Oxidation, efflux, and isotopic fractionation of methane during autumnal turnover in a polyhumic, boreal lake

[1] We studied the oxidation and efflux of methane (CH4) in a small, polyhumic lake, Mekkojarvi (southern Finland), during 6 weeks in autumn when the stability of the water mass first weakened, temporarily restabilized, and finally mixed completely. During the summer stratification period, CH4 had accumulated in the anoxic hypolimnion to high concentrations (>150 mmol m−3). Gradual mixing of the water column during the autumn allowed access to both oxygen and CH4 by aerobic methane-oxidizing bacteria (MOB) deeper in the water column. Thus the bulk (∼83–88%) of the CH4 accumulated in the hypolimnion was subsequently consumed by MOB while only 12–17% was lost from the lake to the atmosphere a…

Recycling and fluxes of carbon gases in a stratified boreal lake following experimental carbon addition

Partly anoxic stratified humic lakes are important sources of methane (CH4) and carbon dioxide (CO2) to the atmosphere. We followed the fate of CH4 and CO2 in a small boreal stratified lake, Alinen Mustajärvi, during 2007–2009. In 2008 and 2009 the lake received additions of dissolved organic carbon (DOC) with stable carbon isotope ratio (δ13C) around 16‰ higher than that of local allochthonous DOC. Carbon transformations in the water column were studied by measurements of δ13C of CH4 and of the dissolved inorganic carbon (DIC). Furthermore, CH4 and CO2 production, consumption and emissions were estimated. Methane oxidation was estimated by a diffusion gradient method. The amount, lo…

Stable carbon isotopic composition of peat columns, subsoil and vegetation on natural and forestry-drained boreal peatlands

We studied natural and forestry-drained peatlands to examine the effect of over 34 years lowered water table on the δ13C values of vegetation, bulk peat and subsoil. In the seven studied sites, δ13C in the basal peat layer was 1.1 and 1.2 ‰ lower than that of the middle-layer and surface layer, respectively. Furthermore, there was a positive correlation between the δ13C values of the basal and surface peat layers, possibly due to carbon (C) recycling within the peat column. In the same mire complex, natural fen peat δ13C values were lower than those of the nearby bog, possibly due to the dominance of vascular plants on fen and the generally larger share of recycled C in the fens than in the…

Genetic and Environmental Controls on Nitrous Oxide Accumulation in Lakes

We studied potential links between environmental factors, nitrous oxide (N2O) accumulation, and genetic indicators of nitrite and N2O reducing bacteria in 12 boreal lakes. Denitrifying bacteria were investigated by quantifying genes encoding nitrite and N2O reductases (nirS/nirK and nosZ, respectively, including the two phylogenetically distinct clades nosZ(I) and nosZ(II)) in lake sediments. Summertime N2O accumulation and hypolimnetic nitrate concentrations were positively correlated both at the inter-lake scale and within a depth transect of an individual lake (Lake Vanajavesi). The variability in the individual nirS, nirK, nosZ(I), and nosZ(II) gene abundances was high (up to tenfold) a…

The role ofSphagnummosses 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 ba…

Novel Autotrophic Organisms Contribute Significantly to the Internal Carbon Cycling Potential of a Boreal Lake

ABSTRACT Oxygen-stratified lakes are typical for the boreal zone and also a major source of greenhouse gas emissions in the region. Due to shallow light penetration, restricting the growth of phototrophic organisms, and large allochthonous organic carbon inputs from the catchment area, the lake metabolism is expected to be dominated by heterotrophic organisms. In this study, we test this assumption and show that the potential for autotrophic carbon fixation and internal carbon cycling is high throughout the water column. Further, we show that during the summer stratification carbon fixation can exceed respiration in a boreal lake even below the euphotic zone. Metagenome-assembled genomes an…

Gammaproteobacterial methanotrophs dominate methanotrophy in aerobic and anaerobic layers of boreal lake waters

Small oxygen-stratified humic lakes of the boreal zone are important sources of methane to the atmosphere. Although stable isotope profiling has indicated that a substantial part of methane is already oxidized in the anaerobic water layers in these lakes, the contributions of aerobic and anaerobic methanotrophs in the process are unknown. We used next-generation sequencing of mcrA and 16S rRNA genes to characterize the microbial communities in the water columns of 2 boreal lakes in Finland, Lake Alinen-Mustajärvi and Lake Mekkojärvi, and complemented this with a shotgun metagenomic analysis from Alinen-Mustajärvi and an analysis of pmoA genes and 16S rRNA, mcrA, and pmoA transcripts from Me…

