Ericoid mycorrhizal fungi mediate the response of ombrotrophic peatlands to fertilization: a modeling study.

Ericaceous shrubs adapt to the nutrient-poor conditions in ombrotrophic peatlands by forming symbiotic associations with ericoid mycorrhizal (ERM) fungi. Increased nutrient availability may diminish the role of ERM pathways in shrub nutrient uptake, consequently altering the biogeochemical cycling within bogs. To explore the significance of ERM fungi in ombrotrophic peatlands, we developed the model MWMmic (a peat cohort-based biogeochemical model) into MWMmic-NP by explicitly incorporating plant-soil nitrogen (N) and phosphorus (P) cycling and ERM fungi processes. The new model was applied to simulate the biogeochemical cycles in the Mer Bleue (MB) bog in Ontario, Canada, and their responses to fertilization. MWMmic_NP reproduced the carbon(C)-N-P cycles and vegetation dynamics observed in the MB bog, and their responses to fertilization. Our simulations showed that fertilization increased shrub biomass by reducing the C allocation to ERM fungi, subsequently suppressing the growth of underlying Sphagnum mosses, and decreasing the peatland C sequestration. Our species removal simulation further demonstrated that ERM fungi were key to maintaining the shrub-moss coexistence and C sink function of bogs. Our results suggest that ERM fungi play a significant role in the biogeochemical cycles in ombrotrophic peatlands and should be considered in future modeling efforts.

Integrating McGill Wetland Model (MWM) with peat cohort tracking and microbial controls.

Peatlands store a large amount of organic carbon and are vulnerable to climate change and human disturbances. However, ecosystem-scale peatland models often do not explicitly simulate the decrease in peat substrate quality, i.e., decomposability or the dynamics of decomposers during peat decomposition, which are key controls in determining peat carbon's response to a changing environment. In this paper, we incorporated the tracking of each year's litter input (a cohort) and controls of microbial processes into the McGill Wetland Model (MWMmic) to address this discrepancy. Three major modifications were made: (1) the simple acrotelm-catotelm decomposition model in MWM was changed into a time-aggregated cohort model, to track the decrease in peat quality with decomposition age; (2) microbial dynamics: growth, respiration and death were incorporated into the model and decomposition rates are regulated by microbial biomass; and (3) vertical and horizontal transport of the dissolved organic carbon (DOC) were added and used to regulate the growth of microbial biomass. MWMmic was evaluated against measurements from the Mer Bleue peatland, a raised ombrotrophic bog located in southern Ontario, Canada. The model was able to replicate microbial and DOC dynamics, while at the same time reproduce the ecosystem-level CO2 and DOC fluxes. Sensitivity analysis with MWMmic showed increased peatland resilience to perturbations compared to the original MWM, because of the tracking of peat substrate quality. The analysis revealed the most important parameters in the model to be microbial carbon use efficiency (CUE) and turnover rate. Simulated microbial adaptation with those two physiological parameters less sensitive to disturbances leads to a significantly larger peat C loss in response to warming and water table drawdown. Thus, the rarely explored peatland microbial physiological traits merit further research. This work paves the way for further model development to examine important microbial controls on peatland's biogeochemical cycling.

Beyond the usual suspects: methanogenic communities in eastern North American peatlands are also influenced by nickel and copper concentrations.

Peatlands both accumulate carbon and release methane, but their broad range in environmental conditions means that the diversity of microorganisms responsible for carbon cycling is still uncertain. Here, we describe a community analysis of methanogenic archaea responsible for methane production in 17 peatlands from 36 to 53 N latitude across the eastern half of North America, including three metal-contaminated sites. Methanogenic community structure was analysed through Illumina amplicon sequencing of the mcrA gene. Whether metal-contaminated sites were included or not, metal concentrations in peat were a primary driver of methanogenic community composition, particularly nickel, a trace element required in the F430 cofactor in methyl-coenzyme M reductase that is also toxic at high concentrations. Copper was also a strong predictor, likely due to inhibition at toxic levels and/or to cooccurrence with nickel, since copper enzymes are not known to be present in anaerobic archaea. The methanogenic groups Methanocellales and Methanosarcinales were prevalent in peatlands with low nickel concentrations, while Methanomicrobiales and Methanomassiliicoccales were abundant in peatlands with higher nickel concentrations. Results suggest that peat-associated trace metals are predictors of methanogenic communities in peatlands.

Peatland Microbial Community Composition Is Driven by a Natural Climate Gradient.

