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Despite growing concerns regarding the long-range transport (LRT) and ecological risks of organophosphate esters (OPEs), information on the environmental behaviors of OPEs in polar terrestrial ecosystems remains inadequate. In the present study, 10 OPEs were analyzed in soil and vegetation samples collected from Fildes Peninsula, Antarctica. The OPE concentrations in Antarctic soils, mosses, and lichens ranged from 0.87 to 15.7 ng/g dry weight (dw), 9.8 to 113 ng/g dw, and 3.6 to 75.2 ng/g dw, respectively. Non-chlorinated OPEs predominated in terrestrial matrices, accounting for approximately 76 % of the OPE composition. Source identification indicated that OPE contamination in Antarctica likely resulted from local anthropogenic sources and LRT. Moreover, the bioaccumulation behavior of OPEs from soil to vegetation was assessed using bioconcentration factors (BCFs), revealing a significant non-linear trend of initial increase and subsequent decrease in BCFs relative to the lipophilicities of the octanol-air partition coefficient (log K OA ) and octanol-water partition coefficient (log K OW ). While low levels of OPEs in Antarctic terrestrial environments were reported in this study, their sustained inputs and potential ecological risks in polar regions warrant further attention., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

The study of moss calyptra form and function began almost 250 years ago, but calyptra research has remained a niche endeavor focusing on only a small number of species. Recent advances have focused on calyptra cuticular waxes, which function in dehydration protection of the immature sporophyte apex. The physical presence of the calyptra also plays a role in sporophyte development, potentially via its influence on auxin transport. Progress developing genomic resources for mosses beyond the model Physcomitrium patens, specifically for species with larger calyptrae and taller sporophytes, in combination with advances in CRISPR-Cas9 genome editing will enable the influence of the calyptra on gene expression and the production of RNAs and proteins that coordinate sporophyte development to be explored., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Jessica M. Budke reports financial support was provided by US National Science Foundation. If there are other authors, they declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

Mosses play a vital role in environmental research as reliable biomonitoring tools. This study aims to understand the accumulation and distribution patterns of Cu and Cd in the acrocarpous moss [Campylopus schmidii (Müll. Hal.) A. Jaeger] (C.schmidii). In controlled in vitro experiments, C.schmidii cultures were exposed to varying concentrations of copper (Cu) and cadmium (Cd) stress (0, 10, 25, 50 μmol/L) in aquatic media. The study systematically evaluated the moss's response, including observing appearance features, oxidative traits, and accumulation characteristics. Scanning electron microscopy with energy-dispersive X-ray spectroscopy analyses were employed. They aimed to characterize and determine the distribution of metal particles in different parts of the mosses under high concentration treatments (50 μmol/L Cd, 50 μmol/L Cu, 50 μmol/L Cu and Cd). Results indicated that C.schmidii exhibited greater tolerance to Cu compared to Cd, as evidenced by significantly higher soluble protein content and lipid peroxidation with increasing concentrations. However, Cd stress induced severe damage, including widespread chlorosis, reduced chlorophyll content, and surface fragmentation. Both Cu and Cd were found to stimulate antioxidant levels by increasing the activity of hydrogen peroxide and peroxidase, thus reducing the accumulation of free radicals in C.schmidii. Additionally, the results revealed differential metal distribution. Higher Cu (2.23%) and lower Cd (0.54%) accumulation were observed at the bottom of gametophores, with Cd content 180.46% higher than Cu at the top. This study provides valuable insights into the potential application of acrocarpous mosses for biomonitoring and phytoremediation. It suggests specific strategies for metal deposition and absorption, such as utilizing upper, younger parts for Cd absorption and lower parts for Cu remediation in soil., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Ltd. All rights reserved.)

The hyperaccumulation potential of zinc (Zn) and cadmium (Cd) and their synergistic effects were examined in relation to Christmas moss (Vesicularia montagnei (Bél) Broth., Hypnaceae), an aquatic and terrestrial moss, dosed with Cd (Cd1 and Cd2), Zn (Zn1 and Zn2) and combined Zn and Cd (Cd1Zn1 and Cd2Zn2). Zinc promoted plant growth and development, particularly in the highest Zn and combined Zn/Cd treatments (Zn2 and Cd2Zn2). The Zn treatment resulted in substantial moss chlorophyll content and highest percentage relative growth rate in biomass value (0.23 mg L -1 and 106.8%, respectively); however, the Cd2Zn2 treatment achieved maximal production of chlorophyll a and total chlorophyll (0.29 and 0.51 mg L -1 , respectively) due to synergistic effects. These findings suggest that Christmas moss is a highly metal-tolerant and adaptable bryophyte species. Zinc was essential for reducing the detrimental effects of Cd while simultaneously promoting moss growth and biomass development. Furthermore, Christmas moss exhibited hyperaccumulation potential for Cd and Zn in the Cd2Zn2 and Zn alone treatments, as evidenced by highest Cd and Zn values in gametophores (1002 and 18,596 mg per colony volume, respectively). Using energy dispersive X-ray fluorescence (EDXRF) spectrometry, atomic percentages of element concentrations in moss gametophores in the Zn2, Cd2 and combined Zn/Cd treatments were generally in the order: K > Ca > P > Zn > Cd. When comparing the atomic percentages of Zn and Cd in gametophores, it is likely that the higher atomic percentage of Zn was because this element is essential for plant growth and development., (© 2024. The Author(s).)

This study investigated the responses of Didymodon constrictus and Hypnum plumaeforme to different light qualities emitted by light-emitting diodes (LEDs), including white light (WL), red light (RL), blue light (BL), yellow light (YL), green light (GL), and a combination of red and blue light (R1B1L). The research analyzed the fluorescence imaging, photosynthetic pigments, coloration, and growth characteristics related to antioxidant enzymes in these two moss species. The results indicated that R1B1L significantly enhanced the content of photosynthetic pigments, maximum relative electron transport rate (rETRmax), saturation light intensity (IK), and the greenness of the moss. RL improved the maximum quantum yield (Fv/Fm), the light energy efficiency of H. plumaeforme and effective quantum yield in both moss species. In contrast, BL notably increased non-photochemical quenching (NPQ), photochemical quenching (qp), and the steady-state fluorescence decrease ratio (RFD) in H. plumaeforme. The application of GL significantly increases the maximum photon yield (Fv/Fm) in D. constrictus, as well as the light energy efficiency and elongation length, resulting in a shift in the color composition of both moss species towards yellow. Among the light treatments, R1B1L had the highest induction rate and promotional effect on the growth of both moss species. These mosses absorbed GL and RL effectively, while BL played a crucial role in the dissipation of heat and electron transfer in H. plumaeforme. This research provides valuable insights for the regulation of LED light environments and the physiological adaptability of moss in artificial cultivation., Competing Interests: Declaration of competing interest The authors declare the following financial interests/personal relationships which may be considered as potential competing interests: Xiurong Wang reports financial support was provided by National Natural Science Foundation of China. If there are other authors, they declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier Masson SAS. All rights reserved.)

The plant cuticle is a hydrophobic barrier, which seals the epidermal surface of most aboveground organs. While the cuticle biosynthesis of angiosperms has been intensively studied, knowledge about its existence and composition in nonvascular plants is scarce. Here, we identified and characterized homologs of Arabidopsis thaliana fatty acyl-CoA reductase (FAR) ECERIFERUM 4 (AtCER4) and bifunctional wax ester synthase/acyl-CoA:diacylglycerol acyltransferase 1 (AtWSD1) in the liverwort Marchantia polymorpha (MpFAR2 and MpWSD1) and the moss Physcomitrium patens (PpFAR2A, PpFAR2B, and PpWSD1). Although bryophyte harbor similar compound classes as described for angiosperm cuticles, their biosynthesis may not be fully conserved between the bryophytes M. polymorpha and P. patens or between these bryophytes and angiosperms. While PpFAR2A and PpFAR2B contribute to the production of primary alcohols in P. patens, loss of MpFAR2 function does not affect the wax profile of M. polymorpha. By contrast, MpWSD1 acts as the major wax ester-producing enzyme in M. polymorpha, whereas mutations of PpWSD1 do not affect the wax ester levels of P. patens. Our results suggest that the biosynthetic enzymes involved in primary alcohol and wax ester formation in land plants have either evolved multiple times independently or undergone pronounced radiation followed by the formation of lineage-specific toolkits., (© 2024 The Authors. New Phytologist © 2024 New Phytologist Foundation.)

