Statistical upscaling of ecosystem CO2 fluxes across the terrestrial tundra and boreal domain: Regional patterns and uncertainties.

The regional variability in tundra and boreal carbon dioxide (CO2 ) fluxes can be high, complicating efforts to quantify sink-source patterns across the entire region. Statistical models are increasingly used to predict (i.e., upscale) CO2 fluxes across large spatial domains, but the reliability of different modeling techniques, each with different specifications and assumptions, has not been assessed in detail. Here, we compile eddy covariance and chamber measurements of annual and growing season CO2 fluxes of gross primary productivity (GPP), ecosystem respiration (ER), and net ecosystem exchange (NEE) during 1990-2015 from 148 terrestrial high-latitude (i.e., tundra and boreal) sites to analyze the spatial patterns and drivers of CO2 fluxes and test the accuracy and uncertainty of different statistical models. CO2 fluxes were upscaled at relatively high spatial resolution (1 km2 ) across the high-latitude region using five commonly used statistical models and their ensemble, that is, the median of all five models, using climatic, vegetation, and soil predictors. We found the performance of machine learning and ensemble predictions to outperform traditional regression methods. We also found the predictive performance of NEE-focused models to be low, relative to models predicting GPP and ER. Our data compilation and ensemble predictions showed that CO2 sink strength was larger in the boreal biome (observed and predicted average annual NEE -46 and -29 g C m-2  yr-1 , respectively) compared to tundra (average annual NEE +10 and -2 g C m-2  yr-1 ). This pattern was associated with large spatial variability, reflecting local heterogeneity in soil organic carbon stocks, climate, and vegetation productivity. The terrestrial ecosystem CO2 budget, estimated using the annual NEE ensemble prediction, suggests the high-latitude region was on average an annual CO2 sink during 1990-2015, although uncertainty remains high.

Lakes as nitrous oxide sources in the boreal landscape.

Estimates of regional and global freshwater N2 O emissions have remained inaccurate due to scarce data and complexity of the multiple processes driving N2 O fluxes the focus predominantly being on summer time measurements from emission hot spots, agricultural streams. Here, we present four-season data of N2 O concentrations in the water columns of randomly selected boreal lakes covering a large variation in latitude, lake type, area, depth, water chemistry, and land use cover. Nitrate was the key driver for N2 O dynamics, explaining as much as 78% of the variation of the seasonal mean N2 O concentrations across all lakes. Nitrate concentrations varied among seasons being highest in winter and lowest in summer. Of the surface water samples, 71% were oversaturated with N2 O relative to the atmosphere. Largest oversaturation was measured in winter and lowest in summer stressing the importance to include full year N2 O measurements in annual emission estimates. Including winter data resulted in fourfold annual N2 O emission estimates compared to summer only measurements. Nutrient-rich calcareous and large humic lakes had the highest annual N2 O emissions. Our emission estimates for Finnish and boreal lakes are 0.6 and 29 Gg N2 O-N/year, respectively. The global warming potential of N2 O from lakes cannot be neglected in the boreal landscape, being 35% of that of diffusive CH4 emission in Finnish lakes.

Increased nitrous oxide emissions from Arctic peatlands after permafrost thaw.

Permafrost in the Arctic is thawing, exposing large carbon and nitrogen stocks for decomposition. Gaseous carbon release from Arctic soils due to permafrost thawing is known to be substantial, but growing evidence suggests that Arctic soils may also be relevant sources of nitrous oxide (N2O). Here we show that N2O emissions from subarctic peatlands increase as the permafrost thaws. In our study, the highest postthaw emissions occurred from bare peat surfaces, a typical landform in permafrost peatlands, where permafrost thaw caused a fivefold increase in emissions (0.56 ± 0.11 vs. 2.81 ± 0.6 mg N2O m-2 d-1). These emission rates match those from tropical forest soils, the world's largest natural terrestrial N2O source. The presence of vegetation, known to limit N2O emissions in tundra, did decrease (by ∼90%) but did not prevent thaw-induced N2O release, whereas waterlogged conditions suppressed the emissions. We show that regions with high probability for N2O emissions cover one-fourth of the Arctic. Our results imply that the Arctic N2O budget will depend strongly on moisture changes, and that a gradual deepening of the active layer will create a strong noncarbon climate change feedback.

