Jingke Weng, John Ralph, Takuya Akiyama, Clint ChappleAbstract:Syringyl lignin, an important component of the secondary cell wall, has traditionally been considered to be a hallmark of angiosperms because ferns and gymnosperms in general lack lignin of this type. Interestingly, syringyl lignin was also detected in Selaginella, a genus that represents an extant lineage of the most basal of the vascular plants, the lycophytes. In angiosperms, syringyl lignin biosynthesis requires the activity of ferulate 5-hydroxylase (F5H), a cytochrome P450-dependent monooxygenase, and caffeic acid/5-hydroxyferulic acid O-methyltransferase (COMT). Together, these two enzymes divert metabolic flux from the biosynthesis of guaiacyl lignin, a lignin type common to all vascular plants, toward syringyl lignin. Selaginella has independently evolved an alternative lignin biosynthetic pathway in which syringyl subunits are directly derived from the precursors of p-hydroxyphenyl lignin, through the action of a dual specificity phenylpropanoid meta-hydroxylase, Sm F5H. Here, we report the characterization of an O-methyltransferase from Selaginella moellendorffii, COMT, the coding sequence of which is clustered together with F5H at the adjacent genomic locus. COMT is a bifunctional phenylpropanoid O-methyltransferase that can methylate phenylpropanoid meta-hydroxyls at both the 3- and 5-position and function in concert with F5H in syringyl lignin biosynthesis in S. moellendorffii. Phylogenetic analysis reveals that Sm COMT, like F5H, evolved independently from its angiosperm counterparts.
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independent origins of syringyl lignin in vascular plants
Proceedings of the National Academy of Sciences of the United States of America, 2008Co-Authors: Jingke Weng, Jake Stout, Xu Li, Clint ChappleAbstract:Lycophytes arose in the early Silurian (≈400 Mya) and represent a major lineage of vascular plants that has evolved in parallel with the ferns, gymnosperms, and angiosperms. A hallmark of vascular plants is the presence of the phenolic lignin heteropolymer in xylem and other sclerified cell types. Although syringyl lignin is often considered to be restricted in angiosperms, it has been detected in lycophytes as well. Here we report the characterization of a cytochrome P450-dependent monooxygenase from the lycophyte Selaginella moellendorffii. Gene expression data, cross-species complementation experiments, and in vitro enzyme assays indicate that this P450 is a ferulic acid/coniferaldehyde/coniferyl alcohol 5-hydroxylase (F5H), and is capable of diverting guaiacyl-substituted intermediates into syringyl lignin biosynthesis. Phylogenetic analysis indicates that the Selaginella F5H represents a new family of plant P450s and suggests that it has evolved independently of angiosperm F5Hs.
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Parallels in lignin biosynthesis: A study in Selaginella moellendorffii reveals convergence across 400 million years of evolution.
Communicative & Integrative Biology, 2008Co-Authors: Jingke Weng, Jo Ann Banks, Clint ChappleAbstract:A hallmark of vascular plants is the development of a complex water-conducting system, which is physically reinforced by the heterogeneous aromatic polymer lignin. Syringyl lignin, a major building block of lignin, is often thought to be uniquely characteristic of angiosperms; however, it was demonstrated over fifty years ago that that syringyl lignin is found in another group of plants, known as the lycophytes, the ancestors of which diverged from all the other vascular plant lineages 400 million years ago.1 To determine the biochemical basis for this common biosynthetic ability, we isolated and characterized cytochrome P450-dependent monooxygenases (P450s) from the lycophyte Selaginella moellendorffii and compared them to the enzyme that is required for syringyl lignin synthesis in angiosperms. Our results showed that one of these P450s encodes an enzyme that is functionally analogous to but phylogenetically independent from its angiosperm counterpart. Here, we discuss the evolution of lignin biosynthes...
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Functional analysis and comparative genomics of expressed sequence tags from the lycophyte Selaginella moellendorffii
BMC Genomics, 2005Co-Authors: Jingke Weng, Milos Tanurdzic, Clint ChappleAbstract:Background The lycophyte Selaginella moellendorffii is a member of one of the oldest lineages of vascular plants on Earth. Fossil records show that the lycophyte clade arose 400 million years ago, 150–200 million years earlier than angiosperms, a group of plants that includes the well-studied flowering plant Arabidopsis thaliana. S. moellendorffii has a genome size of approximately 100 Mbp, as small or smaller than that of A. thaliana. S. moellendorffii has the potential to provide significant comparative information to better understand the evolution of vascular plants.
