Sphagnum growth under N saturation: interactive effects of water level and P or K fertilization.

Sphagnum biomass is a promising material that could be used as a substitute for peat in growing media and can be sustainably produced by converting existing drainage-based peatland agriculture into wet, climate-friendly agriculture (paludiculture). Our study focuses on yield maximization of Sphagnum as a crop. We tested the effects of three water level regimes and of phosphorus or potassium fertilization on the growth of four Sphagnum species (S. papillosum, S. palustre, S. fimbriatum, S. fallax). To simulate field conditions in Central and Western Europe we carried out a glasshouse experiment under nitrogen-saturated conditions. A constant high water table (remaining at 2 cm below capitulum during growth) led to highest productivity for all tested species. Water table fluctuations between 2 and 9 cm below capitulum during growth and a water level 2 cm below capitulum at the start but falling relatively during plant growth led to significantly lower productivity. Fertilization had no effect on Sphagnum growth under conditions with high atmospheric deposition such as in NW Germany (38 kg N, 0.3 kg P, 7.6 kg K·ha-1 ·year-1 ). Large-scale maximization of Sphagnum yields requires precise water management, with water tables just below the capitula and rising with Sphagnum growth. The nutrient load in large areas of Central and Western Europe from atmospheric deposition and irrigation water is high but, with an optimal water supply, does not hamper Sphagnum growth, at least not of regional provenances of Sphagnum.

Response of C and N cycles to N fertilization in Sphagnum and Molinia-dominated peat mesocosms.

Plant communities play an important role in the C-sink function of peatlands. However, global change and local perturbations are expected to modify peatland plant communities, leading to a shift from Sphagnum mosses to vascular plants. Most studies have focused on the direct effects of modification in plant communities or of global change (such as climate warming, N fertilization) in peatlands without considering interactions between these disturbances that may alter peatlands' C function. We set up a mesocosm experiment to investigate how Greenhouse Gas (CO2, CH4, N2O) fluxes, and dissolved organic carbon (DOC) and total dissolved N (TN) contents are affected by a shift from Sphagnum mosses to Molinia caerulea dominated peatlands combined with N fertilization. Increasing N deposition did not alter the C fluxes (CO2 exchanges, CH4 emissions) or DOC content. The lack of N effect on the C cycle seems due to the capacity of Sphagnum to efficiently immobilize N. Nevertheless, N supply increased the N2O emissions, which were also controlled by the plant communities with the presence of Molinia caerulea reducing N2O emissions in the Sphagnum mesocosms. Our study highlights the role of the vegetation composition on the C and N fluxes in peatlands and their responses to the N deposition. Future research should now consider the climate change in interaction to plants community modifications due to their controls of peatland sensitivity to environmental conditions.

Evidence on the effectiveness of mosses for biomonitoring of microplastics in fresh water environment.

Mosses are well known as biomonitors of fresh water for metal pollutants, but no studies were reported so far about their ability to intercept plastic particles, although this kind of pollution has become an urgent issue worldwide. In the present work, the interaction between the moss Sphagnum palustre L. cultured in vitro and polystyrene nanoparticles (NPs) was studied for the first time in a laboratory experiment, in the view of using moss transplants for detecting microplastics in fresh water environments. The ability of S. palustre to intercept and retain polystyrene, and the effects of vitality and post-exposure washing on NP retention by moss were tested. Fluorescence microscope observations showed that polystyrene NPs were retained by moss leaves in form of small (the most abundant fraction) and large aggregates. Particle count analysis highlighted that the number of particles increased while increasing the exposure time. Moreover, moss devitalization favored NP accumulation, likely because of cell membrane damages occurred in dead moss material. Post-exposure washing induced a loss of larger aggregates, suggesting that exposure time is a key point to be carefully evaluated in field conditions. These results encourage the use of S. palustre transplants for monitoring microplastics contamination of fresh water environments.

Nitrogen deposition does not enhance Sphagnum decomposition.