Peatlands are important players in climate change-biosphere feedbacks via long-term net carbon (C) accumulation in soil organic matter and as potential net C sources including the potent greenhouse gas methane (CH4). Interactions of climate, site-hydrology, plant community, and groundwater chemical factors influence peatland development and functioning, including C dioxide (CO2) and CH4 fluxes, but the role of microbial community composition is not well understood. To assess microbial functional and taxonomic dissimilarities, we used high throughput sequencing of the small subunit ribosomal DNA (SSU rDNA) to determine bacterial and archaeal community composition in soils from twenty North American peatlands. Targeted DNA metabarcoding showed that although Proteobacteria, Acidobacteria, and Actinobacteria were the dominant phyla on average, intermediate and rich fens hosted greater diversity and taxonomic richness, as well as an array of candidate phyla when compared with acidic and nutrient-poor poor fens and bogs. Moreover, pH was revealed to be the strongest predictor of microbial community structure across sites. Predictive metagenome content (PICRUSt) showed increases in specific genes, such as purine/pyrimidine and amino-acid metabolism in mid-latitude peatlands from 38 to 45° N, suggesting a shift toward utilization of microbial biomass over utilization of initial plant biomass in these microbial communities. Overall, there appears to be noticeable differences in community structure between peatland classes, as well as differences in microbial metabolic activity between latitudes. These findings are in line with a predicted increase in the decomposition and accelerated C turnover, and suggest that peatlands north of 37° latitude may be particularly vulnerable to climate change.

Soil nitrogen determines greenhouse gas emissions from northern peatlands under concurrent warming and vegetation shifting.

Boreal peatlands store an enormous pool of soil carbon that is dependent upon - and vulnerable to changes in - climate, as well as plant community composition. However, how nutrient availability affects the effects of climate and vegetation change on ecosystem processes in these nutrient-poor ecosystems remains unclear. Here we show that although warming promoted higher CH4 emissions, the concurrent addition of N counteracted most (79%) of this effect. The regulation effects of the vegetation functional group, associated with the substrate quality, suggest that CH4 emissions from peatlands under future warming will be less than expected with predicted shrub expansion. In contrast, N2O flux will be enhanced under future warming with predicted shrub expansion. Our study suggests that changes in greenhouse gas emissions in response to future warming and shifts in plant community composition depend on N availability, which reveals the complex interactions that occur when N is not a limiting nutrient.

Multi-year net ecosystem carbon balance of a restored peatland reveals a return to carbon sink.

Peatlands after drainage and extraction are large sources of carbon (C) to the atmosphere. Restoration, through re-wetting and revegetation, aims to return the C sink function by re-establishing conditions similar to that of an undrained peatland. However, the time needed to re-establish C sequestration is not well constrained due to the lack of multi-year measurements. We measured over 3 years the net ecosystem exchange of CO2 (NEE), methane ( F CH 4 ), and dissolved organic carbon (DOC) at a restored post-extraction peatland (RES) in southeast Canada (restored 14 years prior to the start of the study) and compared our observations to the C balance of an intact reference peatland (REF) that has a long-term continuous flux record and is in the same climate zone. Small but significant differences in winter respiration driven by temperature were mainly responsible for differences in cumulative NEE between years. Low growing season inter-annual variability was linked to constancy of the initial spring water table position, controlled by the blocked drainage ditches and the presence of water storage structures (bunds and pools). Half-hour F CH 4 at RES was small except when Typha latifolia-invaded drainage ditches were in the tower footprint; this effect at the ecosystem level was small as ditches represent a minor fraction of RES. The restored peatland was an annual sink for CO2 (-90 ± 18 g C m-2  year-1 ), a source of CH4 (4.4 ± 0.2 g C m-2  year-1 ), and a source of DOC (6.9 ± 2.2 g C m-2  year-1 ), resulting in mean net ecosystem uptake of 78 ± 17 g C m-2  year-1 . Annual NEE at RES was most similar to wetter, more productive years at REF. Integrating structures to increase water retention, alongside re-establishing key species, have been effective at re-establishing the net C sink rate to that of an intact peatland.

Dissolved organic carbon in streams within a subarctic catchment analysed using a GIS/remote sensing approach.

