Climate change effects on carbon and nitrogen mineralization in peatlands through changes in soil quality

Climate change will directly affect carbon and nitrogen mineralization through changes in temperature and soil moisture, but it may also indirectly affect mineralization rates through changes in soil quality. We used an experimental mesocosm system to examine the effects of 6‐year manipulations of infrared loading (warming) and water‐table level on the potential anaerobic nitrogen and carbon (as carbon dioxide (CO₂) and methane (CH₄) production) mineralization potentials of bog and fen peat over 11 weeks under uniform anaerobic conditions. To investigate the response of the dominant methanogenic pathways, we also analyzed the stable isotope composition of CH₄ produced in the samples. Bog peat from the highest water‐table treatment produced more CO₂ than bog peat from drier mesocosms. Fen peat from the highest water‐table treatment produced the most CH₄. Cumulative nitrogen mineralization was lowest in bog peat from the warmest treatment and lowest in the fen peat from the highest water‐table treatment. As all samples were incubated under constant conditions, observed differences in mineralization patterns reflect changes in soil quality in response to climate treatments. The largest treatment effects on carbon mineralization as CO₂ occurred early in the incubations and were ameliorated over time, suggesting that the climate treatments changed the size and/or quality of a small labile carbon pool. CH₄ from the fen peat appeared to be predominately from the acetoclastic pathway, while in the bog peat a strong CH₄ oxidation signal was present despite the anaerobic conditions of our incubations. There was no evidence that changes in soil quality have lead to differences in the dominant methanogenic pathways in these systems. Overall, our results suggest that even relatively short‐term changes in climate can alter the quality of peat in bogs and fens, which could alter the response of peatland carbon and nitrogen mineralization to future climate change.     

Effects of short-term drying and irrigation on CO2 and CH4 production and emission from mesocosms of a northern bog and an alpine fen

Atmospheric CO₂ and CH₄ exchange in peatlands is controlled by water table levels and soil moisture, but impacts of short periods of dryness and rainfall are poorly known. We conducted drying-rewetting experiments with mesocosms from an ombrotrophic northern bog and an alpine, minerotrophic fen. Efflux of CO₂ and CH₄ was measured using static chambers and turnover and diffusion rates were calculated from depth profiles of gas concentrations. Due to a much lower macroporosity in the fen compared to the bog peat, water table fluctuated more strongly when irrigation was stopped and resumed, about 11 cm in the fen and 5 cm in the bog peat. Small changes in air filled porosity caused CO₂ and CH₄ concentrations in the fen peat to be insensitive to changes in water table position. CO₂ emission was by a factor of 5 higher in the fen than in the bog mesocosms and changed little with water table position in both peats. This was probably caused by the importance of the uppermost, permanently unsaturated zone for auto- and heterotrophic CO₂ production, and a decoupling of air filled porosity from water table position. CH₄ emission was <0.4 mmol m^?2 day^?1 in the bog peat, and up to >12.6 mmol m^?2 day^?1 in the fen peat, where it was lowered by water table fluctuations. CH₄ production was limited to the saturated zone in the bog peat but proceeded in the capillary fringe of the fen peat. Water table drawdown partly led to inhibition of methanogenesis in the newly unsaturated zone, but CH₄ production appeared to continue after irrigation without time-lag. The identified effects of irrigation on soil moisture and respiration highlight the importance of peat physical properties for respiratory dynamics; but the atmospheric carbon exchange was fairly insensitive to the small-scale fluctuations induced.     