Long-term additions of nitrogen (N) to peatlands have altered bryophyte growth, species dominance, N content in peat and peat water, and often resulted in enhanced Sphagnum decomposition rate. However, these results have mainly been derived from experiments in which N was applied as ammonium nitrate (NH4NO3), neglecting the fact that in polluted areas, wet deposition may be dominated either by NO3(-) or NH4(+). We studied effects of elevated wet deposition of NO3(-) vs. NH4(+) alone (8 or 56kgNha(-1)yr(-1) over and above the background of 8kgNha(-1)yr(-1) for 5 to 11years) or combined with phosphorus (P) and potassium (K) on Sphagnum quality for decomposers, mass loss, and associated changes in hummock pore water in an ombrotrophic bog (Whim). Adding N, especially as NH4(+), increased N concentration in Sphagnum, but did not enhance mass loss from Sphagnum. Mass loss seemed to depend mainly on moss species and climatic factors. Only high applications of N affected hummock pore water chemistry, which varied considerably over time. Overall, C and N cycling in this N treated bog appeared to be decoupled. We conclude that moss species, seasonal and annual variation in climatic factors, direct negative effects of N (NH4(+) toxicity) on Sphagnum production, and indirect effects (increase in pH and changes in plant species dominance under elevated NO3(-) alone and with PK) drive Sphagnum decomposition and hummock C and N dynamics at Whim.

Using devitalized moss for active biomonitoring of water pollution.

This paper presents the results of an experiment carried out for the first time in situ to select a treatment to devitalize mosses for use in active biomonitoring of water pollution. Three devitalizing treatments for the aquatic moss Fontinalis antipyretica were tested (i.e. oven-drying at 100 °C, oven-drying with a 50-80-100 °C temperature ramp, and boiling in water), and the effects of these on loss of material during exposure of the transplants and on the accumulation of different heavy metals and metalloids were determined. The suitability of using devitalized samples of the terrestrial moss Sphagnum denticulatum to biomonitor aquatic environments was also tested. The structure of mosses was altered in different ways by the devitalizing treatments. Devitalization by boiling water led to significantly less loss of material (p < 0.01) than the oven-drying treatments. However, devitalization by oven-drying with a temperature ramp yielded more stable results in relation to both loss of material and accumulation of elements. With the aim of standardizing the moss bag technique, the use of F. antipyretica devitalized by oven-drying with a temperature ramp is recommended, rather than other devitalization treatments or use of S. denticulatum.

Desiccation tolerance of Sphagnum revisited: a puzzle resolved.

As ecosystem engineers, Sphagnum mosses control their surroundings through water retention, acidification and peat accumulation. Because water retention avoids desiccation, sphagna are generally intolerant to drought; however, the literature on Sphagnum desiccation tolerance (DT) provides puzzling results, indicating the inducible nature of their DT. To test this, various Sphagnum species and other mesic bryophytes were hardened to drought by (i) slow drying; (ii) ABA application and (iii) chilling or frost. DT tolerance was assessed as recovery of chlorophyll fluorescence parameters after severe desiccation. We monitored the seasonal course of DT in bog bryophytes. Under laboratory conditions, following initial de-hardening, untreated Sphagnum shoots lacked DT; however, DT was induced by all hardening treatments except chilling, notably by slow drying, and in Sphagnum species of the section Cuspidata. In the field, sphagna in hollows and lawns developed DT several times during the growing season, responding to reduced precipitation and a lowered water table. Hummock and aquatic species developed DT only in late autumn, probably as a response to frost. Sphagnum protonemata failed to develop DT; hence, desiccation may limit Sphagnum establishment in drier habitats with suitable substrate chemistry. Desiccation avoiders among sphagna form compact hummocks or live submerged; thus, they do not develop DT in the field, lacking the initial desiccation experience, which is frequent in hollow and lawn habitats. We confirmed the morpho-physiological trade-off: in contrast to typical hollow sphagna, hummock species invest more resources in water retention (desiccation avoidance), while they have a lower ability to develop physiological DT.