Abiotic and biotic controls on methane formation down to 2.5 km depth within the Precambrian Fennoscandian Shield

Abstract Despite a geological history characterised by high temperature and pressure processes and organic carbon deprived crystalline bedrock, large amounts of hydrocarbons are found in deep groundwaters within Precambrian continental shields. In many sites, methane comprises more that 80% of the dissolved gas phase reaching concentrations of tens of mmol l −1 . In this study, we used isotopic methods to study the carbon isotope systematics and sources of crustal methane within the Fennoscandian Shield. The main study sites were the Outokumpu Deep Drill Hole and the Pyhasalmi mine in Finland, both of which allow groundwater sampling down to 2.5 km depth and have been previously studied for…

Effect of catchment characteristics on aquatic carbon export from a boreal catchment and its importance in regional carbon cycling

Inland waters transport and emit into the atmosphere large amounts of carbon (C), which originates from terrestrial ecosystems. The effect of land cover and land-use practises on C export from terrestrial ecosystems to inland waters is not fully understood, especially in heterogeneous landscapes under human influence. We sampled for dissolved C species in five tributaries with well-determined subcatchments (total size 174.5 km(2)), as well as in various points of two of the subcatchments draining to a boreal lake in southern Finland over a full year. Our aim was to find out how land cover and land-use affect C export from the catchments, as well as CH4 and CO2 concentrations of the streams,…

Effects of alternative electron acceptors on the activity and community structure of methane-producing and consuming microbes in the sediments of two shallow boreal lakes

The role of anaerobic CH4 oxidation in controlling lake sediment CH4 emissions remains unclear. Therefore, we tested how relevant EAs (SO42−, NO3−, Fe3+, Mn4+, O2) affect CH4 production and oxidation in the sediments of two shallow boreal lakes. The changes induced to microbial communities by the addition of Fe3+ and Mn4+ were studied using next-generation sequencing targeting the 16S rRNA and methyl-coenzyme M reductase (mcrA) genes and mcrA transcripts. Putative anaerobic CH4-oxidizing archaea (ANME-2D) and bacteria (NC 10) were scarce (up to 3.4% and 0.5% of archaeal and bacterial 16S rRNA genes, respectively), likely due to the low environmental stability associated with shallow depths.…

Stable carbon isotopic composition of peat columns, subsoil and vegetation on natural and forestry-drained boreal peatlands

We studied natural and forestry-drained peatlands to examine the effect of over 34 years lowered water table on the δ13C values of vegetation, bulk peat and subsoil. In the seven studied sites, δ13C in the basal peat layer was 1.1 and 1.2 ‰ lower than that of the middle-layer and surface layer, respectively. Furthermore, there was a positive correlation between the δ13C values of the basal and surface peat layers, possibly due to carbon (C) recycling within the peat column. In the same mire complex, natural fen peat δ13C values were lower than those of the nearby bog, possibly due to the dominance of vascular plants on fen and the generally larger share of recycled C in the fens than in the…

Vertical stratification of bacteria and archaea in sediments of a small boreal humic lake

ABSTRACT Although sediments of small boreal humic lakes are important carbon stores and greenhouse gas sources, the composition and structuring mechanisms of their microbial communities have remained understudied. We analyzed the vertical profiles of microbial biomass indicators (PLFAs, DNA and RNA) and the bacterial and archaeal community composition (sequencing of 16S rRNA gene amplicons and qPCR of mcrA) in sediment cores collected from a typical small boreal lake. While microbial biomass decreased with sediment depth, viable microbes (RNA and PLFA) were present all through the profiles. The vertical stratification patterns of the bacterial and archaeal communities resembled those in mar…

A synthesis of methane emissions from 71 northern, temperate, and subtropical wetlands

Wetlands are the largest natural source of atmospheric methane. Here, we assess controls on methane flux using a database of approximately 19 000 instantaneous measurements from 71 wetland sites located across subtropical, temperate, and northern high latitude regions. Our analyses confirm general controls on wetland methane emissions from soil temperature, water table, and vegetation, but also show that these relationships are modified depending on wetland type (bog, fen, or swamp), region (subarctic to temperate), and disturbance. Fen methane flux was more sensitive to vegetation and less sensitive to temperature than bog or swamp fluxes. The optimal water table for methane flux was consi…

Whole-Lake Sugar Addition Demonstrates Trophic Transfer of Dissolved Organic Carbon to Top Consumers