Climate change projections show that temperature and precipitation increases can alter the exchange of greenhouse gases between the atmosphere and high latitude landscapes, including their freshwaters. Dissolved organic carbon (DOC) plays an important role in greenhouse gas emissions, but the impact of catchment productivity on DOC release to subarctic waters remains poorly known, especially at regional scales. We test the hypothesis that increased terrestrial productivity, as indicated by the normalized difference vegetation index (NDVI), generates higher stream DOC concentrations in the Stordalen catchment in subarctic Sweden. Furthermore, we aimed to determine the degree to which other generic catchment properties (elevation, slope) explain DOC concentration, and whether or not land cover variables representing the local vegetation type (e.g., mire, forest) need to be included to obtain adequate predictive models for DOC delivered into rivers. We show that the land cover type, especially the proportion of mire, played a dominant role in the catchment's release of DOC, while NDVI, slope, and elevation were supporting predictor variables. The NDVI as a single predictor showed weak and inconsistent relationships to DOC concentrations in recipient waters, yet NDVI was a significant positive regulator of DOC in multiple regression models that included land cover variables. Our study illustrates that vegetation type exerts primary control in DOC regulation in Stordalen, while productivity (NDVI) is of secondary importance. Thus, predictive multiple linear regression models for DOC can be utilized combining these different types of explanatory variables.

Modelling CO2 emissions from water surface of a boreal hydroelectric reservoir.

To quantify CO2 emissions from water surface of a reservoir that was shaped by flooding the boreal landscape, we developed a daily time-step reservoir biogeochemistry model. We calibrated the model using the measured concentrations of dissolved organic and inorganic carbon (C) in a young boreal hydroelectric reservoir, Eastmain-1 (EM-1), in northern Quebec, Canada. We validated the model against observed CO2 fluxes from an eddy covariance tower in the middle of EM-1. The model predicted the variability of CO2 emissions reasonably well compared to the observations (root mean square error: 0.4-1.3gCm-2day-1, revised Willmott index: 0.16-0.55). In particular, we demonstrated that the annual reservoir surface effluxes were initially high, steeply declined in the first three years, and then steadily decreased to ~115gCm-2yr-1 with increasing reservoir age over the estimated "engineering" reservoir lifetime (i.e., 100years). Sensitivity analyses revealed that increasing air temperature stimulated CO2 emissions by enhancing CO2 production in the water column and sediment, and extending the duration of open water period over which emissions occur. Increasing the amount of terrestrial organic C flooded can enhance benthic CO2 fluxes and CO2 emissions from the reservoir water surface, but the effects were not significant over the simulation period. The model is useful for the understanding of the mechanism of C dynamics in reservoirs and could be used to assist the hydro-power industry and others interested in the role of boreal hydroelectric reservoirs as sources of greenhouse gas emissions.

Modeling surface energy fluxes and thermal dynamics of a seasonally ice-covered hydroelectric reservoir.

The thermal dynamics of human created northern reservoirs (e.g., water temperatures and ice cover dynamics) influence carbon processing and air-water gas exchange. Here, we developed a process-based one-dimensional model (Snow, Ice, WAater, and Sediment: SIWAS) to simulate a full year's surface energy fluxes and thermal dynamics for a moderately large (>500km(2)) boreal hydroelectric reservoir in northern Quebec, Canada. There is a lack of climate and weather data for most of the Canadian boreal so we designed SIWAS with a minimum of inputs and with a daily time step. The modeled surface energy fluxes were consistent with six years of observations from eddy covariance measurements taken in the middle of the reservoir. The simulated water temperature profiles agreed well with observations from over 100 sites across the reservoir. The model successfully captured the observed annual trend of ice cover timing, although the model overestimated the length of ice cover period (15days). Sensitivity analysis revealed that air temperature significantly affects the ice cover duration, water and sediment temperatures, but that dissolved organic carbon concentrations have little effect on the heat fluxes, and water and sediment temperatures. We conclude that the SIWAS model is capable of simulating surface energy fluxes and thermal dynamics for boreal reservoirs in regions where high temporal resolution climate data are not available. SIWAS is suitable for integration into biogeochemical models for simulating a reservoir's carbon cycle.

The uncertain climate footprint of wetlands under human pressure.

Significant climate risks are associated with a positive carbon-temperature feedback in northern latitude carbon-rich ecosystems, making an accurate analysis of human impacts on the net greenhouse gas balance of wetlands a priority. Here, we provide a coherent assessment of the climate footprint of a network of wetland sites based on simultaneous and quasi-continuous ecosystem observations of CO2 and CH4 fluxes. Experimental areas are located both in natural and in managed wetlands and cover a wide range of climatic regions, ecosystem types, and management practices. Based on direct observations we predict that sustained CH4 emissions in natural ecosystems are in the long term (i.e., several centuries) typically offset by CO2 uptake, although with large spatiotemporal variability. Using a space-for-time analogy across ecological and climatic gradients, we represent the chronosequence from natural to managed conditions to quantify the "cost" of CH4 emissions for the benefit of net carbon sequestration. With a sustained pulse-response radiative forcing model, we found a significant increase in atmospheric forcing due to land management, in particular for wetland converted to cropland. Our results quantify the role of human activities on the climate footprint of northern wetlands and call for development of active mitigation strategies for managed wetlands and new guidelines of the Intergovernmental Panel on Climate Change (IPCC) accounting for both sustained CH4 emissions and cumulative CO2 exchange.