Ke Li Chen - One of the best experts on this subject based on the ideXlab platform.
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a purified biflavonoid extract from Selaginella moellendorffii alleviates gout arthritis via nlrp3 asc caspase 1 axis suppression
Frontiers in Pharmacology, 2021Co-Authors: Xue-yan Zhang, Guang-rui Deng, Yingbo Liu, Bisheng Huang, Guoyin Kai, Ke Li ChenAbstract:Background: Activation of nucleotide oligomerization domain-like receptor protein 3 (NLRP3) inflammasome plays a crucial role in gout. Selaginella moellendorffii has been confirmed effective for the treatment of gout in hospital preparations. Flavonoids, such as amentoflavone (AM), are the main active components of this medicine. Purpose: We aimed to investigate the flavonoid extract (TF) and AM's effects on NLRP3 inflammasome in vitro and their preventive effects on gout in vivo. Methods: LC-MS method was employed to investigate the chemical profile of TF. The cellular inflammation model was established by lipopolysaccharide (LPS) or monosodium urate (MSU) stimulation. The cell membrane integrality and morphological characteristics were determined by using Lactate dehydrogenase (LDH) assay kits, propidium iodide (PI) stain, and scanning electron microscopy (SEM). The inflammatory cytokines and NLRP3 inflammasome activation were determined using enzyme-linked immunosorbent assay (ELISA), quantitative real-time polymerase chain reaction (RT-PCR), immunofluorescence staining, and western blotting. The acute gout mouse model was induced by MSU injection into footpads, and then the paw edema, inflammatory mediators, and histological examination (HE) were analyzed. Results: The main constituents in TF are AM and robustaflavone. In the cellular inflammation model, TF down-regulated the levels of nitric oxide (NO), TNF-α, and LDH, suppressed NLRP3 inflammasome-derived interleukin-1β (IL-1β) secretion, decreased caspase-1 activation, repressed mature IL-1β expression, inhibited ASC speck formation and NLRP3 protein expression. In an acute gout mouse model, oral administration of TF to mice effectively alleviated paw edema, reduced inflammatory features, and decreased the levels of IL-1β in mouse foot tissue. Similarly, the characteristic constituent AM was also able to down-regulated the levels of NO, TNF-α, and LDH, down-regulate the mRNA expression of IL-1β, TNF-α, caspase-1, and NLRP3. Besides, the foot thickness, lymphocyte infiltration, and IL-1β level were also prevented by AM. Conclusion: The results indicated that TF and its main constituent AM alleviate gout arthritis via NLRP3/ASC/Caspase-1 axis suppression.
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screening the best compatibility of Selaginella moellendorffii prescription on hyperuricemia and gouty arthritis and its mechanism
Evidence-based Complementary and Alternative Medicine, 2019Co-Authors: Xue-yan Zhang, Ping Zhao, Jing Cheng, Ke Li ChenAbstract:Objectives. The Selaginella moellendorffii prescription (SMP) consists of S. moellendorffii Herba (SM), Smilacis glabrae Rhizoma (SGR), and Plantaginis Semen (PS). It has been commonly used to treat hyperuricemia and acute gouty arthritis as a hospital preparation. This study was aimed at investigating the best compatibility ratio of SMP on hyperuricemia and gouty arthritis and getting better insight of the possible mechanism. Methods. In vitro, anti-inflammatory activity of SMP was evaluated by lipopolysaccharide (LPS) induced RAW264.7 cells. The release of nitric oxide (NO) was screened by Griess assay, and NF-κB p65 and NLRP3 proteins expression was examined by immunofluorescence assay. Then, the levels of creatinine (Cr), blood urea nitrogen (BUN), and uric acid (UA) were detected in mice induced by potassium oxonate, and the paw oedema, inflammatory mediators, and histological examination were analyzed in rats induced by monosodium urate (MSU). HPLC method was employed to investigate the chemical profile of this preparation. Results. In vitro, SMP-3 (the ratio of SMP:SGR:PS was 3:1:1) exhibited the most potent anti-NO production activity without obvious toxicity. This anti-inflammatory effect was associated with suppression of NF-κB p65 nuclear translocation and NLRP3 protein expression. In animal experiments, the levels of BUN and Cr in SMP-3 group were lower than other extract groups, and the level of UA was also remarkably decreased by SMP-3 in hyperuricemic mice (P<0.01). Besides, SMP-3 extract was able to prevent the paw edema, reduce gouty joint inflammatory features, and decrease the levels IL-1β, PGE-2, IL-8, and NO in gouty arthritis rats. Furthermore, 6-C-β-D-xylopyranosyl-8-C-β-D-glucopyranosyl, apigenin, and astilbin were identified from SMP-3 extract. Conclusions. In summary, SMP-3 may be a potential therapeutic agent for the prevention of hyperuricemic and gout.