Terrestrial dissolved organic carbon (DOC) provides an external carbon source to lake ecosystems. However, there is ongoing debate about whether external DOC that enters a lake can pass up the food web to support top consumers. We show, from experimental manipulation of a whole lake, that externally loaded DOC can contribute appreciably to fish biomass. Monthly additions of cane sugar with a distinct carbon stable isotope value during 2 years rapidly enriched the 13C content of zooplankton and macroinvertebrates, with a more gradual 13C enrichment of fish. After sugar addition stopped, the 13C content of consumers reverted towards original values. A simple isotope mixing model indicated tha…

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 (CO 2 ) and methane (CH 4 ) 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…

Distinct and diverse anaerobic bacterial communities in boreal lakes dominated by candidate division OD1

Lakes have a central role in the carbon cycle of the boreal landscape. These systems typically stratify in summer and their hypolimnetic microbial communities influence burial of biogenic organic matter in sediments. The composition of bacterial communities in these suboxic habitats was studied by pyrosequencing of 16S rRNA amplicons from five lakes with variable dissolved organic carbon (DOC) concentrations. Bacterioplankton communities in the hypolimnetic waters were clearly different from the surface layer with candidate division OD1, Chlorobi and Bacteroidetes as dominant community members. Several operational taxonomic units (OTUs) affiliated with candidate division OD1 were abundant a…

Lowered nutritional quality of prey decrease the growth and biomolecule content of rainbow trout fry

Diet quality is crucial for the development of offspring. Here, we examined how the nutritional quality of prey affects somatic growth and the lipid, carbohydrate, protein, amino acid, and polyunsaturated fatty acid content of rainbow trout (Oncorhynchus mykiss) fry using a three-trophic-level experimental setup. Diets differed especially in their content of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA), which are physiologically essential polyunsaturated fatty acids for a fish fry. Trout were fed with an artificial diet (fish feed, DHA-rich), marine zooplankton diet (krill/Mysis, DHA-rich), or freshwater zooplankton diet (Daphnia, Cladocera, DHA-deficient). The Daphnia were gr…

Stable carbon isotopic composition of peat columns, subsoil and vegetation on natural and forestry-drained boreal peatlands.

We studied natural and forestry-drained peatlands to examine the effect of over 34 years lowered water table on the δ13C values of vegetation, bulk peat and subsoil. In the seven studied sites, δ13C in the basal peat layer was 1.1 and 1.2 ‰ lower than that of the middle-layer and surface layer, respectively. Furthermore, there was a positive correlation between the δ13C values of the basal and surface peat layers, possibly due to carbon (C) recycling within the peat column. In the same mire complex, natural fen peat δ13C values were lower than those of the nearby bog, possibly due to the dominance of vascular plants on fen and the generally larger share of recycled C in the fens than in the…

Vertical stratification patterns of methanotrophs and their genetic controllers in water columns of oxygen-stratified boreal lakes

ABSTRACT The vertical structuring of methanotrophic communities and its genetic controllers remain understudied in the water columns of oxygen-stratified lakes. Therefore, we used 16S rRNA gene sequencing to study the vertical stratification patterns of methanotrophs in two boreal lakes, Lake Kuivajärvi and Lake Lovojärvi. Furthermore, metagenomic analyses were performed to assess the genomic characteristics of methanotrophs in Lovojärvi and the previously studied Lake Alinen Mustajärvi. The methanotroph communities were vertically structured along the oxygen gradient. Alphaproteobacterial methanotrophs preferred oxic water layers, while Methylococcales methanotrophs, consisting of putative…

Priming effect increases with depth in a boreal forest soil

Abstract Climate warming increases labile carbon (C) inputs to soil through increased photosynthesis and C allocation belowground. This could counterintuitively lead to losses of soil C via priming effects (PE): the stimulation of soil organic matter (SOM) decomposition caused by labile C addition. Systematic quantification of PEs in different ecosystems is needed. We measured PEs of free-living soil microbes in different layers of a boreal forest soil, and found that the relative magnitude of the PE increased with soil depth. The relationship between relative PE and the added glucose amount also depended on the soil layer. Our results indicate that the decomposition of SOM in deeper soil l…

Stable isotope profiles of nitrogen gas indicate denitrification in oxygen-stratified humic lakes

Mid-summer N(2) profiles were analyzed from nine oxygen-stratified, humic-acid-rich lakes using a continuous flow isotope ratio mass spectrometer and a Gasbench II device. Sample preparation steps were performed under water to avoid air contamination. The instrument precision for the δ(15)N measurement was high (0.03‰), but for the whole sampling and analysis procedure the mean deviation between replicate samples was 0.13‰ for the δ(15)N measurements and 5.5% for the N(2) gas concentration analysis. The results show that the Gasbench peripheral was suitable for measurement of the (15)N natural abundance of dissolved nitrogen gas, with denitrification indicated by the oversaturation and slig…

