Synergistic effects of zinc and cadmium on phytoremediation potential of Christmas moss (Vesicularia montagnei).

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.

Quantitative cell biology of tip growth in moss.

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.

Spatial and Temporal Patterns of Endopolyploidy in Mosses.

Somatic polyploidy or endopolyploidy is common in the plant kingdom; it ensures growth and allows adaptation to the environment. It is present in the majority of plant groups, including mosses. Endopolyploidy had only been previously studied in about 65 moss species, which represents less than 1% of known mosses. We analyzed 11 selected moss species to determine the spatial and temporal distribution of endopolyploidy using flow cytometry to identify patterns in ploidy levels among gametophytes and sporophytes. All of the studied mosses possessed cells with various ploidy levels in gametophytes, and four of six species investigated in sporophytic stage had endopolyploid sporophytes. The proportion of endopolyploid cells varied among organs, parts of gametophytes and sporophytes, and ontogenetic stages. Higher ploidy levels were seen in basal parts of gametophytes and sporophytes than in apical parts. Slight changes in ploidy levels were observed during ontogenesis in cultivated mosses; the youngest (apical) parts of thalli tend to have lower levels of endopolyploidy. Differences between parts of cauloid and phylloids of Plagiomnium ellipticum and Polytrichum formosum were also documented; proximal parts had higher levels of endopolyploidy than distal parts. Endopolyploidy is spatially and temporally differentiated in the gametophytes of endopolyploid mosses and follows a pattern similar to that seen in angiosperms.

Bryophyte community diversity and structure associated with Asplenium auritum fern (Aspleniaceae) in a Brazilian Atlantic forest fragment / Diversidad y estructura de la comunidad de briófitos asociada con el helecho Asplenium auritum (Aspleniaceae) en un fragmento del bosque atlántico brasileño

Introducción: Los filtros ambientales locales ejercen influencia en la diversidad y en la estructura de las comunidades de briófitas, pero existen pocos estudios que analicen cómo las relaciones bióticas influyen en estas comunidades. Objetivo: Evaluar la influencia de la cobertura del helecho Asplenium auritum sobre los cambios en la riqueza de especies y la composición de las comunidades briófitos, en un bosque remanente semideciduo en Minas Gerais, Brasil. Métodos: Seleccionamos un transecto de 300 m en cada una de las seis áreas donde A. auritum estaba presente. En cada transecto, estimamos la cubierta vegetal en 39 parcelas de 10 × 10 cm distribuidas al azar. Comparamos la riqueza estimada de especies y la composición de briófitos entre los niveles de cobertura del helecho. Resultados: Encontramos 60 especies en 35 géneros y 23 familias. Las formas de vida más representativas fueron las de tipo trama y estera. Los diferentes niveles de cobertura del helecho no mostraron efecto significativosobre la riqueza de especies y la composición de las comunidades de briófitos. Conclusiones: La cobertura de briófitos parece favorecer el desarrollo del helecho, podría promover la coexistencia de varias especies de briófitos y podría predecir variaciones en la riqueza de especies e impulsar el ensamble de especies en esta comunidad.

Introduction: Local environmental filters have shown the influence on the bryophyte diversity and structure communities, but there are limited studies that analyze how biotic relationships filters influence these communities. Objective: To evaluate whether the influence of fern Asplenium auritum cover determine changes in species richness and composition of bryophyte communities, in a semideciduous remnant forest, Minas Gerais, Brazil. Methods: We selected one transect of 300 m in each of six areas where Asplenium auritum was present. In all transect we estimated plant cover in 39 10 × 10 cm plots randomly distributed. We compare the estimated species richness and composition of bryophyte between fern coverage levels. Results: We found 60 species in 35 genera and 23 families. Weft and mat were the most represented life-forms. Fern cover had no significant effect on bryophyte species richness and composition. Conclusions: Bryophyte cover appears to improve fern development and promote the coexistence of several bryophyte species. Bryophyte cover probably predicts variations in species richness and drives the species assemblage in this community.

An Ancestral Role for CONSTITUTIVE TRIPLE RESPONSE1 Proteins in Both Ethylene and Abscisic Acid Signaling.