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corrigendum to pharmacological basis for use of Selaginella moellendorffii in gouty arthritis antihyperuricemic anti inflammatory and xanthine oxidase inhibition
Evidence-based Complementary and Alternative Medicine, 2017Co-Authors: Ping Zhao, Ke Li Chen, Guo-li Zhang, Guang-rui DengAbstract:This study was aimed at evaluating the effects of Selaginella moellendorffii Hieron. (SM) on gouty arthritis and getting an insight of the possible mechanisms. HPLC method was developed for chemical analysis. The paw oedema, the neutrophil accumulation, inflammatory mediators, lipid peroxidation, and histopathological changes of the joints were analyzed in gouty arthritis rat model, and the kidney injury and serum urate were detected in hyperuricemic mice. Pharmacokinetic result demonstrated that the main apigenin glycosides might be quantitatively transformed into apigenin in the mammalian body. Among these compounds, the apigenin exhibited the strongest effect on xanthine oxidase (XOD). SM aqueous extract has proved to be active in reducing hyperuricemia in dose-dependent manner, and the levels of blood urea nitrogen (BUN) and creatinine (Cr) in high dose group were decreased significantly as compared with hyperuricemic control group (P < 0.01). The high dose of SM extract could significantly prevent the paw swelling, reduce gouty joint inflammatory features, reduce the release of IL-1β and TNF-α, lower malondialdehyde (MDA) and myeloperoxidase (MPO) levels, and increase superoxide dismutase (SOD) level (P < 0.01). For the first time, this study provides a rational basis for the traditional use of SM aqueous extract against gout in folk medicine.
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Pharmacological Basis for Use of Selaginella moellendorffii in Gouty Arthritis: Antihyperuricemic, Anti-Inflammatory, and Xanthine Oxidase Inhibition
Hindawi Limited, 2017Co-Authors: Ping Zhao, Ke Li Chen, Guo-li Zhang, Guang-rui DengAbstract:This study was aimed at evaluating the effects of Selaginella moellendorffii Hieron. (SM) on gouty arthritis and getting an insight of the possible mechanisms. HPLC method was developed for chemical analysis. The paw oedema, the neutrophil accumulation, inflammatory mediators, lipid peroxidation, and histopathological changes of the joints were analyzed in gouty arthritis rat model, and the kidney injury and serum urate were detected in hyperuricemic mice. Pharmacokinetic result demonstrated that the main apigenin glycosides were almost quantitatively transformed into apigenin in the mammalian body. Among these compounds, the apigenin exhibited the strongest effect on xanthine oxidase (XOD). SM aqueous extract has proved to be active in reducing hyperuricemia in dose-dependent manner, and the levels of blood urea nitrogen (BUN) and creatinine (Cr) in high dose group were decreased significantly as compared with hyperuricemic control group (P
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Antiviral Activity of Total Flavonoid Extracts from Selaginella moellendorffii Hieron against Coxsackie Virus B3 In Vitro and In Vivo.