Effect of inundation, oxygen and temperature on carbon mineralization in boreal ecosystems.

The inundation of boreal forests and peatlands through the construction of hydroelectric reservoirs can increase carbon dioxide (CO2) and methane (CH4) emission. To establish controls on emission rates, we incubated samples of forest and peat soils, spruce litter, forest litter and peatland litter collected from boreal ecosystems in northern Quebec for 16 weeks and measured CO2 and CH4 production rates under flooded or non-flooded conditions and varying oxygen concentration and temperature. CO2 production under flooded conditions was less than under non-flooded conditions (5-71 vs. 5-85 mg Cg(-1) C), but CH4 production under flooded conditions was larger than under non-flooded conditions (1-8158 vs. 0-86 µg Cg(-1) C). The average CO2 and CH4 production rate factor for flooded:non-flooded conditions was 0.76 and 1.32, respectively. Under flooded conditions, high oxygen concentrations increased CO2 production in peat soils but decreased CH4 production in forest and peat soils and spruce litter. Warmer temperatures (from 4 to 22°C) raised both CO2 production in peat soils and peatland litter, and CH4 production in peat soils and spruce litter. This study shows that the direction and/or strength of CO2 and CH4 fluxes change once boreal forests and peatlands are inundated.

Permafrost conditions in peatlands regulate magnitude, timing, and chemical composition of catchment dissolved organic carbon export.

Permafrost thaw in peatlands has the potential to alter catchment export of dissolved organic carbon (DOC) and thus influence downstream aquatic C cycling. Subarctic peatlands are often mosaics of different peatland types, where permafrost conditions regulate the hydrological setting of each type. We show that hydrological setting is key to observed differences in magnitude, timing, and chemical composition of DOC export between permafrost and nonpermafrost peatland types, and that these differences influence the export of DOC of larger catchments even when peatlands are minor catchment components. In many aspects, DOC export from a studied peatland permafrost plateau was similar to that of a forested upland catchment. Similarities included low annual export (2-3 g C m(-2) ) dominated by the snow melt period (~70%), and how substantial DOC export following storms required wet antecedent conditions. Conversely, nonpermafrost fens had higher DOC export (7 g C m(-2) ), resulting from sustained hydrological connectivity during summer. Chemical composition of catchment DOC export arose from the mixing of highly aromatic DOC from organic soils from permafrost plateau soil water and upland forest surface horizons with nonaromatic DOC from mineral soil groundwater, but was further modulated by fens. Increasing aromaticity from fen inflow to outlet was substantial and depended on both water residence time and water temperature. The role of fens as catchment biogeochemical hotspots was further emphasized by their capacity for sulfate retention. As a result of fen characteristics, a 4% fen cover in a mixed catchment was responsible for 34% higher DOC export, 50% higher DOC concentrations and ~10% higher DOC aromaticity at the catchment outlet during summer compared to a nonpeatland upland catchment. Expansion of fens due to thaw thus has potential to influence landscape C cycling by increasing fen capacity to act as biogeochemical hotspots, amplifying aquatic C cycling, and increasing catchment DOC export.

Greenhouse gas emissions from Canadian peat extraction, 1990-2000: a life-cycle analysis.

This study uses life-cycle analysis to examine the net greenhouse gas (GHG) emissions from the Canadian peat industry for the period 1990-2000. GHG exchange is estimated for land-use change, peat extraction and processing, transport to market, and the in situ decomposition of extracted peat. The estimates, based on an additive GHG accounting model, show that the peat extraction life cycle emitted 0.54 x 10(6) t of GHG in 1990, increasing to 0.89 x 10(6) t in 2000 (expressed as CO2 equivalents using a 100-y time horizon). Peat decomposition associated with end use was the largest source of GHGs, comprising 71% of total emissions during this 11-y period. Land use change resulted in a switch of the peatlands from a GHG sink to a source and contributed an additional 15%. Peat transportation was responsible for 10% of total GHG emissions, and extraction and processing contributed 4%. It would take approximately 2000 y to restore the carbon pool to its original size if peatland restoration is successful and the cutover peatland once again becomes a net carbon sink.