The role of forest ecosystems in the global mercury (Hg) biogeochemical cycle is widely recognized; however, using litterfall as a surrogate to assess the Hg sink function of forests encounters limitations. We investigated the accumulation characteristics and influencing factors of Hg in mosses from two remote subalpine forests in southwestern China. The results indicated that there was high Hg accumulation in subalpine forest mosses, with average concentrations of 82 ± 49 ng g -1 for total mercury (THg) and 1.3 ± 0.8 ng g -1 for methylmercury (MeHg). We demonstrated that the accumulation capacity of Hg in mosses was significantly dependent on species and substrates (micro-habitats), the mosses on tree trunks exhibited significantly elevated Hg accumulation levels (THg 132 ± 56 ng g -1 , MeHg 1.6 ± 0.2 ng g -1 ) compared to mosses in other substrates. The surface morphologies and biochemical components of leaf (phyllidia), such as cation exchange capacity (CEC), pectin, uronic acid, and metallothionein, play a crucial role in the accumulation of Hg by mosses. These findings provide valuable insights into Hg accumulation in forest mosses. Suggesting that the contribution of mosses Hg accumulation should be considered when assessing atmospheric Hg sinks of forests., Competing Interests: Declaration of Competing Interest The authors declare there is no conflict of interest regarding the publication of this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

Bryophytes desiccate rapidly when relative humidity decreases. The capacity to withstand dehydration depends on several ecological and physiological factors. Volatile organic compounds (VOCs) may have a role in enhancing tolerance to desiccating bryophytes. However, the functions of VOCs in bryophytes have received little attention so far. We aimed to investigate the impact of a dehydration-rehydration treatment on primary carbon metabolism and volatile terpenes (VTs) in three bryophytes with contrasting ecological traits: Vessicularia dubyana, Porella platyphylla and Pleurochaete squarrosa. First, we confirmed the desiccation sensitivity gradient of the species. Under fully hydrated conditions, the photosynthetic rate (A) was inversely associated with stress tolerance, with a lower rate in more tolerant species. The partial recovery of A in P. platyphylla and P. squarrosa after rehydration confirmed the desiccation tolerance of these two species. On the other hand, A did not recover after rehydration in V. dubyana. Regarding VT, each species exhibited a distinct VT profile under optimum hydration, with the highest VT pool found in the more desiccation-sensitive species (V. dubyana). However, the observed species-specific VT pattern could be associated with the ecological habitat of each species. P. squarrosa, a moss of dry habitats, may synthesize mainly non-volatile secondary metabolites as stress-defensive compounds. On the other hand, V. dubyana, commonly found submerged, may need to invest photosynthetically assimilated carbon to synthesize a higher amount of VTs to cope with transient water stress occurrence. Further research on the functions of VTs in bryophytes is needed to deepen our understanding of their ecological significance., (© 2024 The Author(s). Physiologia Plantarum published by John Wiley & Sons Ltd on behalf of Scandinavian Plant Physiology Society.)

Plant movements for survival are nontrivial. Antheridia in the moss Physcomitrium patens (P. patens) use motion to eject sperm in the presence of water. However, the biological and mechanical mechanisms that actuate the process are unknown. Here, the burst of the antheridium of P. patens, triggered by water, results from elastic instability and is determined by an asymmetric change in cell geometry. The tension generated in jacket cell walls of antheridium arises from turgor pressure, and is further promoted when the inner walls of apex burst in hydration, causing water and cellular contents of apex quickly influx into sperm chamber. The outer walls of the jacket cells are strengthened by NAC transcription factor VNS4 and serve as key morphomechanical innovations to store hydrostatic energy in a confined space in P. patens. However, the antheridium in liverwort Marchantia polymorpha (M. polymorpha) adopts a different strategy for sperm release; like jacket cell outer walls of P. patens, the cells surrounding the antheridium of M. polymorpha appear to play a similar role in the storage of energy. Collectively, the work shows that plants have evolved different ingenious devices for sperm discharge and that morphological innovations can differ., (© 2024 The Authors. Advanced Science published by Wiley‐VCH GmbH.)

The model moss species Physcomitrium patens has long been used for studying divergence of land plants spanning from bryophytes to angiosperms. In addition to its phylogenetic relationships, the limited number of differential tissues, and comparable morphology to the earliest embryophytes provide a system to represent basic plant architecture. Based on plant-fungal interactions today, it is hypothesized these kingdoms have a long-standing relationship, predating plant terrestrialization. Mortierellaceae have origins diverging from other land fungi paralleling bryophyte divergence, are related to arbuscular mycorrhizal fungi but are free-living, observed to interact with plants, and can be found in moss microbiomes globally. Due to their parallel origins, we assess here how two Mortierellaceae species, Linnemannia elongata and Benniella erionia, interact with P. patens in coculture. We also assess how Mollicute-related or Burkholderia-related endobacterial symbionts (MRE or BRE) of these fungi impact plant response. Coculture interactions are investigated through high-throughput phenomics, microscopy, RNA-sequencing, differential expression profiling, gene ontology enrichment, and comparisons among 99 other P. patens transcriptomic studies. Here we present new high-throughput approaches for measuring P. patens growth, identify novel expression of over 800 genes that are not expressed on traditional agar media, identify subtle interactions between P. patens and Mortierellaceae, and observe changes to plant-fungal interactions dependent on whether MRE or BRE are present. Our study provides insights into how plants and fungal partners may have interacted based on their communications observed today as well as identifying L. elongata and B. erionia as modern fungal endophytes with P. patens., (© 2024 The Authors. The Plant Journal published by Society for Experimental Biology and John Wiley & Sons Ltd.)

Cytidine (C)-to-uridine (U) RNA editing in plant organelles relies on specific RNA-binding pentatricopeptide repeat (PPR) proteins. In the moss Physcomitrium patens, all such RNA editing factors feature a C-terminal DYW domain that acts as the cytidine deaminase for C-to-U conversion. PPR78 of Physcomitrium targets 2 mitochondrial editing sites, cox1eU755SL and rps14eU137SL. Remarkably, the latter is edited to highly variable degrees in different mosses. Here, we aimed to unravel the coevolution of PPR78 and its 2 target sites in mosses. Heterologous complementation in a Physcomitrium knockout line revealed that the variable editing of rps14eU137SL depends on the PPR arrays of different PPR78 orthologues but not their C-terminal domains. Intriguingly, PPR78 has remained conserved despite the simultaneous loss of editing at both known targets among Hypnales (feather mosses), suggesting it serves an additional function. Using a recently established RNA editing assay in Escherichia coli, we confirmed site-specific RNA editing by PPR78 in the bacterium and identified 4 additional off-targets in the bacterial transcriptome. Based on conservation profiles, we predicted ccmFNeU1465RC as a candidate editing target of PPR78 in moss mitochondrial transcriptomes. We confirmed editing at this site in several mosses and verified that PPR78 targets ccmFNeU1465RC in the bacterial editing system, explaining the conservation and functional adaptation of PPR78 during moss evolution., Competing Interests: Conflict of interest statement. None declared., (© The Author(s) 2023. Published by Oxford University Press on behalf of American Society of Plant Biologists. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

This is the first report on the molecular characterization of isoprene synthase (ISPS) from the moss Calohypnum plumiforme. After isoprene emission from C. plumiforme was confirmed, the cDNA encoding C. plumiforme ISPS (CpISPS) was narrowed down using a genome database associated with protein structure prediction, and a CpISPS gene was identified. The recombinant CpISPS, produced in Escherichia coli, converted dimethylallyl diphosphate to isoprene. Phylogenetic analysis indicated similarity between the amino acid sequences of CpISPS and moss diterpene cyclases (DTCs) but not ISPSs of higher plants, implying that CpISPS is derived from moss DTCs and is evolutionarily unrelated to canonical ISPSs of higher plants. CpISPS is a novel class I cyclase of the terpene synthase-c subfamily harboring αβ domains. This study will help further study of isoprene biosynthesis and the physiological functions of isoprene in mosses., (© 2023 Federation of European Biochemical Societies.)