Ecosystem carbon response of an Arctic peatland to simulated permafrost thaw.

Permafrost peatlands are biogeochemical hot spots in the Arctic as they store vast amounts of carbon. Permafrost thaw could release part of these long-term immobile carbon stocks as the greenhouse gases (GHGs) carbon dioxide (CO2 ) and methane (CH4 ) to the atmosphere, but how much, at which time-span and as which gaseous carbon species is still highly uncertain. Here we assess the effect of permafrost thaw on GHG dynamics under different moisture and vegetation scenarios in a permafrost peatland. A novel experimental approach using intact plant-soil systems (mesocosms) allowed us to simulate permafrost thaw under near-natural conditions. We monitored GHG flux dynamics via high-resolution flow-through gas measurements, combined with detailed monitoring of soil GHG concentration dynamics, yielding insights into GHG production and consumption potential of individual soil layers. Thawing the upper 10-15 cm of permafrost under dry conditions increased CO2 emissions to the atmosphere (without vegetation: 0.74 ± 0.49 vs. 0.84 ± 0.60 g CO2 -C m-2  day-1 ; with vegetation: 1.20 ± 0.50 vs. 1.32 ± 0.60 g CO2 -C m-2  day-1 , mean ± SD, pre- and post-thaw, respectively). Radiocarbon dating (14 C) of respired CO2 , supported by an independent curve-fitting approach, showed a clear contribution (9%-27%) of old carbon to this enhanced post-thaw CO2 flux. Elevated concentrations of CO2 , CH4 , and dissolved organic carbon at depth indicated not just pulse emissions during the thawing process, but sustained decomposition and GHG production from thawed permafrost. Oxidation of CH4 in the peat column, however, prevented CH4 release to the atmosphere. Importantly, we show here that, under dry conditions, peatlands strengthen the permafrost-carbon feedback by adding to the atmospheric CO2 burden post-thaw. However, as long as the water table remains low, our results reveal a strong CH4 sink capacity in these types of Arctic ecosystems pre- and post-thaw, with the potential to compensate part of the permafrost CO2 losses over longer timescales.

Tundra landscape heterogeneity, not interannual variability, controls the decadal regional carbon balance in the Western Russian Arctic.

Across the Arctic, the net ecosystem carbon (C) balance of tundra ecosystems is highly uncertain due to substantial temporal variability of C fluxes and to landscape heterogeneity. We modeled both carbon dioxide (CO2 ) and methane (CH4 ) fluxes for the dominant land cover types in a ~100-km2 sub-Arctic tundra region in northeast European Russia for the period of 2006-2015 using process-based biogeochemical models. Modeled net annual CO2 fluxes ranged from -300 g C m-2  year-1 [net uptake] in a willow fen to 3 g C m-2  year-1 [net source] in dry lichen tundra. Modeled annual CH4 emissions ranged from -0.2 to 22.3 g C m-2  year-1 at a peat plateau site and a willow fen site, respectively. Interannual variability over the decade was relatively small (20%-25%) in comparison with variability among the land cover types (150%). Using high-resolution land cover classification, the region was a net sink of atmospheric CO2 across most land cover types but a net source of CH4 to the atmosphere due to high emissions from permafrost-free fens. Using a lower resolution for land cover classification resulted in a 20%-65% underestimation of regional CH4 flux relative to high-resolution classification and smaller (10%) overestimation of regional CO2 uptake due to the underestimation of wetland area by 60%. The relative fraction of uplands versus wetlands was key to determining the net regional C balance at this and other Arctic tundra sites because wetlands were hot spots for C cycling in Arctic tundra ecosystems.

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.

Warming of subarctic tundra increases emissions of all three important greenhouse gases - carbon dioxide, methane, and nitrous oxide.