Recovery of Methanogenesis Following Summer Drought in Soils from Two Cool Temperate Peatlands,New York State,USA

Following a summer drought, intact cores of peat soil from two cool temperate peatlands (a rain-fed bog and a groundwater-fed swamp) were exposed experimentally to three different water table levels. The goal was to examine recovery of anaerobic methanogenesis and to evaluate peat soil decomposition to methane (CH₄), carbon dioxide (CO₂), and dissolved organic carbon (DOC) upon rewetting. Methane emission from soils to the atmosphere was greatest (mean = 80 μmol m^?2 s^?1) when the entire peat core was rewetted quickly; emission was negligible at low water level and when peat cores were rewetted gradually. Rates of CO₂ emission (mean = 1.0 μmol m^?2 s^?1) were relatively insensitive to water level. Concentrations of CH₄ in soil air spaces suggest that onset of methanogenesis induces, but later represses, aerobic oxidation of CH₄ above the water table. Concentrations of CO₂ suggest production at the soil surface of swamp peat versus at greater depths in bog peat. Portions of peat soil incubated in vitro without oxygen (O₂) exhibited a lag before the onset of methanogenesis, and the lag time was less in peat from the cores rewetted quickly. The inhibition of methanogenesis by the selective inhibitor 2-bromoethanesulfonic acid (BES) decreased CO₂ production by 20 to 30% but resulted in an increase in concentrations of DOC by 2 to 5 times. The results show that methanogens in peat soils tolerate moderate drought, and recovery varies among different peat types. In peat soils, the inhibition of methanogenesis might enhance DOC availability.     

Winter fluxes of methane from Minnesota peatlands

Winter fluxes of methane were investigated in northern Minnesota during 1988–89 and 1989–90. Two bogs and a fen emitted methane throughout the snow-covered season (November through March). Fluxes decreased to a low level of 3–16 mg CH₄ m^–2 d^–1 in late March, reflecting decreasing peat temperatures and (in 1989–90) increasing depth of frost in the peat. Winter fluxes calculated by integration for an open poor fen, an open bog, a forested bog hollow, and a hummock site in the forested bog averaged 49, 12, 13, and 5 mg m^–2 d^–1, respectively, in 1989–1990 (the year most measurements were made). These comprised 11%, 4%, 15%, and 21% of total annual flux.     

Methanogenesis in McLean Bog,an Acidic Peat Bog in Upstate New York: Stimulation by H2/CO2 in the Presence of Rifampicin,or by Low Concentrations of Acetate

Acidic peat bog soils produce CH₄ and although molecular biological studies have demonstrated the presence of diverse methano-genic populations in them, few studies have sustained methanogenesis by adding the CH₄ precursors H₂/CO₂ or acetate, and few indigenous methanogens have been cultured. McLean Bog is a small (ca. 70 m across), acidic (pH 3.4–4.3) Sphagnum -dominated bog in upstate New York. Although addition of H₂/CO₂ or 10 mM acetate stimulated methanogenesis in soils from a nearby circumneutral-pH fen, neither of these substrates led to sustained methanogenesis in McLean Bog soil slurries. After a brief period of stimulation by H₂/CO₂, methanogenesis in McLean Bog soil declined, which could be attributed to buildup of large amounts of acetic acid produced from the H₂/CO₂ by acetogens. Addition of the antibiotic rifampicin inhibited acetogenesis (carried out by Bacteria) and allowed methanogenesis (carried out by Archaea) to continue. Using rifampicin, we were able to study effects of temperature, pH, and salts on methanogenesis from H₂/CO₂ in McLean Bog soil samples. The enriched H₂/CO₂-utilizing methanogens showed an optimum for activity near pH 5, and a temperature optimum near 35°C. Methanogenesis was not stimulated by addition of 10 mM acetate, but it was stimulated by 1 mM acetate, and multiple additions were consumed at increasing rates and nearly stoichiometrically converted to CH₄. In conclusion, we have found that both hydrogentrophic and aceticlastic methanogens are present in McLean Bog soils, and that methanogenic activity can be stimulated using H₂/CO₂ in the presence of rifampicin, or using low concentrations of acetate.     