Cytochrome P450 monooxygenase 98 (CYP98) is a critical rate-limiting enzyme of the phenylpropanoid pathway. One of the end-product of the phenylpropanoid pathway is a lignin monomer, although the occurrence of lignin in bryophytes is controversial. Here we investigated the functions of PpCYP98 in Physcomitrium patens by transcriptome and metabolome analyses. We identified 5266 differentially expressed genes (DEGs) and 68 differentially abundant secondary metabolites between wild-type and ΔPpCYP98 gametophores. Of the identified metabolites, 23 phenolic acids were identified, with only one showing upregulation. Among the phenolic acids, 4-coumaroyl tartaric acid and chlorogenic acid showed significant decreases. Declines were also observed in coniferylaldehyde and coniferin, precursor substances and downstream products of the lignin monomer coniferyl alcohol, respectively. Thus, the pre-lignin synthesis pathway already exists in bryophytes, and PpCYP98 plays vital roles in this pathway. Besides, most flavonoids show significant reductions, including eriodyctiol, dihydroquecetin, and dihydromyricetin, whereas naringenin chalone and dihydrokaempferol were increased after PpCYP98 knockout. Therefore, the synthesis of flavonoids shares the core pathway with phenylpropanoids and mainly starts from caffeoyl-CoA, that is the compound of divergence between the two pathways in moss. PpCYP98 showed systemic effects on metabolisms, including carbohydrate, fatty acid, and hormonal signaling transductions, suggesting that PpCYP98 might indirectly regulate carbon influx allocation. Our results demonstrated roles of PpCYP98 were essential for the development of the early landing plant., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Masson SAS. All rights reserved.)

In response to varying light conditions, light-harvesting complexes (LHCs) switch from a light-harvesting state to a quenched state to protect the photosynthetic organism from excessive light irradiation in a strategy known as nonphotochemical quenching (NPQ). NPQ is activated by an acidification of the thylakoid lumen, which is sensed directly or indirectly by the LHC, resulting in a conformational change of the complex that leads to the quenched state. The conformational changes responsible for NPQ activation and their connection to specific quenching mechanisms are still unknown. Here, we investigate the pH-triggered conformational changes in the light-harvesting complex stress-related (LHCSR) of mosses. By combining constant-pH molecular dynamics and enhanced sampling techniques, we find that the pH sensitivity of the complex is driven by the coupled protonation of three residues modulating the conformation of the short amphipathic helix placed at the lumen side of the embedding membrane. Combining these results with quantum mechanics/molecular mechanics calculations, we show that the quenching mechanism sensitive to the pH goes through a charge-transfer between a carotenoid and an excited chlorophyll, which is controlled by the protein conformation.

Antarctic organisms are consistently suffering from multiple environmental pressures, especially the strong UV radiation caused by the loss of the ozone layer. The mosses and lichens dominate the vegetation of the Antarctic continent, which grow and propagate in these harsh environments. However, the molecular mechanisms and related regulatory networks of these Antarctic plants against UV-B radiation are largely unknown. Here, we used an integrated multi-omics approach to study the regulatory mechanism of long non-coding RNAs (lncRNAs) of an Antarctic moss ( Pohlia nutans ) in response to UV-B radiation. We identified a total of 5729 lncRNA sequences by transcriptome sequencing, including 1459 differentially expressed lncRNAs (DELs). Through functional annotation, we found that the target genes of DELs were significantly enriched in plant-pathogen interaction and the flavonoid synthesis pathway. In addition, a total of 451 metabolites were detected by metabonomic analysis, and 97 differentially change metabolites (DCMs) were found. Flavonoids account for 20% of the total significantly up-regulated metabolites. In addition, the comprehensive transcriptome and metabolome analyses revealed the co-expression pattern of DELs and DCMs of flavonoids. Our results provide insights into the regulatory network of lncRNA under UV-B radiation and the adaptation of Antarctic moss to the polar environments.

The auxin efflux PIN-FORMED (PIN) proteins are conserved in all land plants and important players in plant development. In the moss Physcomitrella (Physcomitrium patens), three canonical PINs (PpPINA-C) are expressed in the leafy shoot (gametophore). PpPINA and PpPINB show functional activity in vegetative growth and sporophyte development. Here, we examined the role of PpPINC in the life cycle of Physcomitrella. We established reporter and knockout lines for PpPINC and analysed vegetative and reproductive tissues using microscopy and transcriptomic sequencing of moss gametangia. PpPINC is expressed in immature leaves, mature gametangia and during sporophyte development. The sperm cells (spermatozoids) of pinC knockout mutants exhibit increased motility and an altered flagella phenotype. Furthermore, the pinC mutants have a higher portion of differentially expressed genes related to spermatogenesis, increased fertility and an increased abortion rate of premeiotic sporophytes. Here, we show that PpPINC is important for spermatogenesis and sporophyte retention. We propose an evolutionary conserved way of polar growth during early moss embryo development and sporophyte attachment to the gametophore while suggesting the mechanical function in sporophyte retention of a ring structure, the Lorch ring., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

The moss Physcomitrium patens is crucial for studying plant development and evolution. Although the P. patens genome includes genes acquired from bacteria, fungi and viruses, the functions and evolutionary significance of these acquired genes remain largely unclear. Killer protein 4 (KP4) is a toxin secreted by the phytopathogenic fungus Ustilago maydis that inhibits the growth of sensitive target strains by blocking their calcium uptake. Here, we show that KP4 genes in mosses were acquired from fungi through at least three independent events of horizontal gene transfer. Two paralogous copies of KP4 (PpKP4-1 and PpKP4-2) exist in P. patens. Knockout mutants ppkp4-1 and ppkp4-2 showed cell death at the protonemal stage, and ppkp4-2 also exhibited defects in tip growth. We provide experimental evidence indicating that PpKP4-1/2 affects P. patens protonemal cell development by mediating cytoplasmic calcium and that KP4 genes are functionally conserved between P. patens and fungi. The present study provides additional insights into the role of horizontal gene transfer in land plant development and evolution., (© 2022 The Authors. The Plant Journal published by Society for Experimental Biology and John Wiley & Sons Ltd.)

Soloist is a member of a distinct and small subfamily within the AP2/ERF transcriptional factor family that play important roles in plant biotic and abiotic stress responses. There are limited studies of Soloist genes and their functions are poorly understood. We characterized the abiotic and biotic stress tolerance function of the ScSoloist gene (designated as ScAPD1-like) from the desert moss Syntrichia caninervis. ScAPD1-like responded to multiple abiotic, biotic stresses and plant hormone treatments. ScAPD1-like protein located to the nucleus and bound to several DNA elements. Overexpression of ScAPD1-like in Arabidopsis did not alter abiotic stress resistance or inhibit Pseudomonas syringae pv. tomato (Pst) DC3000 infection. However, overexpression of ScAPD1-like significantly increased the resistance of transgenic Arabidopsis and S. caninervis to Verticillium dahliae infection, decreased reactive oxygen species accumulation and improved reactive oxygen species scavenging activity. ScAPD1-like overexpression plants altered the abundance of transcripts for lignin synthesis and promoted lignin accumulation in Arabidopsis. ScAPD1-like directly bind to RAV1, AC elements, and TATA-box in the promoters of AtPAL1 and AtC4H genes, respectively, in vitro. Chromatin immunoprecipitation-quantitative polymerase chain reaction assays demonstrated ScAPD1-like directly bound to PAL and C4H genes promoters in Arabidopsis and their homologs in S. caninervis. In S. caninervis, ScAPD1-like overexpression and RNAi directly regulated the abundance of ScPAL and ScC4H transcripts and modified the metabolites of phenylpropanoid pathway. We provide insight into the function of Soloist in plant defense mechanisms that likely occurs through activation of the phenylpropanoid biosynthesis pathway. ScAPD1-like is a promising candidate gene for breeding strategies to improve resistance to Verticillium wilt., (© 2022 Society for Experimental Biology and John Wiley & Sons Ltd.)