Parasite infection alters host stable-isotope composition under controlled feeding

1) Stable isotopes are widely used for studying trophic relationships, but variation driven by environmental conditions or food availability complicates the interpretation of trophic dynamics. Parasites are ubiquitous and known to affect physiological functions of their hosts, but only few studies have assessed the effects of parasites on isotope composition of hosts. 2) We measured the changes in two of the most commonly used stable isotopes in food-web studies, nitrogen (i.e. 15N:14N ratio; denoted as δ15N) and carbon (13C:12C; δ13C) in Daphnia hosts exposed to infection by a parasitic micosporidian in the laboratory. Isotopic signatures of hosts fed a standardised controlled diet were co…

The impact of long-term water level draw-down on microbial biomass : A comparative study from two peatland sites with different nutrient status

We examined the effects of long-term (51 years) drainage on peat microbial communities using phospholipid fatty acid (PLFA) analysis. We analysed the peat profiles of natural and adjacent drained fen and bog sites. Viable microbes (i.e. microbial PLFA) were present in relatively large amounts even in the deepest peat layers of both peatland sites, a finding that warrants further investigation. Microbial biomass was generally higher in the fen than in the bog. Microbial community structure (indexed from PLFA) differed between the fen and bog sites and among depths. Although we did not exclude other factors, the effect of drainage on the total microbial biomass and community structure was not…

The influence of lipid content and taxonomic affiliation on methane and carbon dioxide production from phytoplankton biomass in lake sediment

The greenhouse gases methane (CH4) and carbon dioxide (CO2) are end products of microbial anaerobic degradation of organic matter (OM) in lake sediments. Although previous research has shown that phytoplankton lipid content influences sediment methanogenesis, current understanding on how OM quality affects methanogenesis is still limited. Such information is needed to more accurately assess how lake greenhouse gas emissions may change in response to anthropogenic activities. We cultured 11 phytoplankton species from five classes and studied how taxonomic identity, C : N ratio, lipid content, and fatty acid composition of phytoplankton biomass affects the CH4 and net CO2 production in anaero…

The influence of lipid content and taxonomic affiliation on methane and carbon dioxide production from phytoplankton biomass in lake sediment

Bacterial and phytoplankton responses to nutrient amendments in a boreal lake differ according to season and to taxonomic resolution

Nutrient limitation and resource competition in bacterial and phytoplankton communities may appear different when considering different levels of taxonomic resolution. Nutrient amendment experiments conducted in a boreal lake on three occasions during one open water season revealed complex responses in overall bacterioplankton and phytoplankton abundance and biovolume. In general, bacteria were dominant in spring, while phytoplankton was clearly the predominant group in autumn. Seasonal differences in the community composition of bacteria and phytoplankton were mainly related to changes in observed taxa, while the differences across nutrient treatments within an experiment were due to chang…

Ecosystem responses to increased organic carbon concentration: comparing results based on long-term monitoring and whole-lake experimentation

Recent increases in terrestrial dissolved organic carbon (DOC) concentrations in northern inland waters have many ecological consequences. We examined available data on carbon cycles and food webs of 2 boreal headwater lakes in southern Finland. Basic limnology and catchment characteristics of a pristine lake, Valkea-Kotinen (VK), were monitored over the past 25 years while the lake has undergone browning and DOC increased from similar to 11 to 13 mg L-1. Pronounced changes in the early 2000s represent a regime shift in DOC concentration and color. Lake Alinen Mustajarvi (AM) was manipulated for 2 years by additions of labile DOC (cane sugar), raising the DOC concentration from similar to 1…

Ecosystem responses to increased organic carbon concentration: comparing results based on long-term monitoring and whole-lake experimentation

Recent increases in terrestrial dissolved organic carbon (DOC) concentrations in northern inland waters have many ecological consequences. We examined available data on carbon cycles and food webs of 2 boreal headwater lakes in southern Finland. Basic limnology and catchment characteristics of a pristine lake, Valkea-Kotinen (VK), were monitored over the past 25 years while the lake has undergone browning and DOC increased from ∼11 to 13 mg L−1. Pronounced changes in the early 2000s represent a regime shift in DOC concentration and color. Lake Alinen Mustajärvi (AM) was manipulated for 2 years by additions of labile DOC (cane sugar), raising the DOC concentration from ∼10 to 12 mg L−1, but …

Appendix A. Potential methane oxidation rates detailed by Sphagnum species, habitat, water level, and methane concentration.

Potential methane oxidation rates detailed by Sphagnum species, habitat, water level, and methane concentration.