Winter CO2 CH4 and N2O fluxes on some natural and drained boreal peatlands

CO₂ and CH₄ fluxes during the winter were measured at natural and drained bog and fen sites in eastern Finland using both the closed chamber method and calculations of gas diffusion along a concentration gradient through the snowpack. The snow diffusion results were compared with those obtained by chamber, but the winter flux estimates were derived from chamber data only. CH₄ emissions from a poor bog were lower than those from an oligotrophic fen, while both CO₂ and CH₄ fluxes were higher in theCarex rostrata- occupied marginal (lagg) area of the fen than in the slightly less fertile centre. Average estimated winter CO₂-C losses from virgin and drained forested peatlands were 41 and 68 g CO₂-C m^–2, respectively, accounting for 23 and 21% of the annual total CO₂ release from the peat. The mean release of CH₄-C was 1.0 g in natural bogs and 3.4 g m^–2 in fens, giving rise to winter emissions averaging to 22% of the annual emission from the bogs and 10% of that from the fens. These wintertime carbon gas losses in Finnish natural peatlands were even greater than reported average long-term annual C accumulation values (less than 25g C m^–2). The narrow range of 10–30% of the proportion of winter CO₂ and CH₄ emissions from annual emissions found in Finnish peatlands suggest that a wider generalization in the boreal zone is possible. Drained forested bogs emitted 0.3 g CH₄-C m^–2 on the average, while the effectively drained fens consumed an average of 0.01 g CH₄-C m^–2. Reason for the low CH₄. efflux or net oxidation in drained peatlands probably lies in low substrate supply and thus low CH₄ production in the anoxic deep peat layers. N₂O release from a fertilized grassland site in November–May was 0.7 g N₂O m^–2, accounting for 38% of the total annual emission, while a forested bog released none and two efficiently drained forested fens 0.09 (28% of annual release) and 0.04 g N₂O m^–2 (27%) during the winter, respectively.     

Seasonal variation in CH4 emissions and production and oxidation potentials at microsites on an oligotrophic pine fen

 Temporal and spatial variation in CH₄ emissions was studied at hummock, Eriophorum lawn, flark and Carex lawn microsites in an oligotrophic pine fen over the growing season using a static chamber method, and CH₄ production and oxidation potentials in peat profiles from hummock and flark were determined in laboratory incubation experiments. Emissions were lowest in the hummocks, and decreased with increasing hummock height, while in the lawns and flarks they increased with increasing sedge cover. Statistical response functions with water table and peat temperature as independent variables were calculated in order to reconstruct seasonal CH₄ emissions by reference to the time series for peat temperature and water table specific to each microsite type. Mean CH₄ emissions in the whole area in the snow-free period of 1993, weighted in terms of the proportions of the microsites, were 1.7 mol CH₄ m^–2. Potential CH₄ production and oxidation rates were very low in the hummocks rising above the groundwater table, but were relatively similar when expressed per dry weight of peat both in the hummocks and flarks below the water table. The CH₄ production potential increased in autumn at both microsites and CH₄ oxidation potential seemed to decrease. The decrease in temperature in autumn certainly reduced in situ decomposition processes, possibly leaving unused substrates in the peat, which would explain the increase in CH₄ production potential. Received: 16 August 1996 / Accepted: 30 November 1996     

Evidence for Anaerobic CH 4 Oxidation in Freshwater Peatlands

This study involved in vitro assays of peat soil to investigate the occurrence, importance and potential mechanism(s) of anaerobic methane oxidation (AOM) in several northern peatlands ranging from ombrotrophic bog to minerotrophic fen. Although strong evidence suggests that AOM is linked to sulfate reduction in marine sediments, very little is known about AOM in freshwater systems such as northern peatlands, which have large methane (CH ₄ ) production and are a significant source of atmospheric CH ₄ . Our results showed a mean net AOM rate of 17 ± 2.6 nmol kg ^{? 1} s ^{? 1} with a maximum rate of 176 nmol kg ^{? 1} s ^{? 1} for a minerotrophic fen in central New York. AOM was demonstrated with three independent methods to verify our results: (a) additions of methanogenic inhibitors, (b) stable isotope enrichment ( ¹³ C-CH ₄ ), and (c) natural abundance stable isotope analysis ( ¹³ C-CH ₄ ). These experiments confirmed that AOM occurs simultaneously with methanogenesis, consumes a significant portion of gross CH ₄ production, and significantly fractionates C isotopes (～ ?127‰). Experiments using a variety of potential electron acceptors demonstrated that Fe(III) and SO₄ ^{2 ?} are not quantitatively important, while the role of NO ₃ ^? is uncertain and deserves more attention. The exact mechanism(s) for AOM in peat soils remains unclear; however the AOM rates reported in this study are similar to those reported for CH ₄ production and aerobic CH ₄ oxidation in northern peatlands, suggesting that AOM may be an important control on CH ₄ fluxes in northern peatland ecosystems.     