C2H2 zinc finger protein (C2H2-ZFP) plays an important role in regulating plant growth, development, and response to abiotic stress. To date, there have been no analyses of the C2H2-ZFP family in desiccation-tolerant moss. In this study, we identified 57 BaZFP transcripts across the Bryum argenteum ( B. argenteum ) transcriptome. The BaZFP proteins were phylogenetically divided into four groups (I-IV). Additionally, we studied the BaZFP1 gene, which is a nuclear C2H2-ZFP and acts as a positive regulator of growth and development in both moss and Arabidopsis thaliana . The complete coding sequence of the BaZFP1 gene was isolated from B. argenteum cDNA, which showed a high expression level in a dehydration-rehydration treatment process. The overexpression of the BaZFP1 gene in the Physcomitrium patens and B. argenteum promoted differentiation and growth of gametophytes. Heterologous expression in Arabidopsis regulated the whole growth and development cycle. In addition, we quantitatively analyzed the genes related to growth and development in transgenic moss and Arabidopsis , including HLS1 , HY5 , ANT , LFY , FT , EIN3 , MUS , APB4 , SEC6 , and STM1 , and found that their expression levels changed significantly. This study may pave the way for substantial insights into the role of C2H2-ZFPs in plants as well as suggest appropriate candidate genes for crop breeding.

Bryophyta comprises one of the earliest lineages of land plants that had implemented remarkable innovations to their lipid metabolic systems for successful adaptation to terrestrial habitat. This study presents a comprehensive investigation of fatty acid profiles of mosses from Eastern Himalayas with an aim to trace their chemotaxonomic and evolutionary implications. Fatty acid compositions of 40 random mosses belonging to major families of Bryophyta were explored by gas chromatographic analysis. A diverse array of saturated, monounsaturated and polyunsaturated fatty acids including rare acetylenic fatty acids were detected. Hexadecanoic acid (C16:0), 9,12 (Z,Z)-octadecadienoic acid (C18:2n6) and 9,12,15 (Z,Z,Z)-octadecatrienoic acid (C18:3n3) were the predominant fatty acids in all the mosses. However, quantitative variation of C20 polyunsaturated fatty acids (PUFAs), specifically 5,8,11,14 (Z,Z,Z,Z)-eicosatetraenoic acid (C20:4n6), among the investigated mosses was the most prominent outcome. The diplolepidous members of Bryidae, especially the mosses of Hypnales, Bryales and Bartramiales contained higher amount of C20 PUFAs compared with the haplolepidous orders. Principal component analyses based on individual fatty acids and other related parameters validated C20:4n6 content and the ratio of C20:4n6/C18:2n6 as the apparent chemotaxonomic discriminants. The prevalent notion of considering 9,12,15-octadecatrien-6-ynoic acid (C18:4a) as the chemomarker of Dicranaceae has also been challenged, since the compound was detected not only in different families of Dicranales, but also in a Pottiales member, Leptodontium viticulosoides. Therefore, an ensemble of fatty acids instead of a single one can be considered as the chemical signature for taxonomic interpretation which may also be vital from an evolutionary standpoint., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Austria, part of Springer Nature.)

Bryophytes are the group of land plants with the lowest photosynthetic rates, which was considered to be a consequence of their higher anatomical CO2 diffusional limitation compared with tracheophytes. However, the most recent studies assessing limitations due to biochemistry and mesophyll conductance in bryophytes reveal discrepancies based on the methodology used. In this study, we compared data calculated from two different methodologies for estimating mesophyll conductance: variable J and the curve-fitting method. Although correlated, mesophyll conductance estimated by the curve-fitting method was on average 4-fold higher than the conductance obtained by the variable J method; a large enough difference to account for the scale of differences previously shown between the biochemical and diffusional limitations to photosynthesis. Biochemical limitations were predominant when the curve-fitting method was used. We also demonstrated that variations in bryophyte relative water content during measurements can also introduce errors in the estimation of mesophyll conductance, especially for samples which are overly desiccated. Furthermore, total chlorophyll concentration and soluble proteins were significantly lower in bryophytes than in tracheophytes, and the percentage of proteins quantified as Rubisco was also significantly lower in bryophytes (<6.3% in all studied species) than in angiosperms (>16% in all non-stressed cases). Photosynthetic rates normalized by Rubisco were not significantly different between bryophytes and angiosperms. Our data suggest that the biochemical limitation to photosynthesis in bryophytes is more relevant than so far assumed., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

The data available so far indicate that the photosynthetic and relative growth rates of bryophytes are 10% of those reported for tracheophytes. By examining the existing literature and reanalysing data published in over 100 studies, this review examines the ecophysiological, biochemical, and structural reasons behind this phenomenon. The limiting Rubisco content and surface for gas exchange are the internal factors that can explain the low photosynthetic and growth rates of bryophytes. The role of the thicker cell walls of bryophytes in limiting CO2 diffusion is unclear, due to the current uncertainties regarding their porosity and permeability to CO2. From this review, it is also evident that, despite bryophytes having low photosynthetic rates, their positive carbon balance is tightly related to their capacity to deal with extreme conditions. Contributing factors include their capacity to deal with large daily temperature oscillations, and their capacity to delay the cessation of photosynthesis under water deficit (or to tolerate desiccation in extreme situations). Although further studies on bryophytes are needed before more solid conclusions can be drawn, it seems that their success relies on their remarkable tolerance to a highly variable environment, possibly at the expense of their maximum photosynthetic rate., (© The Author(s) 2022. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Sewage and industrial waste discharges have been found to have a deleterious effect on plant growth and environmental safety through the accumulation of trace metal mercury (Hg) in soils. Although the effects of Hg on vascular plants have been reported in terms of enzyme activity, oxidative damage and physiology, few studies have been done on non-vascular plants. A simulation experiment including 7 Hg concentrations (0, 10, 20, 30, 40, 50, 75 µM) was conducted to investigate the influence of Hg stress on ultrastructure and physiological properties of biocrust moss Syntrichia. caninervis across 7 consecutive days. The results showed that the lowest lethal concentration of S. caninervis was 30 µM Hg. The mortality rate of the plants increased significantly with Hg concentrations. The ultrastructure did not change significantly at Hg concentration ≤ 20 µM, while exceeding which, cell walls began to separate, nuclei began to blur, and chloroplasts began to expand. The soluble sugars (SS), peroxidase (POD), and superoxide dismutase (SOD) activities increased initially and then decreased with the increase of concentration in the time gradient, with the largest values at 20 µM. The contents of malondialdehyde (MDA) and proline (Pro) increased with the increase of Hg concentration, both reached peak value at 50 µM. However, chlorophyll (Chl) contents continued to decrease along both the concentration and time gradients. Pearson correlation and principal component analysis showed that two principal components (PC1 and PC2) explained 73.9% of the variance in plant adaptation to Hg stress. SOD, POD, Chl, SS, and Pro all responded well to Hg in S. caninervis. Our study showed that Hg stress caused changes in ultrastructure and physiological metabolism of S. caninervis. 20 µM was the maximum concentration of Hg that biocrust moss S. caninervis can tolerate. S. caninervis mainly adopted two adaptation strategies related to exclusion and accumulation to reduce Hg stress., (© 2022. The Author(s).)

The CLAVATA pathway is a key regulator of stem cell function in the multicellular shoot tips of Arabidopsis, where it acts via the WUSCHEL transcription factor to modulate hormone homeostasis. Broad-scale evolutionary comparisons have shown that CLAVATA is a conserved regulator of land plant stem cell function, but CLAVATA acts independently of WUSCHEL-like (WOX) proteins in bryophytes. The relationship between CLAVATA, hormone homeostasis and the evolution of land plant stem cell functions is unknown. Here we show that in the moss, Physcomitrella (Physcomitrium patens), CLAVATA affects stem cell activity by modulating hormone homeostasis. CLAVATA pathway genes are expressed in the tip cells of filamentous tissues, regulating cell identity, filament branching, plant spread and auxin synthesis. The receptor-like kinase PpRPK2 plays the major role, and Pprpk2 mutants have abnormal responses to cytokinin, auxin and auxin transport inhibition, and show reduced expression of PIN auxin transporters. We propose a model whereby PpRPK2 modulates auxin gradients in filaments to determine stem cell identity and overall plant form. Our data indicate that CLAVATA-mediated auxin homeostasis is a fundamental property of plant stem cell function, probably exhibited by the last shared common ancestor of land plants., (© 2022 The Authors. New Phytologist © 2022 New Phytologist Foundation.)