Rapidly rising temperatures in the Arctic might cause a greater release of greenhouse gases (GHGs) to the atmosphere. To study the effect of warming on GHG dynamics, we deployed open-top chambers in a subarctic tundra site in Northeast European Russia. We determined carbon dioxide (CO2 ), methane (CH4 ), and nitrous oxide (N2 O) fluxes as well as the concentration of those gases, inorganic nitrogen (N) and dissolved organic carbon (DOC) along the soil profile. Studied tundra surfaces ranged from mineral to organic soils and from vegetated to unvegetated areas. As a result of air warming, the seasonal GHG budget of the vegetated tundra surfaces shifted from a GHG sink of -300 to -198 g CO2 -eq m-2 to a source of 105 to 144 g CO2 -eq m-2 . At bare peat surfaces, we observed increased release of all three GHGs. While the positive warming response was dominated by CO2 , we provide here the first in situ evidence of increasing N2 O emissions from tundra soils with warming. Warming promoted N2 O release not only from bare peat, previously identified as a strong N2 O source, but also from the abundant, vegetated peat surfaces that do not emit N2 O under present climate. At these surfaces, elevated temperatures had an adverse effect on plant growth, resulting in lower plant N uptake and, consequently, better N availability for soil microbes. Although the warming was limited to the soil surface and did not alter thaw depth, it increased concentrations of DOC, CO2, and CH4 in the soil down to the permafrost table. This can be attributed to downward DOC leaching, fueling microbial activity at depth. Taken together, our results emphasize the tight linkages between plant and soil processes, and different soil layers, which need to be taken into account when predicting the climate change feedback of the Arctic.

The role of Sphagnum mosses in the methane cycling of a boreal mire.

Peatlands are a major natural source of atmospheric methane (CH4). Emissions from Sphagnum-dominated mires are lower than those measured from other mire types. This observation may partly be due to methanotrophic (i.e., methane-consuming) bacteria associated with Sphagnum. Twenty-three of the 41 Sphagnum species in Finland can be found in the peatland at Lakkasuo. To better understand the Sphagnum-methanotroph system, we tested the following hypotheses: (1) all these Sphagnum species support methanotrophic bacteria; (2) water level is the key environmental determinant for differences in methanotrophy across habitats; (3) under dry conditions, Sphagnum species will not host methanotrophic bacteria; and (4) methanotrophs can move from one Sphagnum shoot to another in an aquatic environment. To address hypotheses 1 and 2, we measured the water table and CH4 oxidation for all Sphagnum species at Lakkasuo in 1-5 replicates for each species. Using this systematic approach, we included Sphagnum spp. with narrow and broad ecological tolerances. To estimate the potential contribution of CH4 to moss carbon, we measured the uptake of delta13C supplied as CH4 or as carbon dioxide dissolved in water. To test hypotheses 2-4, we transplanted inactive moss patches to active sites and measured their methanotroph communities before and after transplantation. All 23 Sphagnum species showed methanotrophic activity, confirming hypothesis 1. We found that water level was the key environmental factor regulating methanotrophy in Sphagnum (hypothesis 2). Mosses that previously exhibited no CH4 oxidation became active when transplanted to an environment in which the microbes in the control mosses were actively oxidizing CH4 (hypothesis 4). Newly active transplants possessed a Methylocystis signature also found in the control Sphagnum spp. Inactive transplants also supported a Methylocystis signature in common with active transplants and control mosses, which rejects hypothesis 3. Our results imply a loose symbiosis between Sphagnum spp. and methanotrophic bacteria that accounts for potentially 10-30% of Sphagnum carbon.

Author Correction: Neglecting diurnal variations leads to uncertainties in terrestrial nitrous oxide emissions.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A plant-microbe interaction framework explaining nutrient effects on primary production.