Change in fluxes of carbon dioxide,methane and nitrous oxide due to forest drainage of mire sites of different trophy

Northern peatlands accumulate atmospheric CO₂ thus counteracting climate warming. However, CH₄ which is more efficient as a greenhouse gas than CO₂, is produced in the anaerobic decomposition processes in peat. When peatlands are taken for forestry their water table is lowered by ditching. We studied long-term effects of lowered water table on the development of vegetation and the annual emissions of CO₂, CH₄ and N₂O in an ombrotrophic bog and in a minerotrophic fen in Finland. Reclamation of the peat sites for forestry had changed the composition and coverage of the field and ground layer species, and increased highly the growth of tree stand at the drained fen. In general, drainage increased the annual CO₂ emissions but the emissions were also affected by the natural fluctuations of water table. In contrast to CO₂, drainage had decreased the emissions of CH₄, the drained fen even consumed atmospheric CH₄. CO₂ and CH₄ emissions were higher in the virgin fen than in the virgin bog. There were no N₂O emissions from neither type of virgin sites. Drainage had, however, highly increased the N₂O emissions from the fen. The results suggest that post-drainage changes in gas fluxes depend on the trophy of the original mires.     

Effect of peat re-wetting on carbon and nutrient fluxes, greenhouse gas production and diversity of methanogenic archaeal community

Many peatlands were affected by drainage in the past, and restoration of their water regime aims to bring back their original functions. The purpose of our study was to simulate re-wetting of soils of different types of drained peatlands (bogs and minerotrophic mires, located in the Sumava Mountains, Czech Republic) under laboratory conditions (incubation for 15 weeks) and to assess possible risks of peatland water regime restoration - especially nutrient leaching and the potentials for CO₂ and CH₄ production. After re-wetting of soils sampled from drained peatlands (simulated by anaerobic incubation) (i) phosphorus concentration (SRP) did not change in any soil, (ii) concentration of ammonium and dissolved organic nitrogen (DON) increased, but only in a drained fen, (iii) DOC increased significantly in the drained fen and degraded drained bog, (iv) CO₂ production decreased, (v) CH₄ production and the number of methanogens increased in all soils, and (vi) archaeal methanogenic community composition was also affected by re-wetting; it differed significantly between drained and pristine fens, whereas it was more similar between drained and pristine bogs. Overall, the soils from fens reacted more dynamically to re-wetting than the bogs, and therefore, some nutrients (especially nitrogen) and DOC leaching may be expected from drained fens after their water regime restoration. However, if compared to their state before restoration, ammonium and phosphorus leaching should not increase and leaching of nitrates and DON should even decrease after restoration, especially during the vegetation season. Further, CO₂ production in soils of fens and bogs should decrease after their water regime restoration, whereas CH₄ production in soils should increase. However, we cannot derive any clear conclusions about CH₄ emissions from the ecosystems based on this study, as they depend strongly on environmental factors and on the actual activity of methanotrophs in situ.     