Land plants have evolved adaptive regulatory mechanisms enabling the survival of environmental stresses associated with terrestrial life. Here, we focus on the evolution of the regulatory CONSTITUTIVE TRIPLE RESPONSE1 (CTR1) component of the ethylene signaling pathway that modulates stress-related changes in plant growth and development. First, we compare CTR1-like proteins from a bryophyte, Physcomitrella patens (representative of early divergent land plants), with those of more recently diverged lycophyte and angiosperm species (including Arabidopsis [Arabidopsis thaliana]) and identify a monophyletic CTR1 family. The fully sequenced P. patens genome encodes only a single member of this family (PpCTR1L). Next, we compare the functions of PpCTR1L with that of related angiosperm proteins. We show that, like angiosperm CTR1 proteins (e.g. AtCTR1 of Arabidopsis), PpCTR1L modulates downstream ethylene signaling via direct interaction with ethylene receptors. These functions, therefore, likely predate the divergence of the bryophytes from the land-plant lineage. However, we also show that PpCTR1L unexpectedly has dual functions and additionally modulates abscisic acid (ABA) signaling. In contrast, while AtCTR1 lacks detectable ABA signaling functions, Arabidopsis has during evolution acquired another homolog that is functionally distinct from AtCTR1. In conclusion, the roles of CTR1-related proteins appear to have functionally diversified during land-plant evolution, and angiosperm CTR1-related proteins appear to have lost an ancestral ABA signaling function. Our study provides new insights into how molecular events such as gene duplication and functional differentiation may have contributed to the adaptive evolution of regulatory mechanisms in plants.

Hybridization is associated with changes in sexual system in the bryophyte genus Atrichum.

PREMISE OF THE STUDY: Over 50% of bryophytes have separate sexes, and numerous transitions have occurred between combined and separate sexes. Polyploidy and hybridization is one proximate mechanism hypothesized to cause evolutionary transitions to hermaphroditism in bryophytes because sex is expressed at the haploid stage and in nonpolyploid dioecious species males have a single V chromosome and females a U. Hermaphroditism can arise if gametophytes of allopolyploids have both U and V chromosomes. We examined the association between polyploidy and hermaphroditism in the bryophyte genus Atrichum, which has species where gametophytes can be haploid, diploid, or triploid, and some species have hermaphroditic individuals. METHODS: We generated phylogenies of Atrichum from sequences of three plastid regions (rbcL, rps4, and trnL-trnF) and the second intron for the nuclear gene Leafy/Floricaula to further understand the relationships among haploid, diploid, and triploid species, and those with combined or separate sexes. KEY RESULTS: The existence of multiple sequences of Leafy/Floricaula in diploid and triploid, but not haploid, individuals is consistent with independent allopolyploid origins of the diploid and triploid species. Allopolyploidy was associated with a likely gain in hermaphroditism in triploid Atrichum undulatum and possibly diploid A. altecristatum, but not in the allopolyploid A. crispulum (diploid at the gametophyte level). CONCLUSIONS: These results highlight a role for hybridization and polyploidy in sexual system evolution, but the presence of diploid (allopolyploid) dioecious species suggest that other factors may influence the maintenance of sexual systems after an evolutionary transition.

Chryseobacterium polytrichastri sp. nov., isolated from a moss (Polytrichastrum formosum), and emended description of the genus Chryseobacterium.

A Gram-stain negative, rod-shaped and non-endospore forming bacterium, designated strain YG4-6(T), was isolated from Polytrichastrum formosum collected from Gawalong glacier in Tibet, China and characterized by using a polyphasic taxonomic approach. The predominant fatty acids of strain YG4-6(T) were identified as iso-C15:0 (29.3 %), summed feature 3 (C16:1 ω7c and/or C16:1 ω6c as defined by MIDI, 23.5 %) and iso-C17:0 3-OH (16.5 %). The major polar lipids were found to consist of five unidentified aminolipids and three unidentified lipids. Strain YG4-6(T) was found to contain MK-6 as the dominant menaquinone and the G+C content of its genomic DNA was determined to be 37.3 mol%. The phylogenetic analysis based on 16S rRNA gene sequences showed that strain YG4-6(T) is affiliated to Chryseobacterium species, and its closest related species were Chryseobacterium aahli T68(T) (97.9 % sequence similarity), Chryseobacterium zeae JM-1085(T) (97.8 % sequence similarity), Chryseobacterium yeoncheonense DCY67(T) (97.6 % sequence similarity) and Chryseobacterium soldanellicola NBRC 100864(T) (97.2 % sequence similarity). However, the DNA-DNA relatedness values between these strains and strain YG4-6(T) were found to be clearly below 70 %. Based on the phylogenetic inference and phenotypic data, strain YG4-6(T) is considered to represent a novel species of the genus Chryseobacterium, for which the name Chryseobacterium polytrichastri sp. nov. is proposed. The type strain is YG4-6(T) (=CGMCC 1.12491(T) = DSM 26899(T)). An emended description of the genus Chryseobacterium is also proposed.