Plant GH19 chitinases have several loop structures, which may define their enzymatic properties. Among these loops, the longest loop, Loop-III, is most frequently conserved in GH19 enzymes. A GH19 chitinase from the moss Bryum coronatum (BcChi-A) has only one loop structure, Loop-III, which is connected to the catalytically important β-sheet region. Here, we produced and characterized a Loop-III-deleted mutant of BcChi-A (BcChi-A-ΔIII) and found that its stability and chitinase activity were strongly reduced. The deletion of Loop-III also moderately affected the chitooligosaccharide binding ability as well as the binding mode to the substrate-binding groove. The crystal structure of an inactive mutant of BcChi-A-ΔIII was successfully solved, revealing that the remaining polypeptide chain has an almost identical fold to that of the original protein. Loop-III is not necessarily essential for the folding of the enzyme protein. However, closer examination of the crystal structure revealed that the deletion of Loop-III altered the arrangement of the catalytic triad, Glu61, Glu70 and Ser102, and the orientation of the Trp103 side chain, which is important for sugar residue binding. We concluded that Loop-III is not directly involved in the enzymatic activity but assists the enzyme function by stabilizing the conformation of the β-sheet region and the adjacent substrate-binding platform from behind the core-functional regions., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Antarctic mosses are the dominant vegetation in the Antarctic continent. Because of stratospheric ozone depletion, they can withstand physiological extreme UV. The formation of CPD and 6-4PP is one of the most harmful damages of UV to DNA. DNA damage can interfere with replication and transcription, resulting in mutation and death. Two types of photolyase, CPD photolyase and 6-4 photolyase, are capable of specific binding CPD or 6-4PP and repairing these lesions. However, there is little research on photolyase in Antarctic moss. Here, we isolated a gene encoding class II CPD photolyase (PnCPDPhr) and a gene encoding 6-4 photolyase (Pn6-4Phr) from Antarctic moss P. nutans M211. When exposed to UVB, CPDs accumulated in gametophytes and the gene expressions of PnCPDPhr and Pn6-4Phr were both up-regulated. In addition, the in vitro expression and photoreactivation assays of PnCPDPhr and Pn6-4Phr were performed. Our results demonstrated that PnCPDPhr and Pn6-4Phr have an effective activity of DNA repair. This is the first study to determine the CPD accumulation in Antarctic moss as well as the first report isolating CPD photolyase and 6-4 photolyase from Antarctic moss. These results will enrich the knowledge of photolyase family and benefit the exploitation of functioning gene in Antarctic moss., (© 2021 American Society for Photobiology.)

Key Message: Here we review, from a quantitative point of view, the cell biology of protonemal tip growth in the model moss Physcomitrium patens. We focus on the role of the cytoskeleton, vesicle trafficking, and cell wall mechanics, including reviewing some of the existing mathematical models of tip growth. We provide a primer for existing cell biological tools that can be applied to the future study of tip growth in moss. Polarized cell growth is a ubiquitous process throughout the plant kingdom in which the cell elongates in a self-similar manner. This process is important for nutrient uptake by root hairs, fertilization by pollen, and gametophyte development by the protonemata of bryophytes and ferns. In this review, we will focus on the tip growth of moss cells, emphasizing the role of cytoskeletal organization, cytoplasmic zonation, vesicle trafficking, cell wall composition, and dynamics. We compare some of the existing knowledge on tip growth in protonemata against what is known in pollen tubes and root hairs, which are better-studied tip growing cells. To fully understand how plant cells grow requires that we deepen our knowledge in a variety of forms of plant cell growth. We focus this review on the model plant Physcomitrium patens, which uses tip growth as the dominant form of growth at its protonemal stage. Because mosses and vascular plants shared a common ancestor more than 450 million years ago, we anticipate that both similarities and differences between tip growing plant cells will provide mechanistic information of tip growth as well as of plant cell growth in general. Towards this mechanistic understanding, we will also review some of the existing mathematical models of plant tip growth and their applicability to investigate protonemal morphogenesis. We attempt to integrate the conclusions and data across cell biology and physical modeling to our current state of knowledge of polarized cell growth in P. patens and highlight future directions in the field., (© 2021. The Author(s), under exclusive licence to Springer Nature B.V.)

Key Message: Most known phytohormones regulate moss development. We present a comprehensive view of the synthesis and signaling pathways for the most investigated of these compounds in mosses, focusing on the model Physcomitrium patens. The last 50 years of research have shown that most of the known phytohormones are synthesized by the model moss Physcomitrium patens (formerly Physcomitrella patens) and regulate its development, in interaction with responses to biotic and abiotic stresses. Biosynthesis and signaling pathways are best described in P. patens for the three classical hormones auxins, cytokinins and abscisic acid. Furthermore, their roles in almost all steps of development, from early filament growth to gametophore development and sexual reproduction, have been the focus of much research effort over the years. Evidence of hormonal roles exist for ethylene and for CLE signaling peptides, as well as for salicylic acid, although their possible effects on development remain unclear. Production of brassinosteroids by P. patens is still debated, and modes of action for these compounds are even less known. Gibberellin biosynthesis and signaling may have been lost in P. patens, while gibberellin precursors such as ent-kaurene derivatives could be used as signals in a yet to discover pathway. As for jasmonic acid, it is not used per se as a hormone in P. patens, but its precursor OPDA appears to play a corresponding role in defense against abiotic stress. We have tried to gather a comprehensive view of the biosynthesis and signaling pathways for all these compounds in mosses, without forgetting strigolactones, the last class of plant hormones to be reported. Study of the strigolactone response in P. patens points to a novel signaling compound, the KAI2-ligand, which was likely employed as a hormone prior to land plant emergence., (© 2021. The Author(s), under exclusive licence to Springer Nature B.V.)

As an adaptive innovation in plant terrestrialization, cuticle covers the plant surface and greatly contributes to the development and stress tolerance in land plants. Although past decades have seen great progress in understanding the molecular mechanism of cuticle biosynthesis in flowering plants with the contribution of cuticle biosynthesis mutants and advanced cuticle composition profiling techniques, origins and evolution of cuticle biosynthesis are poorly understood. Recent chemical, phylogenomic, and molecular genetic studies on cuticle biosynthesis in early-diverging extant land plant lineages, the bryophytes, shed novel light on the origins and evolution of plant cuticle biosynthesis. In this mini-review, we highlighted these recent advances in the molecular biology of cuticle biosynthesis in bryophytes, and provided evolutionary insights into plant cuticle biosynthesis.

Mosses are mainly the object of ecological and taxonomic research. This group of plants are still underestimated by scientists in other aspects of research. Recent research has shown that these plants contain remarkable and unique substances with high biological activity. Five species of mosses from a large urban ecosystem were identified for present study. In order to determine their biological potential, multifaceted studies were carried out, including: total phenolics content, antioxidant activity, antimicrobial and antifungal study, cytotoxicity evaluation, and scratch assay to assess pro-regenerative effect in the context of their possible use as the ingredients of biologically active cosmetics. Additionally, determination of individual phenolic compounds in selected extracts of the tested mosses was made. Research showed that Ceratodon purpureus and Dryptodon pulvinatus extracts had the greatest potential as antioxidants and antimicrobial activity. The cytotoxicity assessment indicated that the extracts from Dryptodon pulvinatus and Rhytidiadelphus squarossus exerted the strongest negative effect on mouse fibroblast line L929 viability at higher concentrations. While, the extract from Tortulla muralis best stimulated human foreskin fibroblast line HFF-1 proliferation and wound healing. The research on individual phenolic compounds content in the extracts tested indicated over 20 peaks on UPLC chromatograms. The conducted study has shown that mosses, especially so far unexplored species of open ecosystems, and e.g. epilytic habitats, may be a valuable source of biologically active substances and thus may constitute important medical and cosmetic possibilities., Competing Interests: The authors have declared that no competing interests exist.