Active Methanotrophs in Two Contrasting North American Peatland Ecosystems Revealed Using DNA-SIP

The active methanotroph community was investigated in two contrasting North American peatlands, a nutrient-rich sedge fen and nutrient-poor Sphagnum bog using in vitro incubations and ¹³C-DNA stable-isotope probing (SIP) to measure methane (CH₄) oxidation rates and label active microbes followed by fingerprinting and sequencing of bacterial and archaeal 16S rDNA and methane monooxygenase (pmoA and mmoX) genes. Rates of CH₄ oxidation were slightly, but significantly, faster in the bog and methanotrophs belonged to the class Alphaproteobacteria and were similar to other methanotrophs of the genera Methylocystis, Methylosinus, and Methylocapsa or Methylocella detected in, or isolated from, European bogs. The fen had a greater phylogenetic diversity of organisms that had assimilated ¹³C, including methanotrophs from both the Alpha- and Gammaproteobacteria classes and other potentially non-methanotrophic organisms that were similar to bacteria detected in a UK and Finnish fen. Based on similarities between bacteria in our sites and those in Europe, including Russia, we conclude that site physicochemical characteristics rather than biogeography controlled the phylogenetic diversity of active methanotrophs and that differences in phylogenetic diversity between the bog and fen did not relate to measured CH₄ oxidation rates. A single crenarchaeon in the bog site appeared to be assimilating ¹³C in 16S rDNA; however, its phylogenetic similarity to other CO₂-utilizing archaea probably indicates that this organism is not directly involved in CH₄ oxidation in peat.     

Soil–Methanogen Interactions in Two Peatlands (Bog,Fen) in Central New York State

Rates of methanogenesis vary widely in peat soils, yet the reasons are poorly known. We examined rates of methanogenesis and methanogen diversity in relation to soil chemical and biological characteristics in 2 peatlands in New York State. One was an acidic (pH < 4.5) bog dominated by Sphagnum mosses and ericaceous shrubs, although deeper peat was derived from sedges. The other was a fen dominated by Carex lacustris sedges with near-neutral pH soil. At both sites, the most active rates of methanogenesis occurred in the top 20 cm of the peat profile, even when using a substrate-induced methanogenesis technique with added glucose that stimulated rates up to 2 μ mol g ^{? 1} day ^?1 in the bog and 6 μ mol g ^?1 day ^?1 in the fen. Rates of anaerobic CO ₂ production were greater in the bog (0–36 μ mol g ^?1 day ^?1 ) than in the fen (0–5 μ mol g ^?1 day ^?1 ), and added glucose induced greater rates in the sedge-derived peat from the bog than the fen. The peat soil was much more decomposed throughout the profile in the fen. Analysis of chemical elements in the peat profile revealed a striking anomaly: a very high concentration of Pb in surface peat of the bog, which might have constrained methanogenesis. Application of T-RFLP analysis to methanogens revealed dominance by a Methanomicrobiales E2 clade of H ₂ /CO ₂ users in the acidic peat soil of the bog, whereas deeper peat had a different Methanomicrobiales E1 clade, uncultured euryarchaeal rice cluster (RC)-I and RC-II groups, marine benthic group D (MBD) and a new cluster called subaqueous cluster (SC). In contrast, T-RFLP analysis of peat from the fen revealed co-dominance by Methanosaetaceae and Methanomicrobiales E1. The results showed complex relationships between rates of methanogenesis, methanogen populations and metabolic substrate availability with idiosyncratic interactions of trace chemical elements.     

Methane Biogeochemistry and Methanogen Communities in Two Northern Peatland Ecosystems,New York State

