Ericoid shrub encroachment shifts aboveground-belowground linkages in three peatlands across Europe and Western Siberia.

In northern peatlands, reduction of Sphagnum dominance in favour of vascular vegetation is likely to influence biogeochemical processes. Such vegetation changes occur as the water table lowers and temperatures rise. To test which of these factors has a significant influence on peatland vegetation, we conducted a 3-year manipulative field experiment in Linje mire (northern Poland). We manipulated the peatland water table level (wet, intermediate and dry; on average the depth of the water table was 17.4, 21.2 and 25.3 cm respectively), and we used open-top chambers (OTCs) to create warmer conditions (on average increase of 1.2°C in OTC plots compared to control plots). Peat drying through water table lowering at this local scale had a larger effect than OTC warming treatment per see on Sphagnum mosses and vascular plants. In particular, ericoid shrubs increased with a lower water table level, while Sphagnum decreased. Microclimatic measurements at the plot scale indicated that both water-level and temperature, represented by heating degree days (HDDs), can have significant effects on the vegetation. In a large-scale complementary vegetation gradient survey replicated in three peatlands positioned along a transitional oceanic-continental and temperate-boreal (subarctic) gradient (France-Poland-Western Siberia), an increase in ericoid shrubs was marked by an increase in phenols in peat pore water, resulting from higher phenol concentrations in vascular plant biomass. Our results suggest a shift in functioning from a mineral-N-driven to a fungi-mediated organic-N nutrient acquisition with shrub encroachment. Both ericoid shrub encroachment and higher mean annual temperature in the three sites triggered greater vascular plant biomass and consequently the dominance of decomposers (especially fungi), which led to a feeding community dominated by nematodes. This contributed to lower enzymatic multifunctionality. Our findings illustrate mechanisms by which plants influence ecosystem responses to climate change, through their effect on microbial trophic interactions.

Responses of the species complex Fallopia × bohemica to single-metal contaminations to Cd, Cr or Zn: growth traits, metal accumulation and secondary metabolism.

Plant responses to heavy metals and their storage constitute a crucial step to understand the environmental impacts of metallic trace elements (MTEs). In controlled experiments, we previously demonstrated the tolerance and resilience of Japanese knotweed to soil artificial polymetallic contamination. Using the same experimental design, we tested here the effect of three individual MTEs on Fallopia × bohemica performance traits. Rhizome fragments from three different sites (considered as distinct morphotypes) were grown in a greenhouse for 1 month on a prairial soil artificially contaminated with either Cd, Cr (VI) or Zn at concentrations corresponding to relatively highly polluted soils. Our results confirmed the high tolerance of Bohemian knotweed to metal stress, though, plant response to MTE pollution was dependant on MTE identity. Bohemian knotweed was stimulated by Cr (VI) (increased root and aerial masses), did not display any measurable change in performance traits under Cd at the high dose of 10 mg kg-1, and uptook all MTEs in its rhizome, but only Zn was transferred to its aerial parts. We also highlighted changes in root secondary metabolism that were more accentuated with Zn, including the increase of anthraquinone, stilbene and biphenyl derivatives. These results compared to multi-contamination experiments previously published suggest complex interactions between metals and plant, depending principally on metal identity and also suggest a potential role of soil microbes in the interactions.

Sphagnum Species Modulate their Phenolic Profiles and Mycorrhizal Colonization of Surrounding Andromeda polifolia along Peatland Microhabitats.