In most terrestrial ecosystems, plant growth is limited by nitrogen and phosphorus. Adding either nutrient to soil usually affects primary production, but their effects can be positive or negative. Here we provide a general stoichiometric framework for interpreting these contrasting effects. First, we identify nitrogen and phosphorus limitations on plants and soil microorganisms using their respective nitrogen to phosphorus critical ratios. Second, we use these ratios to show how soil microorganisms mediate the response of primary production to limiting and non-limiting nutrient addition along a wide gradient of soil nutrient availability. Using a meta-analysis of 51 factorial nitrogen-phosphorus fertilization experiments conducted across multiple ecosystems, we demonstrate that the response of primary production to nitrogen and phosphorus additions is accurately predicted by our stoichiometric framework. The only pattern that could not be predicted by our original framework suggests that nitrogen has not only a structural function in growing organisms, but also a key role in promoting plant and microbial nutrient acquisition. We conclude that this stoichiometric framework offers the most parsimonious way to interpret contrasting and, until now, unresolved responses of primary production to nutrient addition in terrestrial ecosystems.

Estimates of microbial quality and concentration of copper in distributed drinking water are highly dependent on sampling strategy.

The numbers of bacteria generally increase in distributed water. Often household pipelines or water fittings (e.g., taps) represent the most critical location for microbial growth in water distribution systems. According to the European Union drinking water directive, there should not be abnormal changes in the colony counts in water. We used a pilot distribution system to study the effects of water stagnation on drinking water microbial quality, concentration of copper and formation of biofilms with two commonly used pipeline materials in households; copper and plastic (polyethylene). Water stagnation for more than 4h significantly increased both the copper concentration and the number of bacteria in water. Heterotrophic plate counts were six times higher in PE pipes and ten times higher in copper pipes after 16 h of stagnation than after only 40 min stagnation. The increase in the heterotrophic plate counts was linear with time in both copper and plastic pipelines. In the distribution system, bacteria originated mainly from biofilms, because in laboratory tests with water, there was only minor growth of bacteria after 16 h stagnation. Our study indicates that water stagnation in the distribution system clearly affects microbial numbers and the concentration of copper in water, and should be considered when planning the sampling strategy for drinking water quality control in distribution systems.

Denitrification in the river estuaries of the northern Baltic Sea.

Estuaries have been suggested to have an important role in reducing the nitrogen load transported to the sea. We measured denitrification rates in six estuaries of the northern Baltic Sea. Four of them were river mouths in the Bothnian Bay (northern Gulf of Bothnia), and two were estuary bays, one in the Archipelago Sea (southern Gulf of Bothnia) and the other in the Gulf of Finland. Denitrification rates in the four river mouths varied between 330 and 905 micromol N m(-2) d(-1). The estuary bays at the Archipelago Sea and the Gulf of Bothnia had denitrification rates from 90 micromol N m(-2) d(-1) to 910 micromol N m(-2) d(-1) and from 230 micromol N m(-2) d(-1) to 320 micromol N m(-2) d(-1), respectively. Denitrification removed 3.6-9.0% of the total nitrogen loading in the river mouths and in the estuary bay in the Gulf of Finland, where the residence times were short. In the estuary bay with a long residence time, in the Archipelago Sea, up to 4.5% of nitrate loading and 19% of nitrogen loading were removed before entering the sea. According to our results, the sediments of the fast-flowing rivers and the estuary areas with short residence times have a limited capacity to reduce the nitrogen load to the Baltic Sea.

Degradation potentials of dissolved organic carbon (DOC) from thawed permafrost peat.

Global warming can substantially affect the export of dissolved organic carbon (DOC) from peat-permafrost to aquatic systems. The direct degradability of such peat-derived DOC, however, is poorly constrained because previous permafrost thaw studies have mainly addressed mineral soil catchments or DOC pools that have already been processed in surface waters. We incubated peat cores from a palsa mire to compare an active layer and an experimentally thawed permafrost layer with regard to DOC composition and degradation potentials of pore water DOC. Our results show that DOC from the thawed permafrost layer had high initial degradation potentials compared with DOC from the active layer. In fact, the DOC that showed the highest bio- and photo-degradability, respectively, originated in the thawed permafrost layer. Our study sheds new light on the DOC composition of peat-permafrost directly upon thaw and suggests that past estimates of carbon-dioxide emissions from thawed peat permafrost may be biased as they have overlooked the initial mineralization potential of the exported DOC.