Rates of methane (CH₄) production vary considerably among northern peat-forming wetlands, and it is not clear whether variability is caused by environmental factors affecting CH₄ production or differences in methanogen communities. We investigated CH₄ production and emission dynamics concomitantly with 16S rRNA gene sequence-based community analysis of Archaea in two contrasting peat-forming northern wetlands, an ombrotrophic bog and a minerotrophic conifer swamp. Individual measurements of CH₄ emissions to the atmosphere followed a lognormal distribution pattern in both sites, and mean rates were 30× greater in the bog site. Rates of CH₄ production measured in vitro were initially 3× greater in the bog than in the conifer swamp; although, after 30 days of incubation, production rates were similar suggesting that in situ environmental conditions limited production in the conifer swamp. Amplified ribosomal DNA restriction analysis (ARDRA) and rarefaction techniques indicated that both sites had similar levels of archaeal richness, with 27 unique taxa in the bog and 23 taxa in the conifer swamp. However, the bog had more pronounced dominance of a few taxa, whereas the conifer swamp had more even distribution among taxa. A 16S rRNA gene sequence-based phylogenetic analysis indicated high levels of diversity with similarity to known methanogenic families Methanosarcinaceae, Methanosaetaceae, Methanobacteriaceae, and likely Methanomicrobiaceae as well as two additional lineages previously characterized as groups of yet uncultivated Euryarchaeota commonly occurring in flooded rice soils. Therefore, sites with low and high rates of CH₄ production supported very diverse methanogenic communities.     

Pathways for Methanogenesis and Diversity of Methanogenic Archaea in Three Boreal Peatland Ecosystems

The main objectives of this study were to uncover the pathways used for methanogenesis in three different boreal peatland ecosystems and to describe the methanogenic populations involved. The mesotrophic fen had the lowest proportion of CH₄ produced from H₂-CO₂. The oligotrophic fen was the most hydrogenotrophic, followed by the ombrotrophic bog. Each site was characterized by a specific group of methanogenic sequences belonging to Methanosaeta spp. (mesotrophic fen), rice cluster-I (oligotrophic fen), and fen cluster (ombrotrophic bog).     

Controls on Microbial Production of Methane and Carbon Dioxide in Three Sphagnum-Dominated Peatland Ecosystems as Revealed by a Reciprocal Field Peat Transplant Experiment

We examined controls on mineralization of carbon to methane (CH₄) and carbon dioxide (CO₂) in Sphagnum (moss)-dominated peatland ecosystems by transplanting surface (5 cm deep) and subsurface (40 cm deep) peat samples reciprocally among three sites for periods ranging from 4 to 25 months. The sites were Big Run Bog in West Virginia, USA, Bog Lake Bog in Minnesota, USA, and Bog 307 in Ontario, Canada. Immediately upon retrieval, we incubated the peat samples in the laboratory at 12 and 22°C under both anoxic and oxic conditions to estimate rates of carbon mineralization. Transplanting affected surface peat more than subsurface peat. Peat incubated within Bog Lake Bog in Minnesota had the highest rates of CH₄ production, regardless of origin, whereas transplanting did not affect rates of CO₂ production measured concomitantly. Peat that originated in Big Run Bog in West Virginia generally maintained higher rates of CH₄ production and CO₂ production than peat from the other two sites after incubation in the field. The temperature dependence (Q ₁₀) of CH₄ production and CO₂ production varied among transplant sites, but not among peat origins, suggesting physiological adaptations of microbial communities to local environmental conditions. Differences in organic matter quality of the peat, particularly lignin chemistry, helped explain the results: (a) CH₄ production correlated with fresher lignin derived from Carex sedges, and (b) CO₂ production correlated with woody lignin. We concluded that, although both site conditions (climate, nutrient status, and microbial communities) and organic matter quality influence carbon mineralization in peat, interactive effects occur and may differ depending on peat temperature. Moreover, CH₄ production and CO₂ production respond differently to environmental regulators.     

Activity and metabolic regulation of methane production in deep peat profiles of boreal bogs