Noble metals nanoparticles (NPs) and metal oxide NPs are widely used in different fields of application and commercial products, exposing living organisms to their potential adverse effects. Recent evidences suggest their presence in the aquifers water and consequently in drinking water. In this work, we have carefully synthesized four types of NPs, namely, silver and gold NPs (Ag NPs and Au NPs) and silica and titanium dioxide NPs (SiO 2 NPs and TiO 2 NPs) having a similar size and negatively charged surfaces. The synthesis of Ag NPs and Au NPs was carried out by colloidal route using silver nitrate (AgNO 3 ) and tetrachloroauric (III) acid (HAuCl 4 ) while SiO 2 NPs and TiO 2 NPs were achieved by ternary microemulsion and sol-gel routes, respectively. Once the characterization of NPs was carried out in order to assess their physico-chemical properties, their impact on living cells was studied. We used the human colorectal adenocarcinoma cells (Caco-2), known as the best representative intestinal epithelial barrier model to understand the effects triggered by NPs through ingestion. Then, we moved to explore how water contamination caused by NPs can be lowered by the ability of three species of aquatic moss, namely, Leptodictyum riparium , Vesicularia ferriei , and Taxiphyllum barbieri, to absorb them. The experiments were conducted using two concentrations of NPs (100 μM and 500 Μm as metal content) and two time points (24 h and 48 h), showing a capture rate dependent on the moss species and NPs type. Then, the selected moss species, able to actively capture NPs, appear as a powerful tool capable to purify water from nanostructured materials, and then, to reduce the toxicity associated to the ingestion of contaminated drinking water.

Detecting and quantifying low-abundance (deoxy)ribonucleotides and (deoxy)ribonucleosides in plants remains difficult; this is a major roadblock for the investigation of plant nucleotide (NT) metabolism. Here, we present a method that overcomes this limitation, allowing the detection of all deoxy- and ribonucleotides as well as the corresponding nucleosides from the same plant sample. The method is characterized by high sensitivity and robustness enabling the reproducible detection and absolute quantification of these metabolites even if they are of low abundance. Employing the new method, we analyzed Arabidopsis thaliana null mutants of CYTIDINE DEAMINASE, GUANOSINE DEAMINASE, and NUCLEOSIDE HYDROLASE 1, demonstrating that the deoxyribonucleotide (dNT) metabolism is intricately interwoven with the catabolism of ribonucleosides (rNs). In addition, we discovered a function of rN catabolic enzymes in the degradation of deoxyribonucleosides in vivo. We also determined the concentrations of dNTs in several mono- and dicotyledonous plants, a bryophyte, and three algae, revealing a correlation of GC to AT dNT ratios with genomic GC contents. This suggests a link between the genome and the metabolome previously discussed but not experimentally addressed. Together, these findings demonstrate the potential of this new method to provide insight into plant NT metabolism., (© American Society of Plant Biologists 2020. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Bryophytes, including liverworts, mosses, and hornworts, are gametophyte-dominant land plants that are derived from a common ancestor and underwent independent evolution from the sporophyte-dominant vascular plants since their divergence. The plant hormone auxin has been shown to play pleiotropic roles in the haploid bodies of bryophytes. Pharmacological and chemical studies identified conserved auxin molecules, their inactivated forms, and auxin transport in bryophyte tissues. Recent genomic and molecular biological studies show deep conservation of components and their functions in auxin biosynthesis, inactivation, transport, and signaling in land plants. Low genetic redundancy in model bryophytes enable unique assays, which are elucidating the design principles of the auxin signaling pathway. In this article, the physiological roles of auxin and regulatory mechanisms of gene expression and development by auxin in Bryophyta are reviewed., (Copyright © 2021 Cold Spring Harbor Laboratory Press; all rights reserved.)

There have been several published reports regarding the growth promoting effect of humic acids (HA) on vascular plants; however, the effect of HA on bryophytes is still unknown. Due to the ecological importance of mosses, which dominate the Antarctic flora, we assessed the effectiveness of HA as a biostimulant using three moss species: Antarctic Ceratodon purpureus KMA5038, Arctic Bryum sp. KMR5045, and Physcomitrella patens which inhabits temperate regions. Natural HA (KS1-3&#95;HA) were extracted through acidic precipitation of alkaline extracts from Antarctic tundra soil. Spectroscopic structural properties of KS1-3&#95;HA were characterized and determined to possess several functional groups such as hydroxyl (R-OH) and carboxyl (R-COOH), implying they could have a growth-related biological function. For two polar mosses, increasing HA concentrations correlated with increased growth and photosynthesis. The efficiency for temperate moss increased at lower concentrations tested, but rather began to reduce at the highest HA concentration, indicating that effective concentrations of HA vary depending on the moss species and habitat. Based on these results, Antarctic HA may have ecological role in enhancing the growth and photosynthesis of Antarctic mosses. We believe this is the first study to establish a positive physiological effect of HA on mosses and hope it may serve as a basis for studying the role of HA in preserving the terrestrial ecosystem of Antarctica., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

The physiological responses of desert moss crusts under four artificial media (Beneck, Part, BG11, and Hogland) were investigated to evaluate the function of culture media during different culture periods. The results showed that the value of malondialdehyde (MDA) was at a maximum at 11d, on the contrary, chlorophyll-a, soluble protein, and soluble sugar were at a minimum. As the time increased, the value of MDA and soluble protein decreased faster in the Hogland, while the value of chlorophyll-a and soluble sugar increased. At the end of the culture period, the value of chlorophyll-a and soluble sugar was at a maximum in the Hogland, while the value of MDA and soluble protein was at a minimum. The results suggested that the Hogland medium had a promoting effect on the growth of desert moss crusts. The selected artificial cultivation medium towards wider and larger scale field applications of cultural desert biocrust was widely anticipated., (© 2020 Wiley-VCH GmbH.)

In conventional light microscopy, the adjacent cell walls of filamentous moss protonemata are seen from its narrow side thereby obscuring the major area of cell-cell connection. Optical sectioning, segmentation and 3D reconstructions allow the tilting and rotation of intracellular structures thereby greatly improving our understanding of interaction between organelles, membranes and the cell wall. Often, the findings also allow for conclusions on the respective functions. The moss Physcomitrium ( Physcomitrella ) patens is a model organism for growth, development and morphogenesis. Its filamentous protonemata are ideal objects for microscopy. Here, we investigated the cell wall between two neighboring cells and the connection of membranes towards this wall after plasmolysis in 0.8 M mannitol. An m-green fluorescent protein (GFP)-HDEL cell line was used to visualize the endoplasmatic reticulum (ER), the plasma membrane (PM) was stained with FM4-64. Our studies clearly show the importance of cell-cell contacts in P. patens protonemata. In 86% of the investigated cell pairs, at least one of the protoplasts remained fully attached to the adjacent cell wall. By tilting of z-stacks, volume renderings and 3D reconstructions, we visualized the amount of attached/detached PM and ER components after plasmolysis and membrane piercings through the wall of cell neighbors.

Understanding the evolution events defining silicon (Si) uptake in plant species is important for the efficient exploration of Si-derived benefits. In the present study, Si accumulation was studied in 456 diverse plant species grown in uniform field conditions, and in a subset of 151 species grown under greenhouse conditions, allowing efficient comparison among the species. In addition, a systematic analysis of nodulin 26-like intrinsic proteins III (NIP-III), which form Si channels, was performed in >1000 species to trace their evolutionary path and link with Si accumulation. Significant variations in Si accumulation were observed among the plant species studied. For their part, species lacking NIP-IIIs systematically showed low Si accumulation. Interestingly, seven NIP-IIIs were identified in three moss species, namely Physcomitrella patens, Andreaea rupestris, and Scouleria aquatica, indicating that the evolution of NIP-IIIs dates back as early as 515 million years ago. These results were further supported from previous reports of Si deposition in moss fossils estimated to be from around the Ordovician era. The taxonomical distribution provided in the present study will be helpful for several other disciplines, such as palaeoecology and geology, that define the biogeochemical cycling of Si. In addition to the prediction of Si uptake potential of plant species based on sequence information and taxonomical positioning, the evolutionary path of the Si uptake mechanism described here will be helpful to understand the Si environment over the different eras of land plant evolution., (© The Author(s) 2020. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Chemical investigation of the secondary metabolites of the whole plant of bryophyte Hypnum plumaeforme Wilson led to the isolation of a new pimarane-type diterpenoid, momilactone F (1), along with seventeen known compounds. Their chemical structures were elucidated based on massive spectroscopic data. The allelopathic and antifungal properties were evaluated. Among them, momilactone F (1), acrenol (2), [11] momilactones A (3) and B (4) showed significant allelopathic activity against Samolus parviflorus Raf. and Lactuca sativa L. var. angustana Irish, as well as selected antifungal property against crop pathogenic fungi strains. On the other hand, 8(14)-podocarpen-13-on-18-oic acid (8) exhibited strong promoting activity on the growth of L. sativa L. var. angustana Irish. The present investigation provided new insights for developing of H. plumaeforme for further application as a potential agricultural tool., (© 2020 Wiley-VHCA AG, Zurich, Switzerland.)