Selected ion monitoring in quantitative gas-liquid chromatographic-mass spectrometric detection of fatty acid methyl esters from environmental samples.

To calculate selected ion monitoring (SIM) gas-liquid chromatography (GLC)-mass spectrometry (MS) results of phospholipid fatty acids (PLFAs) from environmental samples, coefficients were calculated for each fatty acid by dividing the sum of ion intensities in SCAN with that of ions followed in SIM. The SIM chromatogram areas were multiplied with the coefficients, and then processed as in SCAN. The results were compared to those obtained using calibration curves and SCAN. The calibration curve and coefficient based results had the greatest errors of 7.8 and 6.7%, respectively, outside standard deviations of SCAN percentages. The PLFA contents calculated using calibration curves and coefficients were 104.9+/-7.3% and 101.5+/-8.6%, respectively, of SCAN values. SIM increased sensitivity approximately 10-fold from SCAN, and the smallest detectable injected amount was approximately 50 ng (0.18 nmol) for 20 fatty acids, corresponding to 4 x 10(6) cells.

The effects of changing water flow velocity on the formation of biofilms and water quality in pilot distribution system consisting of copper or polyethylene pipes.

We studied the effects of flow velocity on the formation of biofilms and the concentration of bacteria in water in copper and plastic (polyethylene, PE) pipes. The formation of biofilms increased with the flow velocity of water. The increase in microbial numbers and contents of ATP was clearer in the PE pipes than in the copper pipes. This was also seen as increased consumption of microbial nutrients in the pipeline system. This indicates that the mass transfer of nutrients is in major role in the growth of biofilms. However, the increased biomass of biofilms did not affect microbial numbers in the water. Rapid changes in water flow rate resuspended biofilms and sediments which increased the concentrations of bacteria and copper in water. The disturbance caused by the changing water flow was also seen as an increase in the particle counts and water turbidity recorded with online instrumentation.

Heterogeneity of carbon loss and its temperature sensitivity in East-European subarctic tundra soils.

Arctic peatlands store large stocks of organic carbon which are vulnerable to the climate change but their fate is uncertain. There is increasing evidence that a part of it will be lost as a result of faster microbial mineralization. We studied the vulnerability of 3500-5900 years old bare peat uplifted from permafrost layers by cryogenic processes to the surface of an arctic peat plateau. We aimed to find biotic and abiotic drivers of CLOSS from old peat and compare them with those of adjacent, young vegetated soils of the peat plateau and mineral tundra. The soils were incubated in laboratory at three temperatures (4°C, 12°C and 20°C) and two oxygen levels (aerobic, anaerobic). CLOSS was monitored and soil parameters (organic carbon quality, nutrient availability, microbial activity, biomass and stoichiometry, and extracellular oxidative and hydrolytic enzyme pools) were determined. We found that CLOSS from the old peat was constrained by low microbial biomass representing only 0.22% of organic carbon. CLOSS was only slightly reduced by the absence of oxygen and exponentially increased with temperature, showing the same temperature sensitivity under both aerobic and anaerobic conditions. We conclude that carbon in the old bare peat is stabilized by a combination of physical, chemical and biological controls including soil compaction, organic carbon quality, low microbial biomass and the absence of plants.

Neglecting diurnal variations leads to uncertainties in terrestrial nitrous oxide emissions.

Nitrous oxide (N2O) is an important greenhouse gas produced in soil and aquatic ecosystems. Its warming potential is 296 times higher than that of CO2. Most N2O emission measurements made so far are limited in temporal and spatial resolution causing uncertainties in the global N2O budget. Recent advances in laser spectroscopic techniques provide an excellent tool for area-integrated, direct and continuous field measurements of N2O fluxes using the eddy covariance method. By employing this technique on an agricultural site with four laser-based analysers, we show here that N2O exchange exhibits contrasting diurnal behaviour depending upon soil nitrogen availability. When soil N was high due to fertilizer application, N2O emissions were higher during daytime than during the night. However, when soil N became limited, emissions were higher during the night than during the day. These reverse diurnal patterns supported by isotopic analyses may indicate a dominant role of plants on microbial processes associated with N2O exchange. This study highlights the potential of new technologies in improving estimates of global N2O sources.