Sphagnum mosses mediate long-term carbon accumulation in peatlands. Given their functional role as keystone species, it is important to consider their responses to ecological gradients and environmental changes through the production of phenolics. We compared the extent to which Sphagnum phenolic production was dependent on species, microhabitats and season, and how surrounding dwarf shrubs responded to Sphagnum phenolics. We evaluated the phenolic profiles of aqueous extracts of Sphagnum fallax and Sphagnum magellanicum over a 6-month period in two microhabitats (wet lawns versus dry hummocks) in a French peatland. Phenolic profiles of water-soluble extracts were measured by UHPLC-QTOF-MS. Andromeda polifolia mycorrhizal colonization was quantified by assessing the intensity of global root cortex colonization. Phenolic profiles of both Sphagnum mosses were species-, season- and microhabitat- dependant. Sphagnum-derived acids were the phenolics mostly recovered; relative quantities were 2.5-fold higher in S. fallax than in S. magellanicum. Microtopography and vascular plant cover strongly influenced phenolic profiles, especially for minor metabolites present in low abundance. Higher mycorrhizal colonization of A. polifolia was found in lawns as compared to hummocks. Mycorrhizal abundance, in contrast to environmental parameters, was correlated with production of minor phenolics in S. fallax. Our results highlight the close interaction between mycorrhizae such as those colonizing A. polifolia and the release of Sphagnum phenolic metabolites and suggest that Sphagnum-derived acids and minor phenolics play different roles in this interaction. This work provides new insight into the ecological role of Sphagnum phenolics by proposing a strong association with mycorrhizal colonization of shrubs.

To what extent do food preferences explain the trophic position of heterotrophic and mixotrophic microbial consumers in a Sphagnum peatland?

Although microorganisms are the primary drivers of biogeochemical cycles, the structure and functioning of microbial food webs are poorly studied. This is the case in Sphagnum peatlands, where microbial communities play a key role in the global carbon cycle. Here, we explored the structure of the microbial food web from a Sphagnum peatland by analyzing (1) the density and biomass of different microbial functional groups, (2) the natural stable isotope (δ(13)C and δ(15)N) signatures of key microbial consumers (testate amoebae), and (3) the digestive vacuole contents of Hyalosphenia papilio, the dominant testate amoeba species in our system. Our results showed that the feeding type of testate amoeba species (bacterivory, algivory, or both) translates into their trophic position as assessed by isotopic signatures. Our study further demonstrates, for H. papilio, the energetic benefits of mixotrophy when the density of its preferential prey is low. Overall, our results show that testate amoebae occupy different trophic levels within the microbial food web, depending on their feeding behavior, the density of their food resources, and their metabolism (i.e., mixotrophy vs. heterotrophy). Combined analyses of predation, community structure, and stable isotopes now allow the structure of microbial food webs to be more completely described, which should lead to improved models of microbial community function.

Exposure chambers for studying the partitioning of atmospheric PAHs in environmental compartments: validation and calibration using experimental and computational approaches.

The environmental partitioning of atmospheric polycyclic aromatic hydrocarbons (PAHs) conditions their entry into food chains and subsequent risks for human health. The need for new experimental exposure devices for elucidating the mechanisms governing ecosystemic PAH transfer motivated the elaboration of an original small-scale exposure chamber (EC). A dual approach pairing experimentation and computational fluid dynamics (CFD) was selected to provide comprehensive validation of this EC as a tool to study the transfer and biological effects of atmospheric PAH pollution in microsystems. Soil samples and passive air samplers (PASs) were exposed to atmospheric pollution by phenanthrene (PHE), a gaseous PAH, for 2 weeks in examples of the EC being tested, set up under different conditions. Dynamic concentrations of atmospheric PHE and its uptake by PASs were simulated with CFD, results showing homogeneous distribution and constant atmospheric PHE concentrations inside the ECs. This work provides insight into the setting of given concentrations and pollution levels when using such ECs. The combination of experimentation and CFD is a successful ECs calibration method that should be developed with other semivolatile organic pollutants, including those that tend to partition in the aerosol phase.

Using testate amoeba as potential biointegrators of atmospheric deposition of phenanthrene (polycyclic aromatic hydrocarbon) on "moss/soil interface-testate amoeba community" microecosystems.