Spatio-temporal trends of nitrogen deposition and climate effects on Sphagnum productivity in European peatlands.

To quantify potential nitrogen (N) deposition impacts on peatland carbon (C) uptake, we explored temporal and spatial trends in N deposition and climate impacts on the production of the key peat forming functional group (Sphagnum mosses) across European peatlands for the period 1900-2050. Using a modelling approach we estimated that between 1900 and 1950 N deposition impacts remained limited irrespective of geographical position. Between 1950 and 2000 N deposition depressed production between 0 and 25% relative to 1900, particularly in temperate regions. Future scenarios indicate this trend will continue and become more pronounced with climate warming. At the European scale, the consequences for Sphagnum net C-uptake remained small relative to 1900 due to the low peatland cover in high-N areas. The predicted impacts of likely changes in N deposition on Sphagnum productivity appeared to be less than those of climate. Nevertheless, current critical loads for peatlands are likely to hold under a future climate.

Assessment of potential climate change impacts on peatland dissolved organic carbon release and drinking water treatment from laboratory experiments.

Catchments draining peat soils provide the majority of drinking water in the UK. Over the past decades, concentrations of dissolved organic carbon (DOC) have increased in surface waters. Residual DOC can cause harmful carcinogenic disinfection by-products to form during water treatment processes. Increased frequency and severity of droughts combined with and increased temperatures expected as the climate changes, have potentials to change water quality. We used a novel approach to investigate links between climate change, DOC release and subsequent effects on drinking water treatment. We designed a climate manipulation experiment to simulate projected climate changes and monitored releases from peat soil and litter, then simulated coagulation used in water treatment. We showed that the 'drought' simulation was the dominant factor altering DOC release and affected the ability to remove DOC. Our results imply that future short-term drought events could have a greater impact than increased temperature on DOC treatability.

Glasshouse vs field experiments: do they yield ecologically similar results for assessing N impacts on peat mosses?

• Peat bogs have accumulated more atmospheric carbon (C) than any other terrestrial ecosystem today. Most of this C is associated with peat moss (Sphagnum) litter. Atmospheric nitrogen (N) deposition can decrease Sphagnum production, compromising the C sequestration capacity of peat bogs. The mechanisms underlying the reduced production are uncertain, necessitating multifactorial experiments. • We investigated whether glasshouse experiments are reliable proxies for field experiments for assessing interactions between N deposition and environment as controls on Sphagnum N concentration and production. We performed a meta-analysis over 115 glasshouse experiments and 107 field experiments. • We found that glasshouse and field experiments gave similar qualitative and quantitative estimates of changes in Sphagnum N concentration in response to N application. However, glasshouse-based estimates of changes in production--even qualitative assessments-- diverged from field experiments owing to a stronger N effect on production response in absence of vascular plants in the glasshouse, and a weaker N effect on production response in presence of vascular plants compared to field experiments. • Thus, although we need glasshouse experiments to study how interacting environmental factors affect the response of Sphagnum to increased N deposition, we need field experiments to properly quantify these effects.

Mercury accumulation in peatbogs at Czech sites with contrasting pollution histories.