Pipeline materials modify the effectiveness of disinfectants in drinking water distribution systems.

We studied how pipe material can modify the effectiveness of UV- and chlorine disinfection in drinking water and biofilms. This study was done with two pipe materials: copper and composite plastic (polyethylene, PE) in a pilot scale water distribution network. UV-disinfection decreased viable bacterial numbers in the pilot waterworks and outlet water of pipes on average by 79%, but in biofilms its disinfecting effect was minor. Chlorine decreased effectively the microbial numbers in water and biofilms of PE pipes. In outlet water from copper pipes, the effect of chlorination was weaker; microbial numbers increased back to the level before chlorination within a few days. In the biofilms present in the copper pipes, chlorine decreased microbial numbers only in front of the pipeline. One reason for weaker efficiency of chlorine in copper pipes was that its concentration declined more rapidly in the copper pipes than in the PE pipes. These results means that copper pipes may require a higher chlorine dosage than plastic pipes to achieve effective disinfection of the pipes.

Environmental and microbial factors influencing methane and nitrous oxide fluxes in Mediterranean cork oak woodlands: trees make a difference.

Cork oak woodlands (montado) are agroforestry systems distributed all over the Mediterranean basin with a very important social, economic and ecological value. A generalized cork oak decline has been occurring in the last decades jeopardizing its future sustainability. It is unknown how loss of tree cover affects microbial processes that are consuming greenhouse gases in the montado ecosystem. The study was conducted under two different conditions in the natural understory of a cork oak woodland in center Portugal: under tree canopy (UC) and open areas without trees (OA). Fluxes of methane and nitrous oxide were measured with a static chamber technique. In order to quantify methanotrophs and bacteria capable of nitrous oxide consumption, we used quantitative real-time PCR targeting the pmoA and nosZ genes encoding the subunit of particulate methane mono-oxygenase and catalytic subunit of the nitrous oxide reductase, respectively. A significant seasonal effect was found on CH4 and N2O fluxes and pmoA and nosZ gene abundance. Tree cover had no effect on methane fluxes; conversely, whereas the UC plots were net emitters of nitrous oxide, the loss of tree cover resulted in a shift in the emission pattern such that the OA plots were a net sink for nitrous oxide. In a seasonal time scale, the UC had higher gene abundance of Type I methanotrophs. Methane flux correlated negatively with abundance of Type I methanotrophs in the UC plots. Nitrous oxide flux correlated negatively with nosZ gene abundance at the OA plots in contrast to that at the UC plots. In the UC soil, soil organic matter had a positive effect on soil extracellular enzyme activities, which correlated positively with the N2O flux. Our results demonstrated that tree cover affects soil properties, key enzyme activities and abundance of microorganisms and, consequently net CH4 and N2O exchange.

Responses of two Sphagnum moss species and Eriophorum vaginatum to enhanced UV-B in a summer of low UV intensity.

• The flux of ultraviolet (UV)-B radiation to the Earth's surface is increasing, particularly in high latitudes. We studied the sensitivity of some dominant plant species of boreal and subarctic peatlands to this increase. • Intact peat monoliths with the mosses Sphagnum balticum and Sphagnum papillosum, and cotton grass (Eriophorum vaginatum) were exposed to ambient solar UV-B or ambient solar UV-B supplemented by 30% in a field experiment in central Finland. • Although the UV-B dose was low during the growing season, owing to frequent cloudiness, both Sphagnum species showed significantly higher membrane permeability under enhanced UV-B. In S. balticum, UV-B tended to decrease the capitulum dry mass and induced a 30-40% increase in the concentration of chlorophyll and carotenoid pigments. Enhanced UV-B had no effects on leaf morphology, chlorophyll fluorescence or stomatal functioning in E. vaginatum. • The various UV-B responses in the Sphagnum species under investigation indicate that they may be sensitive even to small increases in solar UV-B radiation. By contrast, E. vaginatum appeared to tolerate the UV-B fluxes of the experiment.