Evidence-Based Complementary and Alternative Medicine, 2014Co-Authors: Dan Yin, Yi-mei Liu, Xiang Lei, Zhanqiu Yang, Ke Li ChenAbstract:The antiviral activity of total flavonoid extracts from Selaginella moellendorffii Hieron and its main constituents amentoflavone were investigated against coxsackie virus B3 (CVB3). When added during or after viral infection, the extracts and amentoflavone prevented the cytopathic effect (CPE) of CVB3, as demonstrated in a 3-(4,5-dimethylthiazol-2-yl)-2,5-diphenyl tetrazolium bromide (MTT) colorimetric assay, with a 50% inhibitory concentration (IC50) from to μg/mL and to μg/mL, respectively. KM mice were used as animal models to test the extracts' activity in vivo. Oral administration of the total flavonoid extracts at 300 mg/kg/day significantly reduced mean viral titers in the heart and kidneys as well as mortality after infection for 15 days. The experimental results demonstrate that in vitro and in vivo the model mice infected with CVB3 can be effectively treated by the total flavonoid extracts from Selaginella moellendorffii Hieron.
Cecilia Zumajocardona - One of the best experts on this subject based on the ideXlab platform.
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all type ii classic mads box genes in the lycophyte Selaginella moellendorffii are broadly yet discretely expressed in vegetative and reproductive tissues
Evolution & Development, 2021Co-Authors: Barbara A Ambrose, Tynisha L Smalls, Cecilia ZumajocardonaAbstract:The MADS-box genes constitute a large transcription factor family that appear to have evolved by duplication and diversification of function. Two types of MADS-box genes are distinguished throughout eukaryotes, types I and II. Type II classic MADS-box genes, also known as MIKC-type, are key developmental regulators in flowering plants and are particularly well-studied for their role in floral organ specification. However, very little is known about the role that these genes might play outside of the flowering plants. We investigated the evolution of type II classic MADS-box genes across land plants by performing a maximum likelihood analysis with a particular focus on lycophytes. Here, we present the expression patterns of all three type II classic MADS-box homologs throughout plant development in the lycophyte Selaginella moellendorffii: SmMADS1, SmMADS3, and SmMADS6. We used scanning electron microscopy and histological analyses to define stages of sporangia development in S. moellendorffii. We performed phylogenetic analyses of this gene lineage across land plants and found that lycophyte sequences appeared before the multiple duplication events that gave rise to the major MADS-box gene lineages in seed plants. Our expression analyses by in situ hybridization show that all type II classic MADS-box genes in S. moellendorffii have broad but distinct patterns of expression in vegetative and reproductive tissues, where SmMADS1 and SmMADS6 only differ during late sporangia development. The broad expression during S. moellendorffii development suggests that MADS-box genes have undergone neofunctionalization and subfunctionalization after duplication events in seed plants.
Jo Ann Banks - One of the best experts on this subject based on the ideXlab platform.
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The Glycosyltransferase Repertoire of the Spikemoss Selaginella moellendorffii and a Comparative Study of Its
2016Co-Authors: Cell Wall, Jesper Harholt, Iben Sørensen, Jonatan Fangel, Alison Roberts, William G. T. Willats, Vibe Scheller, Bent Larsen Petersen, Jo Ann Banks, Peter UlvskovAbstract:Spike mosses are among the most basal vascular plants, and one species, Selaginella moellendorffii, was recently selected for full genome sequencing by the Joint Genome Institute (JGI). Glycosyltransferases (GTs) are involved in many aspects of a plant life, including cell wall biosynthesis, protein glycosylation, primary and secondary metabolism. Here, we present a comparative study of the S. moellendorffii genome across 92 GT families and an additional family (DUF266) likely to include GTs. The study encompasses the moss Physcomitrella patens, a non-vascular land plant, while rice and Arabidopsis represent commelinid and non-commelinid seed plants. Analysis of the subset of GT-families particularly relevant to cell wall polysaccharide biosynthesis was complemented by a detailed analysis of S. moellendorffii cell walls. The S. moellendorffii cell wall contains many of the same components as seed plant cell walls, but appears to differ somewhat in its detailed architecture. The S. moellendorffii genome encodes fewer GTs (287 GTs including DUF266s) than the reference genomes. In a few families, notably GT51 and GT78, S. moellendorffii GTs have no higher plant orthologs, but in most families S. moellendorffii GTs have clear orthologies with Arabidopsis and rice. A gene naming convention of GTs is proposed whic