Polyphenol oxidases in Physcomitrella: functional PPO1 knockout modulates cytokinin-dependent development in the moss Physcomitrella patens.

Polyphenol oxidases (PPOs) are copper-binding enzymes of the plant secondary metabolism that oxidize polyphenols to quinones. Although PPOs are nearly ubiquitous in seed plants, knowledge on their evolution and function in other plant groups is missing. This study reports on the PPO gene family in the moss Physcomitrella patens (Hedw.) B.S.G. asan example for an early divergent plant. The P. patens PPO multigene family comprises 13 paralogues. Phylogenetic analyses suggest that plant PPOs evolved with the colonization of land and that PPO duplications within the monophyletic P. patens paralogue clade occurred after the separation of the moss and seed plant lineages. PPO functionality was demonstrated for recombinant PPO6. P. patens was analysed for phenolic compounds and six substances were detected intracellularly by LC-MS analysis: 4-hydroxybenzoic acid, p-cumaric acid, protocatechuic acid, salicylic acid, caffeic acid, and an ester of caffeic acid. Targeted PPO1 knockout (d|ppo1) plants were generated and plants lacking PPO1 exhibited only ~30% of the wild-type PPO activity in the culture medium, thus suggesting extracellular localization of PPO1, which is in contrast to the mostly plastidic PPO localization in seed plants. Further, d|ppo1 lines formed significantly more gametophores with a reduced areal plant size, which could be related to an increase of endogenously produced cytokinins and indicates an impact of PPO1 on plant development. d|ppo1 plants were less tolerant towards applied 4-methylcatechol compared to the wild type, which suggests a role of extracellular PPO1 in establishing appropriate conditions by the removal of inhibitory extracellular phenolic compounds.

Impacts of oil sands process water on fen plants: implications for plant selection in required reclamation projects.

Fen plant growth in peat contaminated with groundwater discharges of oil sands process water (OSPW) was assessed in a greenhouse over two growing seasons. Three treatments (non-diluted OSPW, diluted OSPW and rainwater) were tested on five vascular plants and four mosses. All vascular plants tested can grow in salinity and naphthenic acids levels currently produced by oil sands activity in northwestern Canada. No stress sign was observed after both seasons. Because of plant characteristics, Carex species (C. atherodes and C. utriculata) and Triglochin maritima would be more useful for rapidly restoring vegetation and creating a new peat-accumulating system. Groundwater discharge of OSPW proved detrimental to mosses under dry conditions and ensuring adequate water levels would be crucial in fen creation following oil sands exploitation. Campylium stellatum would be the best choice to grow in contaminated areas and Bryum pseudotriquetrum might be interesting as it has spontaneously regenerated in all treatments.

Practical application of methanol-mediated mutualistic symbiosis between Methylobacterium species and a roof greening moss, Racomitrium japonicum.

Bryophytes, or mosses, are considered the most maintenance-free materials for roof greening. Racomitrium species are most often used due to their high tolerance to desiccation. Because they grow slowly, a technology for forcing their growth is desired. We succeeded in the efficient production of R. japonicum in liquid culture. The structure of the microbial community is crucial to stabilize the culture. A culture-independent technique revealed that the cultures contain methylotrophic bacteria. Using yeast cells that fluoresce in the presence of methanol, methanol emission from the moss was confirmed, suggesting that it is an important carbon and energy source for the bacteria. We isolated Methylobacterium species from the liquid culture and studied their characteristics. The isolates were able to strongly promote the growth of some mosses including R. japonicum and seed plants, but the plant-microbe combination was important, since growth promotion was not uniform across species. One of the isolates, strain 22A, was cultivated with R. japonicum in liquid culture and in a field experiment, resulting in strong growth promotion. Mutualistic symbiosis can thus be utilized for industrial moss production.

Environmental effects of oil and gas lease sites in a grassland ecosystem.