Microecosystem models could allow understanding of the impacts of pollutants such as polycyclic aromatic hydrocarbons on ecosystem functioning. We studied the effects of atmospheric phenanthrene (PHE) deposition on the microecosystem "moss/soil interface-testate amoebae (TA) community" over a 1-month period under controlled conditions. We found that PHE had an impact on the microecosystem. PHE was accumulated by the moss/soil interface and was significantly negatively correlated (0.4 < r(2) < 0.7) with total TA abundance and the abundance of five species of TA (Arcella sp., Centropyxis sp., Nebela lageniformis, Nebela tincta and Phryganella sp.). Among sensitive species, species with a superior trophic level (determined by the test aperture size) were more sensitive than other TA species. This result suggests that links between microbial groups in the microecosystems are disrupted by PHE and that this pollutant had effects both direct (ingestion of the pollutant or direct contact with cell) and/or indirect (decrease of prey) on the TA community. The TA community seems to offer a potential integrative tool to understand mechanisms and processes by which the atmospheric PHE deposition affects the links between microbial communities.

Floristic survey, trace element transfers between soil and vegetation and human health risk at an urban industrial wasteland.

This study aimed to determine the trace element accumulation in the soil and plants in an industrial wasteland and to estimate the extent of transfer to humans to measure the effects on and risks to vegetation and human health and find bioindicator plants representative of the levels of the main contaminants. In areas with the highest extractable trace element levels, we observed decreases in plant biodiversity explained by the disappearance of several families, favouring the coverage of tolerant species, such as Urtica dioica and Hedera helix. Trace elements were also found in the leaves of several plants, especially in a dominant species that is poorly studied, Alliaria petiolata. Indeed, this species had the highest contents of Zn (1750 mg.kg-1 DW), Ni (13.1 mg.kg-1 DW), and Cd (18 mg.kg-1 DW) found at the site and is a potential Zn bioindicator since its leaf contents were also representative of the Zn extractable contents in soil (R² = 0.94). The hazard quotient and carcinogen risk revealed that most of the site had an identified or possible risk, mainly due to Pb and As. Native species, especially A. petiolata, could be used in phytoextraction to manage and limit these human and environmental risks.

Fine-scale horizontal and vertical micro-distribution patterns of testate amoebae along a narrow Fen/Bog gradient.

The ecology of peatland testate amoebae is well studied along broad gradient from very wet (pool) to dry (hummock) micro-sites where testate amoebae are often found to respond primarily to the depth to water table (DWT). Much less is known on their responses to finer-scale gradients, and nothing is known of their possible response to phenolic compounds, which play a key role in carbon storage in peatlands. We studied the vertical (0-3, 3-6, and 6-9 cm sampling depths) micro-distribution patterns of testate amoebae in the same microhabitat (Sphagnum fallax lawn) along a narrow ecological gradient between a poor fen with an almost flat and homogeneous Sphagnum carpet (fen) and a "young bog" (bog) with more marked micro-topography and mosaic of poor fen and bog vegetation. We analyzed the relationships between the testate amoeba data and three sets of variables (1) "chemical" (pH, Eh potential, and conductivity), (2) "physical" (water temperature, altitude, i.e., Sphagnum mat micro-topography, and DWT), and (3) phenolic compounds in/from Sphagnum (water-soluble and primarily bound phenolics) as well as the habitat (fen/bog) and the sampling depth. Testate amoeba Shannon H' diversity, equitability J of communities, and total density peaked in lower parts of Sphagnum, but the patterns differed between the fen and bog micro-sites. Redundancy analyses revealed that testate amoeba communities differed significantly in relation to Eh, conductivity, water temperature, altitude, water-soluble phenolics, habitat, and sampling depth, but not to DWT, pH, or primarily bound phenolics. The sensitivity of testate amoebae to weak environmental gradients makes them particularly good integrators of micro-environmental variations and has implications for their use in paleoecology and environmental monitoring. The correlation between testate amoeba communities and the concentration of water-soluble phenolic suggests direct (e.g., physiological) and/or indirect (e.g., through impact on prey organisms) effects on testate amoebae, which requires further research.

Effect of a temperature gradient on Sphagnum fallax and its associated living microbial communities: a study under controlled conditions.