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The Glycosyltransferase Repertoire of the Spikemoss Selaginella moellendorffii and a Comparative Study of Its
2015Co-Authors: Københavns Universitet, Cell Wall, Jesper Harholt, Iben Sørensen, Jonatan Fangel, Alison Roberts, William G. T. Willats, Vibe Scheller, Bent Larsen Petersen, Jo Ann BanksAbstract:The glycosyltransferase repertoire of the spikemoss Selaginella moellendorffii and a comparative study of its cell wal
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The Glycosyltransferase Repertoire of the Spikemoss Selaginella moellendorffii and a Comparative Study of Its Cell Wall
PLoS ONE, 2012Co-Authors: Jesper Harholt, Iben Sørensen, Jonatan Fangel, William G. T. Willats, Bent Larsen Petersen, Jo Ann Banks, Alison W. Roberts, Henrik Vibe Scheller, Peter UlvskovAbstract:Spike mosses are among the most basal vascular plants, and one species, Selaginella moellendorffii, was recently selected for full genome sequencing by the Joint Genome Institute (JGI). Glycosyltransferases (GTs) are involved in many aspects of a plant life, including cell wall biosynthesis, protein glycosylation, primary and secondary metabolism. Here, we present a comparative study of the S. moellendorffii genome across 92 GT families and an additional family (DUF266) likely to include GTs. The study encompasses the moss Physcomitrella patens, a non-vascular land plant, while rice and Arabidopsis represent commelinid and non-commelinid seed plants. Analysis of the subset of GT-families particularly relevant to cell wall polysaccharide biosynthesis was complemented by a detailed analysis of S. moellendorffii cell walls. The S. moellendorffii cell wall contains many of the same components as seed plant cell walls, but appears to differ somewhat in its detailed architecture. The S. moellendorffii genome encodes fewer GTs (287 GTs including DUF266s) than the reference genomes. In a few families, notably GT51 and GT78, S. moellendorffii GTs have no higher plant orthologs, but in most families S. moellendorffii GTs have clear orthologies with Arabidopsis and rice. A gene naming convention of GTs is proposed which takes orthologies and GT-family membership into account. The evolutionary significance of apparently modern and ancient traits in S. moellendorffii is discussed, as is its use as a reference organism for functional annotation of GTs.
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The Selaginella Genome Identifies Genetic Changes Associated with the Evolution of Vascular Plants
Science, 2011Co-Authors: Jo Ann Banks, Tomoaki Nishiyama, Mitsuyasu Hasebe, John Bowman, Michael Gribskov, Claude De Pamphilis, Victor Albert, Naoki Aono, Tsuyoshi Aoyama, Barbara AmbroseAbstract:Vascular plants appeared ~410 million years ago, then diverged into several lineages of which only two survive: the euphyllophytes (ferns and seed plants) and the lycophytes. We report here the genome sequence of the lycophyte Selaginella moellendorffii (Selaginella), the first nonseed vascular plant genome reported. By comparing gene content in evolutionarily diverse taxa, we found that the transition from a gametophyte- to a sporophyte-dominated life cycle required far fewer new genes than the transition from a nonseed vascular to a flowering plant, whereas secondary metabolic genes expanded extensively and in parallel in the lycophyte and angiosperm lineages. Selaginella differs in posttranscriptional gene regulation, including small RNA regulation of repetitive elements, an absence of the trans-acting small interfering RNA pathway, and extensive RNA editing of organellar genes.
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The highest-copy repeats are methylated in the small genome of the early divergent vascular plant Selaginella moellendorffii.
BMC Genomics, 2008Co-Authors: Agnes P. Chan, Jo Ann Banks, Admasu Melake-berhan, Kimberly O'brien, Stephanie Buckley, Hui Quan, Dan Chen, Matthew Lewis, Pablo D. RabinowiczAbstract:Background The lycophyte Selaginella moellendorffii is a vascular plant that diverged from the fern/seed plant lineage at least 400 million years ago. Although genomic information for S. moellendorffii is starting to be produced, little is known about basic aspects of its molecular biology. In order to provide the first glimpse to the epigenetic landscape of this early divergent vascular plant, we used the methylation filtration technique. Methylation filtration genomic libraries select unmethylated DNA clones due to the presence of the methylation-dependent restriction endonuclease McrBC in the bacterial host.