Mercury (Hg) concentrations and accumulation patterns were studied in (210)Pb-dated peat cores from three ombrotrophic sites in the Czech Republic with contrasting emission histories (Novodomské raseliniste, ND, and Bílá Smedá, BS, in the polluted northern parts of the country, and Jezerní slat, JS, in a relatively pristine southern part of the Czech Republic). The Hg concentration varied significantly between sites. Whereas the sites in the northern part of the Czech Republic yielded a range of higher Hg concentrations (50-750 µg kg(-1) for ND and 30-600 µg kg(-1) for BS), a Hg concentration range of 40-220 µg kg(-1) was reported at JS. At the northern localities, the highest Hg concentrations were detected at depths of 5-10 cm, corresponding to the period between the early 1960s until the late 1980s. In contrast, the highest Hg values at JS were observed at a depth of 10-15 cm, corresponding to the period between the early 1950s and the early 1970s. The maximum Hg accumulation rates were approximately 2× higher at the northern localities (ND: 106 µg m(-2)yr(-1), BS: 90 µg m(-2)yr(-1), JS: 43 µg m(-2)yr(-1)). Although a decrease in the Hg concentration can be observed in the youngest segments of all the peat cores, a slight increase in Hg accumulation rates in the most recent peat segments (living Sphagnum moss) has been reported for all three sites (40-44 µg m(-2)yr(-1)), which is approximately 2× higher than in peat bogs in western and northern Europe. This observation may either be related to a real recent increase in Hg emissions in Central Europe (active coal mining and burning and limited Hg pollution control in thermal power plants) or could indicate a preferential Hg binding mechanism in the living moss at the surface of the peat.

Bioaccumulation and glutathione-mediated detoxification of copper and cadmium in Sphagnum squarrosum Crome Samml.

Physiological and biochemical responses, metal bioaccumulation and tolerance potential of Sphagnum squarrosum Crome Samml. to Cu and Cd were studied to determine its bioindication and bioremediation potential. Results suggest that glutathione treatment increases the metal accumulation potential and plays a definite role in heavy metal scavenging. High abundance of Sphagnum in metal-rich sites strongly suggests its high metal tolerance capabilities. This experiment demonstrates that S. squarrosum is able to accumulate and tolerate a high amount of metals and feasibility of its application as bioindicator and remediator test species of metal-contaminated environment.

Direct and indirect effects of ammonia, ammonium and nitrate on phosphatase activity and carbon fluxes from decomposing litter in peatland.

Here we investigate the response of soils and litter to 5 years of experimental additions of ammonium (NH4), nitrate (NO3), and ammonia (NH3) to an ombrotrophic peatland. We test the importance of direct (via soil) and indirect (via litter) effects on phosphatase activity and efflux of CO2. We also determined how species representing different functional types responded to the nitrogen treatments. Our results demonstrate that additions of NO3, NH4 and NH3 all stimulated phosphatase activity but the effects were dependent on species of litter and mechanism (direct or indirect). Deposition of NH3 had no effect on efflux of CO2 from Calluna vulgaris litter, despite it showing signs of stress in the field, whereas both NO3 and NH4 reduced CO2 fluxes. Our results show that the collective impacts on peatlands of the three principal forms of nitrogen in atmospheric deposition are a result of differential effects and mechanisms on individual components.

[Phenotypic and epigenetic adaptation of the moss clone to mercury].

On agar-Knop medium containing 0.5 microM HgCl2 about one third of microregenerants of the clone from the individual gametophyte cell of the moss Pottia intermedia survived and gave rise to protonemal mats. The high survival percentage testifies to epigenetic nature of adaptation. The latter proved to be correlated to the increase of leaf cell number and of peroxidase activity as well as to intensification of activity zone of peroxidase isoform with MM in limits of 66 kD and to appearance of two isoforms of the enzyme on electrophoregrams. The increase of peroxidase activity, though considerably weaker expressed, has been stated at 0.2 microM HgCl2 when practically all regenerants survived and on the mercury-free medium epigenetically adapted regenerants differed from physiologically adapted ones only in intensification of activity zone of peroxidase isoform with 66 kD. This gives reason to regard the adaptation of the regenerants to 0.5 microM HgCl2 as intensified epigenocopy of modification and indicates the generality of mechanisms of both types of adaptation.

Heatwave 2003: high summer temperature, rather than experimental fertilization, affects vegetation and CO2 exchange in an alpine bog.