We conducted laboratory experiments to determine the lethal temperatures of the shoots of dried Bryum argenteum and to determine how this restoration species responds to extreme environments. We specifically assessed changes in gene expression levels in the shoots of dried B. argenteum plants that were subjected to sudden heat shock (control (20 ± 2°C), 80°C, 100°C, 110°C or 120°C) followed by exposure to heat for an additional 10, 20, 30 or 60 min. After they were exposed to heat, the samples were placed in wet sand medium, and their survival and regeneration abilities were evaluated daily for 56 days. The results showed that lethal temperatures significantly reduced the shoot regeneration potential, delayed both shoot and protonemal emergence times and reduced the protonemal emergence area. In addition, the expression of nine genes (HSF3, HSP70, ERF, LEA, ELIP, LHCA, LHCB, Tr288 and DHN) was induced by temperature stress, as assessed after 30 min of exposure. Additionally, a new thermal tolerance level for dried B. argenteum - 120°C for 20 min - was determined, which was the highest temperature recorded for this moss; this tolerance exceeded the previous record of 110°C for 10 min. These findings help elucidate the survival mechanism of this species under heat shock stress and facilitate the recovery and restoration of destroyed ecosystems., (Copyright © 2020 Xinjiang institute of ecology and geography, the CAS. Published by Elsevier Ltd.. All rights reserved.)

Background: NADP-malic enzyme (NAPD-ME), and pyruvate orthophosphate dikinase (PPDK) are important enzymes that participate in C 4 photosynthesis. However, the evolutionary history and forces driving evolution of these genes in C 4 plants are not completely understood., Results: We identified 162 NADP-ME and 35 PPDK genes in 25 species and constructed respective phylogenetic trees. We classified NADP-ME genes into four branches, A1, A2, B1 and B2, whereas PPDK was classified into two branches in which monocots were in branch I and dicots were in branch II. Analyses of selective pressure on the NAPD-ME and PPDK gene families identified four positively selected sites, including 94H and 196H in the a5 branch of NADP-ME, and 95A and 559E in the e branch of PPDK at posterior probability thresholds of 95%. The positively selected sites were located in the helix and sheet regions. Quantitative RT-PCR (qRT-PCR) analyses revealed that expression levels of 6 NADP-ME and 2 PPDK genes from foxtail millet were up-regulated after exposure to light., Conclusion: This study revealed that positively selected sites of NADP-ME and PPDK evolution in C 4 plants. It provides information on the classification and positive selection of plant NADP-ME and PPDK genes, and the results should be useful in further research on the evolutionary history of C 4 plants.

The early light-induced proteins (ELIPs) are postulated to act as transient pigment-binding proteins that protect the chloroplast from photodamage caused by excessive light energy. Desert mosses such as Syntric h ia caninervis , that are desiccation-tolerant and homoiochlorophyllous, are often exposed to high-light conditions when both hydrated and dry ELIP transcripts are accumulated in response to dehydration. To gain further insights into ELIP gene function in the moss S. caninervis , two ELIP cDNAs cloned from S. caninervis , ScELIP1 and ScELIP2 and both sequences were used as the basis of a transcript abundance assessment in plants exposed to high-light, UV-A, UV-B, red-light, and blue-light. ScELIPs were expressed separately in an Arabidopsis ELIP mutant Atelip . Transcript abundance for ScELIP s in gametophytes respond to each of the light treatments, in similar but not in identical ways. Ectopic expression of either ScELIPs protected PSII against photoinhibition and stabilized leaf chlorophyll content and thus partially complementing the loss of AtELIP 2. Ectopic expression of ScELIPs also complements the germination phenotype of the mutant and improves protection of the photosynthetic apparatus of transgenic Arabidopsis from high-light stress. Our study extends knowledge of bryophyte photoprotection and provides further insight into the molecular mechanisms related to the function of ELIPs., Competing Interests: The authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript, or in the decision to publish the results.

Plants live in close association with microorganisms that can have beneficial or detrimental effects. The activity of bacteria in association with flowering plants has been extensively analysed. Bacteria use quorum-sensing as a way of monitoring their population density and interacting with their environment. A key group of quorum sensing molecules in Gram-negative bacteria are the N-acylhomoserine lactones (AHLs), which are known to affect the growth and development of both flowering plants, including crops, and marine algae. Thus, AHLs have potentially important roles in agriculture and aquaculture. Nothing is known about the effects of AHLs on the earliest-diverging land plants, thus the evolution of AHL-mediated bacterial-plant/algal interactions is unknown. In this paper, we show that AHLs can affect spore germination in a representative of the earliest plants on land, the Bryophyte moss Physcomitrella patens. Furthermore, we demonstrate that sporophytes of some wild isolates of Physcomitrella patens are associated with AHL-producing bacteria.

Nodulin 26-like intrinsic proteins (NIPs) play essential roles in transporting the nutrients silicon and boron in seed plants, but the evolutionary origin of this transport function and the co-permeability to toxic arsenic remains enigmatic. Horizontal gene transfer of a yet uncharacterised bacterial AqpN-aquaporin group was the starting-point for plant NIP evolution. We combined intense sequence, phylogenetic and genetic context analyses and a mutational approach with various transport assays in oocytes and plants to resolve the transorganismal and functional evolution of bacterial and algal and terrestrial plant NIPs and to reveal their molecular transport specificity features. We discovered that aqpN genes are prevalently located in arsenic resistance operons of various prokaryotic phyla. We provided genetic and functional evidence that these proteins contribute to the arsenic detoxification machinery. We identified NIPs with the ancestral bacterial AqpN selectivity filter composition in algae, liverworts, moss, hornworts and ferns and demonstrated that these archetype plant NIPs and their prokaryotic progenitors are almost impermeable to water and silicon but transport arsenic and boron. With a mutational approach, we demonstrated that during evolution, ancestral NIP selectivity shifted to allow subfunctionalisations. Together, our data provided evidence that evolution converted bacterial arsenic efflux channels into essential seed plant nutrient transporters., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

Mosses are frequently used to monitor atmospheric metal contamination but few studies on metal adsorption under controlled conditions are available. Here, the accumulation of the heavy metals copper and zinc was studied in the acrocarp moss Atrichum undulatum. An in vitro culture of A. undulatum was established and the same line, size and equally old remets were exposed to six different treatments representing various metal exposure times and washing scenarios as rain simulation. The metal treatments were done in copper and zinc salts (Cu-acetate, CuSO 4 , ZnSO 4 and ZnCl 2 , respectively). Energy-Dispersive X-ray microanalysis (EDX) was employed to detect bound heavy metals on the moss plantlets. Element distribution in stems and leaves was measured separately. The aqueous solution of metal salts facilitated an adsorption of both elements in the moss tissue as compared to solid medium. Furthermore, A. undulatum can tolerate pollution of zinc and copper in a distinctive extent; our data point towards a higher zinc tolerance whereas copper is rather harmful. However, semi-quantitatively, less zinc was detected within the moss tissue compared to copper. Interestingly, a strong positive correlation between the accumulation of copper/zinc and iron, and a strong negative correlation between copper/zinc and magnesium, respectively, was documented., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The effects of biological soil crusts (BSC) on vascular plant growth can be positive, neutral or negative, and little information is available on the impacts of different BSC successional stages on vascular plant population dynamics. We analysed seedling emergence, survival, plant growth and reproduction in response to different BSC successional stages (i.e. habitats: bare soil, cyanobacteria, lichen and moss crusts) in natural populations of Echinops gmelinii Turcz. in the Tengger Desert of northwest China. The winter annual E. gmelinii is a dominant pioneer herb after sand stabilisation. During the early stages of BSC succession, the studied populations of E. gmelinii were characterised by high density, plant growth and fecundity. As the BSC succession proceeded beyond moss crusts, the fecundity decreased sharply, which limited seedling recruitment. Differences in seedling survival among the successional stages were not evident, indicating that BSC have little effect on survival in arid desert regions. Moreover, E. gmelinii biomass allocation exhibited low plasticity, and only reproductive allocation was sensitive to the various habitats. Our results further suggest that the negative effects of BSC succession on population dynamics are primarily driven by increasing topsoil water-holding capacity and decreasing rain water infiltration into deeper soil. We conclude that BSC succession drives population dynamics of E. gmelinii, primarily via its effect on soil moisture. The primary cause for E. gmelinii population decline during the moss-dominated stage of BSC succession is decreased fecundity of individual plants, with declining seed mass possibly reducing the success of seedling establishment., (© 2019 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.)