Microbial communities living in Sphagnum are known to constitute early indicators of ecosystem disturbances, but little is known about their response (including their trophic relationships) to climate change. A microcosm experiment was designed to test the effects of a temperature gradient (15, 20, and 25°C) on microbial communities including different trophic groups (primary producers, decomposers, and unicellular predators) in Sphagnum segments (0-3 cm and 3-6 cm of the capitulum). Relationships between microbial communities and abiotic factors (pH, conductivity, temperature, and polyphenols) were also studied. The density and the biomass of testate amoebae in Sphagnum upper segments increased and their community structure changed in heated treatments. The biomass of testate amoebae was linked to the biomass of bacteria and to the total biomass of other groups added and, thus, suggests that indirect effects on the food web structure occurred. Redundancy analysis revealed that microbial assemblages differed strongly in Sphagnum upper segments along a temperature gradient in relation to abiotic factors. The sensitivity of these assemblages made them interesting indicators of climate change. Phenolic compounds represented an important explicative factor in microbial assemblages and outlined the potential direct and (or) indirect effects of phenolics on microbial communities.

Statistical hydrology for evaluating peatland water table sensitivity to simple environmental variables and climate changes application to the mid-latitude/altitude Frasne peatland (Jura Mountains, France).

Peatlands are habitats for a range of fragile flora and fauna species. Their eco-physicochemical characteristics make them as outstanding global carbon and water storage systems. These ecosystems occupy 3% of the worldwide emerged land surface but represent 30% of the global organic soil carbon and 10% of the global fresh water volumes. In such systems, carbon speciation depends to a large extent on specific redox conditions which are mainly governed by the depth of the water table. Hence, understanding their hydrological variability, that conditions both their ecological and biogeochemical functions, is crucial for their management, especially when anticipating their future evolution under climate change. This study illustrates how long-term monitoring of basic hydro-meteorological parameters combined with statistical modeling can be used as a tool to evaluate i) the horizontal (type of peat), ii) vertical (acrotelm/catotelm continuum) and iii) future hydrological variability. Using cross-correlations between meteorological data (precipitation, potential evapotranspiration) and water table depth (WTD), we primarily highlight the spatial heterogeneity of hydrological reactivity across the Sphagnum-dominated Frasne peatland (French Jura Mountain). Then, a multiple linear regression model allows performing hydrological projections until 2100, according to regionalized IPCC RCP4.5 and 8.5 scenarios. Although WTD remains stable during the first half of 21th century, seasonal trends beyond 2050 show lower WTD in winter and markedly greater WTD in summer. In particular, after 2050, more frequent droughts in summer and autumn should occur, increasing WTD. These projections are completed with risk evaluations for peatland droughts until 2100 that appear to be increasing especially for transition seasons, i.e. May-June and September-October. Comparing these trends with previous evaluations of phenol concentrations in water throughout the vegetative period, considered as a proxy of plant functioning intensity, highlights that these hydrological modifications during transitional seasons could be a great ecological perturbation, especially by affecting Sphagnum metabolism.

Modelling and mapping trace element accumulation in Sphagnum peatlands at the European scale using a geomatic model of pollutant emissions dispersion.

Trace elements (TEs) transported by atmospheric fluxes can negatively impact isolated ecosystems. Modelling based on moss-borne TE accumulation makes tracking TE deposition in remote areas without monitoring stations possible. Using a single moss species from ombrotrophic hummock peatlands reinforces estimate quality. This study used a validated geomatic model of particulate matter dispersion to identify the origin of Cd, Zn, Pb and Cu accumulated in Sphagnum capillifolium and the distance transported from their emission sources. The residential and industrial sectors of particulate matter emissions showed the highest correlations with the TEs accumulated in S. capillifolium (0.28(Zn)-0.56(Cu)) and (0.27(Zn)-0.47(Cu), respectively). Distances of dispersion varied depending on the sector of emissions and the considered TE. The greatest transportation distances for mean emissions values were found in the industrial (10.6 km when correlating with all TEs) and roads sectors (13 km when correlating with Pb). The residential sector showed the shortest distances (3.6 km when correlating with Cu, Cd, and Zn). The model presented here is a new tool for evaluating the efficacy of air pollution abatement policies in non-monitored areas and provides high-resolution (200 × 200 m) maps of TE accumulation that make it possible to survey the potential impacts of TEs on isolated ecosystems.