The northern Great Plains of Saskatchewan is one of the most significantly modified landscapes in Canada. While the majority of anthropogenic disturbances to Saskatchewan's grasslands are the result of agricultural practices, development of petroleum and natural gas (PNG) resources is of increasing concern for grassland conservation. Although PNG developments require formal assessment and regulatory approval, follow-up and monitoring of the effects of PNG development on grasslands is not common practice. Consequently, the effects of PNG activity on grasslands and the spatial and temporal extent of such impacts are largely unknown. This paper examines the spatial and temporal extent of PNG development infrastructure from 1955 to 2006 in a grassland ecosystem in southwest Saskatchewan. The effects of PNG development on grassland ecology were assessed from measurements of ground cover characteristics, soil properties, and plant community composition at 31 sites in the study area. PNG lease sites were found to have low cover of herbaceous plants, club moss (Lycopodiaceae), litter, and shallow organic (Ah) horizons. Lease sites were also characterized by low diversity of desirable grassland plants and low range health values compared to off-lease reference sites. These impacts were amplified at active and highly productive lease sites. Impacts of PNG development persisted for more than 50 years following well site construction, and extended outward 20 m-25 m beyond the direct physical footprint of PNG well infrastructure. These results have significant implications with regard to the current state of monitoring and follow-up of PNG development, and the cumulative effective of PNG activity on grassland ecosystems over space and time.

[Effects of enhanced UV-B radiation on the growth of five bryophytes in Changbai Mountains].

Five bryophytes (Rhytidium rugosum, Rhytidiadelphus triquetrus, Hylocomium splendens, Hylocomium pyrenaicum, and Polytrichum alpinum) were exposed to 0.2 kJ x m(-2) x d(-1) (visible light under native condition, CK), 3.0 kJ x m(-2) x d(-1) (simulated dose of UV-B irradiance at the tundra in Changbai Mountains, medium dose of UV-B irradiance, T1), and 6.0 kJ x m(-2) x d(-1) (high dose of UV-B irradiance, T2) to investigate the effects of enhanced UV-B radiation on plant height, biomass, and chlorophyll content. The results indicated that medium and high UV-B radiations decreased the plant height, biomass, and chlorophyll content of R. triquetrus and H. splendens by 32.3%, 62.4%, and 81.3%, and 21.4%, 59.4%, and 62.8%, respectively, and the relative growth rates were negative. Enhanced UV-B radiations had less effect on P. alpinum chlorophyll content but doubled its below-ground biomass, and slightly increased the biomass of R. rugosum. P. alpinum and R. rugosum had higher tolerance against UV-B radiation, while R. triquetrus and H. splendens were more sensitive to UV-B radiation.

A new species of Cladophialophora (hyphomycetes) from boreal and montane bryophytes.

During a survey of bryophilous fungi from boreal and montane habitats in central Alberta, a hitherto undescribed species of Cladophialophora was recovered from Polytrichum juniperinum, Aulacomnium palustre, and Sphagnum fuscum. On potato dextrose agar (PDA) colonies grew slowly, attaining a diameter of 25 mm after 30 d, were dark grey, velvety, radially sulcate, and convolute and cracked at the centre. Micronematous conidiophores gave rise to branched chains of small (1-2 x 8-22 microm), cylindrical to fusiform conidia with truncate, swollen scars at each end. Phylogenies built on the ITS and ribosomal SSU regions indicate the isolates form a monophyletic clade within the family Herpotrichiellaceae (Chaetothyriales) that is composed of two geographically based groups, each with 99% within-group sequence similarity and 97-98% between-group sequence similarity. A teleomorph has not been found but would likely be similar to species of Capronia. In vitro inoculation of the isolates onto axenically grown P. juniperinum produced no discernible host symptoms, and host penetration could not be detected using light microscopy. The production of polyphenol oxidases by the fungus and the role of other Cladophialophora species as latent endophytes and saprobes suggest that a potential role for the fungus is the degradation of the polyphenol-rich cell walls of mosses. A dichotomous key to species of the genus Cladophialophora is provided.

Molecular cloning and characterization of a sodium-pump ATPase of the moss Physcomitrella patens.

Physcomitrella patens grew slowly at 600 mm Na+, pH 6.0, affected by the low water potential but without signs of suffering Na+ toxicity. At pH 8.0, tolerance seemed to be lower but it grew at 200 mm Na+, again without signs of Na+ toxicity. The resistance of Physcomitrella cells to the toxic effects of Na+ can be accounted for by their capacity to keep high K+:Na+ ratios and to extrude Na+ by a system that is not dependent on DeltapH. Physcomitrella expresses two P-type ATPases similar in sequence to fungal ENA-type Na+-ATPases. A functional study in yeast demonstrated that one of these ATPases, PpENA1, is an Na+-pump. We also found that P. patens has a plant-type SOS1 Na+/H+ antiporter. We discuss that Na+-ATPases existed in early land plants but that they were lost during the evolution of bryophytes to flowering plants.