Nitrogen and phosphorus were added experimentally in a bog in the southern Alps. It was hypothesized that alleviating nutrient limitation will increase vascular plant cover. As a consequence, more carbon will be fixed through higher rates of net ecosystem CO(2) exchange (NEE). The vascular cover did increase at the expense of Sphagnum mosses. However, such vegetation changes were largely independent of the treatment and were probably triggered by an exceptional heatwave in summer 2003. Contrary to the tested hypothesis, NEE was unaffected by the nutrient treatments but was strongly influenced by temperature and water-table depth. In particular, ecosystem respiration in the hot summer of 2003 increased dramatically, presumably owing to enhanced heterotrophic respiration in an increased oxic peat layer. At the end of the experiment, the Sphagnum cover decreased significantly in the nitrogen-fertilized treatment at hummock microhabitats. In the long term, this will imply a proportionally greater accumulation of vascular litter, more easily decomposable than the recalcitrant Sphagnum litter. As a result, rates of carbon fixation may decrease because of stimulated respiration.

Effects of experimental lead pollution on the microbial communities associated with Sphagnum fallax (Bryophyta).

Ecotoxicological studies usually focus on single microbial species under controlled conditions. As a result, little is known about the responses of different microbial functional groups or individual species to stresses. In an aim to assess the response of complex microbial communities to pollution in their natural habitat, we studied the effect of a simulated lead pollution on the microbial community (bacteria, cyanobacteria, protists, fungi, and micrometazoa) living on Sphagnum fallax. Mosses were grown in the laboratory with 0 (control), 625, and 2,500 microg L(-1) of Pb(2+) diluted in a standard nutrient solution and were sampled after 0, 6, 12, and 20 weeks. The biomasses of bacteria, microalgae, testate amoebae, and ciliates were dramatically and significantly decreased in both Pb addition treatments after 6, 12, and 20 weeks in comparison with the control. The biomass of cyanobacteria declined after 6 and 12 weeks in the highest Pb treatment. The biomasses of fungi, rotifers, and nematodes decreased along the duration of the experiment but were not significantly affected by lead addition. Consequently, the total microbial biomass was lower for both Pb addition treatments after 12 and 20 weeks than in the controls. The community structure was strongly modified due to changes in the densities of testate amoebae and ciliates, whereas the relative contribution of bacteria to the microbial biomass was stable. Differences in responses among the microbial groups suggest changes in the trophic links among them. The correlation between the biomass of bacteria and that of ciliates or testate amoebae increased with increasing Pb loading. We interpret this result as an effect on the grazing pathways of these predators and by the Pb effect on other potential prey (i.e., smaller protists). The community approach used here complements classical ecotoxicological studies by providing clues to the complex effect of pollutant-affecting organisms both directly and indirectly through trophic effects and could potentially find applications for pollution monitoring.

Seasonal pattern of metal bioaccumulation and their toxicity on Sphagnum squarrosum.

Present study was undertaken as an attempt to study the effect of pollutants on biological responses of Sphagnum growing at Kainchi, Kumaon hills (Uttranchal). Sphagnum plants of almost identical size, collected from the marked sites of Kainchi in different seasons viz., monsoon, winter, summer and again in monsoon, were analysed for chlorophyll, protein, shoot length and nitrate reductase and peroxidase activities. Maximum chlorophyll, protein, shoots length and nitrate reductase activities were observed during the monsoon while minimum in summers. The abundance of Sphagnum and two other bryophytes, Marchantia and Plagiochasma was also higher in monsoon than in other seasons. The study also indicated that Sphagnum has more bioaccumulation and tolerance potential for heavy metals than Marchantia and Plagiochasma.

Impact of platinum group metals on the environment: a toxicological, genotoxic and analytical chemistry study.