Above- and belowground linkages in Sphagnum peatland: climate warming affects plant-microbial interactions.

Peatlands contain approximately one third of all soil organic carbon (SOC). Warming can alter above- and belowground linkages that regulate soil organic carbon dynamics and C-balance in peatlands. Here we examine the multiyear impact of in situ experimental warming on the microbial food web, vegetation, and their feedbacks with soil chemistry. We provide evidence of both positive and negative impacts of warming on specific microbial functional groups, leading to destabilization of the microbial food web. We observed a strong reduction (70%) in the biomass of top-predators (testate amoebae) in warmed plots. Such a loss caused a shortening of microbial food chains, which in turn stimulated microbial activity, leading to slight increases in levels of nutrients and labile C in water. We further show that warming altered the regulatory role of Sphagnum-polyphenols on microbial community structure with a potential inhibition of top predators. In addition, warming caused a decrease in Sphagnum cover and an increase in vascular plant cover. Using structural equation modelling, we show that changes in the microbial food web affected the relationships between plants, soil water chemistry, and microbial communities. These results suggest that warming will destabilize C and nutrient recycling of peatlands via changes in above- and belowground linkages, and therefore, the microbial food web associated with mosses will feedback positively to global warming by destabilizing the carbon cycle. This study confirms that microbial food webs thus constitute a key element in the functioning of peatland ecosystems. Their study can help understand how mosses, as ecosystem engineers, tightly regulate biogeochemical cycling and climate feedback in peatlands.

Characterizing the feeding habits of the testate amoebae Hyalosphenia papilio and Nebela tincta along a narrow "fen-bog" gradient using digestive vacuole content and 13C and 15N isotopic analyses.

Population dynamics and feeding habits of the testate amoebae Nebela tincta and Hyalosphenia papilio were studied along a short "fen" to "bog" gradient in a Sphagnum-dominated mire (Jura, France). Samples were collected in living "top segments" (0-3 cm) and early declining "bottom segments" (3-6 cm) of Sphagnum fallax peat. Observations of digestive vacuole content and stable isotope analyses ((13)C and (15)N) were used to establish the feeding behavior of both testate amoeba species. Owing to their vertical distribution, the feeding habit of H. papilio was described from top segments, and that of N. tincta from bottom segments. Among identified food sources, those most frequently ingested by N. tincta were spores and mycelia of fungi (55%), microalgae (25%) and cyanobacteria (8.5%). For H. papilio, the most frequently ingested prey were ciliates (55%) and microalgae (35%). Nonmetric Multidimensional Scaling analysis clearly demonstrated that the two species did not have the same feeding habit along the "fen-bog" gradient, and furthermore that a significant spatial split exists in the feeding behavior of H. papilio. Additionally, isotope analyses suggested that H. papilio and N. tincta did not have the same trophic position in the microbial food web, probably resulting from their different feeding strategies.

Atmospheric phenanthrene pollution modulates carbon allocation in red clover (Trifolium pratense L.).

The influence of atmospheric phenanthrene (PHE) exposure (160 µg m(-3)) during one month on carbon allocation in clover was investigated by integrative (plant growth analysis) and instantaneous (13)CO(2) pulse-labelling approaches. PHE exposure diminished plant growth parameters (relative growth rate and net assimilation rate) and disturbed photosynthesis (carbon assimilation rate and chlorophyll content), leading to a 25% decrease in clover biomass. The root-shoot ratio was significantly enhanced (from 0.32 to 0.44). Photosynthates were identically allocated to leaves while less allocated to stems and roots. PHE exposure had a significant overall effect on the (13)C partitioning among clover organs as more carbon was retained in leaves at the expense of roots and stems. The findings indicate that PHE decreases root exudation or transfer to symbionts and in leaves, retains carbon in a non-structural form diverting photosynthates away from growth and respiration (emergence of an additional C loss process).