Moss is usually as an initial colonizer in alpine glacier retreated regions. We hypothesized that moss can significantly facilitate the toxic metals accumulation in alpine ecosystems based on its strong ability of absorption and the role in soil development. Hence, we investigated the trace element pool sizes and enrichment factors, especially for mercury (Hg) by using the Hg isotopic compositions to determine the source contributions in a moss-dominated ecosystem over glacial erratic in Eastern Tibetan Plateau. Results show that Hg, lead (Pb) and cadmium (Cd) are highly enriched in organic soils. Specifically, Cd concentration is 5-20 times higher than the safety limit of the acid soil (pH ≤ 5.5) in China. Atmospheric depositions dominantly contribute to the Pb and Cd sources in organic soils, and followed by the moraine particles influences. The lowering pH in organic soils increasing with glacial retreated time results in the desorption of Cd in organic soils. Atmospheric Hg 0 uptake by moss predominantly contributes to the Hg sources in organic soils. The average Pb accumulation rate over last 125-year is about 5.6 ± 1.0 mg m -2 yr -1 , and for Cd is 0.4 ± 0.1 mg m -2 yr -1 , and for Hg 0 is 27.6 ± 3.2 μg m -2 yr -1 . These elevated accumulation rates are caused by the high moss biomass and elevated atmospheric Hg, Pb and Cd pollution levels in China and neighbouring regions. Our study indicates that the moss not only as the bioindicator, but also plays an important role in the hazardous metal biogeochemical cycling in alpine regions., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Deposition of reactive nitrogen (N) is a major threat to terrestrial ecosystems associated with impacts on ecosystem properties and functions including carbon (C) and nutrient stocks, soil water quality and nutrient retention. In the oceanic-alpine Racomitrium heath habitat, N deposition is associated with moss mat degradation and a shift from bryophyte to graminoid dominance. To investigate the effects of moss mat decline on C and N stocks and fluxes, we collected Racomitrium heath vegetation/soil cores from sites along a gradient of N deposition in the UK. Cores were maintained under controlled conditions and exposed to scenarios of current (8-40 kg N ha -1 y -1 ), reduced (8 kg N ha -1 y -1 ) and elevated (50 kg N ha -1 y -1 ) N deposition. Cores from high N deposition sites had smaller aboveground C and N stocks and, under current conditions, leached large amounts of inorganic N and had low soil water pH compared with low N deposition sites. With reduced N deposition there was evidence for rapid recovery of soil water quality in terms of reduced N leaching and small increases in pH. Under high N deposition, cores from low N deposition sites retained much of the applied N while those with a history of high N deposition leached large amounts of inorganic N. Carbon fluxes in soil water and net CO 2 fluxes varied according to core source site but were not affected by the N deposition scenarios. We conclude that C and N stocks and cycling in Racomitrium heath are strongly affected by long-term exposure to N deposition but that soil water quality may improve rapidly, if N deposition rates are reduced. The legacy of N deposition impacts on moss mat cover and vegetation composition however, mean that the ecosystem remains sensitive to future pulses in N input., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Efforts to reveal ancestral functions of auxin, a key regulator of plant growth and development, and its importance for evolution have been hampered by a fragmented picture of auxin response domains in early-diverging land plants. We report the mapping of auxin sensing and responses during vegetative moss development using novel reporters. We established a moss-specific ratiometric reporter (PpR2D2) for Auxin Response Element- and AUXIN RESPONSE FACTOR-independent auxin sensing in Physcomitrella patens, and its readout during vegetative development was compared with new promoter-based GmGH3::GFPGUS and DR5revV2::GFPGUS auxin response reporters. The ratiometric reporter responds rapidly to auxin in a time-, dose- and TRANSPORT INHIBITOR RESISTANT1/AUXIN F-BOX-dependent manner and marks known, anticipated and novel auxin sensing domains. It reveals proximal auxin sensing maxima in filamentous tissues and sensing minima in all five vegetative gametophytic stem cell types as well as dividing cells. PpR2D2 readout is compliant with an ancestral function of auxin as a positive regulator of differentiation vs proliferation in stem cell regions. The PpR2D2 reporter is a sensitive tool for high-resolution mapping of auxin sensing, which can increase our knowledge of auxin function in early-diverging land plants substantially, thereby advancing our understanding of its importance for plant evolution., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

Plants undergo a phase transition from vegetative to reproductive development that triggers floral induction. Genes containing an AAI (α-amylase inhibitor) domain form a large gene family, but there have been no comprehensive analyses of this gene family in any plant species. Here, we identified 336 AAI genes from nine plant species including122 AAI genes in cotton (Gossypium hirsutum). The AAI gene family has evolutionarily conserved amino acid residues throughout the plant kingdom. Phylogenetic analysis classified AAI genes into five major clades with significant polyploidization and showing effects of genome duplication. Our study identified 42 paralogous and 216 orthologous gene pairs resulting from segmental and whole-genome duplication, respectively, demonstrating significant contributions of gene duplication to expansion of the cotton AAI gene family. Further, GhAAI66 was preferentially expressed in flower tissue and as responses to phytohormone treatments. Ectopic expression of GhAAI66 in Arabidopsis and silencing in cotton revealed that GhAAI66 triggers a phase transition to induce early flowering. Further, GO (Gene Ontology) and KEGG (Kyoto Encyclopedia of Genes and Genomes) analysis of RNA sequencing data and qRT-PCR (quantitative reverse transcription-PCR) analysis indicated that GhAAI66 integrates multiple flower signaling pathways including gibberellin, jasmonic acid, and floral integrators to trigger an early flowering cascade in Arabidopsis. Therefore, characterization of the AAI family provides invaluable insights for improving cotton breeding., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Multiple adaptations were necessary when plants conquered the land. Among them were soluble phenylpropanoids related to plant protection and lignin necessary for upright growth and long-distance water transport. Cytochrome P450 monooxygenase 98 (CYP98) catalyzes a rate-limiting step in phenylpropanoid biosynthesis. Phylogenetic reconstructions suggest that a single copy of CYP98 founded each major land plant lineage (bryophytes, lycophytes, monilophytes, gymnosperms and angiosperms), and was maintained as a single copy in all lineages but the angiosperms. In angiosperms, a series of independent gene duplications and losses occurred. Biochemical assays in four angiosperm species tested showed that 4-coumaroyl-shikimate, a known intermediate in lignin biosynthesis, was the preferred substrate of one member in each species, while independent duplicates in Populus trichocarpa and Amborella trichopoda each showed broad substrate ranges, accepting numerous 4-coumaroyl-esters and -amines, and were thus capable of producing a wide range of hydroxycinnamoyl conjugates. The gymnosperm CYP98 from Pinus taeda showed a broad substrate range, but preferred 4-coumaroyl-shikimate as its best substrate. In contrast, CYP98s from the lycophyte Selaginella moellendorffii and the fern Pteris vittata converted 4-coumaroyl-shikimate poorly in vitro, but were able to use alternative substrates, in particular 4-coumaroyl-anthranilate. Thus, caffeoyl-shikimate appears unlikely to be an intermediate in monolignol biosynthesis in non-seed vascular plants, including ferns. The best substrate for CYP98A34 from the moss Physcomitrella patens was also 4-coumaroyl-anthranilate, while 4-coumaroyl-shikimate was converted to lower extents. Despite having in vitro activity with 4-coumaroyl-shikimate, CYP98A34 was unable to complement the Arabidopsis thaliana cyp98a3 loss-of-function phenotype, suggesting distinct properties also in vivo., (© 2019 The Authors The Plant Journal © 2019 John Wiley & Sons Ltd.)