Recent studies show particles of Platinum Group Metals (PGMs); primarily platinum, palladium and rhodium; released from automobile catalytic converters are being deposited alongside roadways. This deposition is leading to increasing concentrations of PGMs in the environment, raising concerns about the environmental impact and toxicity of these elements in living organisms. The objective of this study was to determine how PGMs alter the patterns of growth, development, and physiology by studying the toxicological and genotoxic effects of these metals. Two vastly different species were used as models: plant-a wild wetland common Sphagnum moss, and animal-6-week old rats Sprague-Dawley. Both species were exposed, in controlled environments, to different concentrations of the PGMs. Toxicological and genotoxic effects were determined by assessment of plant growth, animal survival and pathology, and influence on DNA in both models. Our results on the uptake of PGMs by Sphagnum showed significant decreases in plant length and biomass as PGM concentration increased. Histological and pathological analysis of the animal model revealed vacuolization, eosinophil inclusion bodies in adrenal glands, shrinkage of glomeruli in the kidney, and enlargement of white pulp in the spleen. In both models, DNA damage was detected. Chemical analysis using ICP-AES atomic absorption demonstrated accumulation of PGMs in plant tissues at all PGM levels, proportional to concentration.

Ozone effects on the ultrastructure of peatland plants: Sphagnum mosses, Vaccinium oxycoccus, Andromeda polifolia and Eriophorum vaginatum.

BACKGROUND AND AIMS: Ozone effects on peatland vegetation are poorly understood. Since stress responses are often first visible in cell ultrastructure, electron microscopy was used to assess the sensitivity of common peatland plants to elevated ozone concentrations. METHODS: Three moss species (Sphagnum angustifolium, S. magellanicum and S. papillosum), a graminoid (Eriophorum vaginatum) and two dwarf shrubs (Vaccinium oxycoccus and Andromeda polifolia), all growing within an intact canopy on peat monoliths, were exposed to a concentration of 0, 50, 100 or 150 ppb ozone in two separate growth chamber experiments simulating either summer or autumn conditions in central Finland. After a 4- or 5-week-long exposure, samples were photographed in a transmission electron microscope and analysed quantitatively using image processing software. KEY RESULTS: In the chlorophyllose cells of the Sphagnum moss leaves from the capitulum, ozone exposure led to a decrease in chloroplast area and in granum stack thickness and various changes in plastoglobuli and cell wall thickness, depending on the species and the experiment. In E. vaginatum, ozone exposure significantly reduced chloroplast cross-sectional areas and the amount of starch, whereas there were no clear changes in the plastoglobuli. In the dwarf shrubs, ozone induced thickening of the cell wall and an increase in the size of plastoglobuli under summer conditions. In contrast, under autumn conditions the cell wall thickness remained unchanged but ozone exposure led to a transient increase in the chloroplast and starch areas, and in the number and size of plastoglobuli. CONCLUSIONS: Ozone responses in the Sphagnum mosses were comparable to typical ozone stress symptoms of higher plants, and indicated sensitivity especially in S. angustifolium. The responses in the dwarf shrubs suggest stimulation of photosynthesis by low ozone concentrations and ozone sensitivity only under cool autumn conditions.

Exposure to Asulox inhibits the growth of mosses.

Asulox is a herbicide used to control bracken. Its effects on mosses were investigated to ascertain whether exposure proved as detrimental as found in parallel studies on pteridophytes. Mature gametophytes of 18 mosses were exposed to a range of concentrations of Asulox under standard conditions and the effects on growth monitored. Plants were cut to a standard length, exposed to Asulox solution for 24 h, grown for 3 weeks and total elongation (main stem and branches) measured. EC50 values were calculated and species ranked according to sensitivity. The effects of exposure on total elongation were compared with those on main stem elongation alone. Under the conditions tested, the total elongation of all species was inhibited after exposure to Asulox. The amount of elongation observed after exposure was different for different species and inhibition of elongation occurred at different exposure concentrations. A single regression equation was not adequate to describe the dose response curves of all species tested. An ability to produce secondary branches may confer increased tolerance to Asulox exposure. It is concluded that mosses suffer detrimental effects after exposure to Asulox at concentrations similar to those that affect fern gametophytes such as bracken.