The potential activity of methane production was determined in the vertical profiles of the peat deposits of three bogs in Tver oblast, which were representative of the boreal zone. In the minerotrophic fen, the rates of methane production measured throughout the profile did not change significantly with depth and comprised 3–6 ng CH₄-C g^?1 h^?1. In ombrotrophic peat bogs, the rate did not exceed 5 ng CH₄-C g^?1 h^?1 in the upper layer of the profile (up to 1.5 m) and increased to 15–30 ng CH₄-C g^?1 h^?1 in the deep layers of the peat deposits. The distribution of fermentative microorganisms and methanogens in the profiles of peat deposits was uniform in all the studied bogs. In bog water samples, the presence of butyrate (up to 14.1 mg 1^?1) and acetate (up to 2.4 mg 1^?1) was revealed throughout the whole profile; in the upper 0.5-m layer of the ombrotrophic bogs, formate (up to 8.9 mg 1^?1) and propionate (up to 0.3 mg 1^?1) were detected as well. The arrangement of local maxima of the fatty acid content and methanogenic activity in the peat deposits, as well as the decrease in the acetate concentrations during summer, support the hypothesis that the initial substrates for methanogenesis come from the upper peat layers. It was established that the addition of sulfate and nitrate inhibits methane production in peat samples; the changes in the concentrations, recorded in situ, may also influence the methane content in peat layers.     

Hydrogenotrophic Methanogenesis by Moderately Acid-Tolerant Methanogens of a Methane-Emitting Acidic Peat

The emission of methane (1.3 mmol of CH₄ m⁻² day⁻¹), precursors of methanogenesis, and the methanogenic microorganisms of acidic bog peat (pH 4.4) from a moderately reduced forest site were investigated by in situ measurements, microcosm incubations, and cultivation methods, respectively. Bog peat produced CH₄ (0.4 to 1.7 μmol g [dry wt] of soil⁻¹ day⁻¹) under anoxic conditions. At in situ pH, supplemental H₂-CO₂, ethanol, and 1-propanol all increased CH₄ production rates while formate, acetate, propionate, and butyrate inhibited the production of CH₄; methanol had no effect. H₂-dependent acetogenesis occurred in H₂-CO₂-supplemented bog peat only after extended incubation periods. Nonsupplemented bog peat initially produced small amounts of H₂ that were subsequently consumed. The accumulation of H₂ was stimulated by ethanol and 1-propanol or by inhibiting methanogenesis with bromoethanesulfonate, and the consumption of ethanol was inhibited by large amounts of H₂; these results collectively indicated that ethanol- or 1-propanol-utilizing bacteria were trophically associated with H₂-utilizing methanogens. A total of 10⁹ anaerobes and 10⁷ hydrogenotrophic methanogens per g (dry weight) of bog peat were enumerated by cultivation techniques. A stable methanogenic enrichment was obtained with an acidic, H₂-CO₂-supplemented, fatty acid-enriched defined medium. CH₄ production rates by the enrichment were similar at pH 4.5 and 6.5, and acetate inhibited methanogenesis at pH 4.5 but not at pH 6.5. A total of 27 different archaeal 16S rRNA gene sequences indicative of Methanobacteriaceae, Methanomicrobiales, and Methanosarcinaceae were retrieved from the highest CH₄-positive serial dilutions of bog peat and methanogenic enrichments. A total of 10 bacterial 16S rRNA gene sequences were also retrieved from the same dilutions and enrichments and were indicative of bacteria that might be responsible for the production of H₂ that could be used by hydrogenotrophic methanogens. These results indicated that in this acidic bog peat, (i) H₂ is an important substrate for acid-tolerant methanogens, (ii) interspecies hydrogen transfer is involved in the degradation of organic carbon, (iii) the accumulation of protonated volatile fatty acids inhibits methanogenesis, and (iv) methanogenesis might be due to the activities of methanogens that are phylogenetic members of the Methanobacteriaceae, Methanomicrobiales, and Methanosarcinaceae.     

Effects of a transient oxic period on mineralization of organic matter to CH4 and CO2 in anoxic peat incubations

Rates of organic matter mineralization in peatlands, and hence production of the greenhouse gases CH₄ and CO₂, are highly dependent on the distribution of oxygen in the peat. Using laboratory incubations of peat, we investigated the sensitivity of the anoxic production of CH₄ and CO₂ to a transient oxic period of a few weeks’ duration. Production rates during 3 successive anoxic periods were compared with rates in samples incubated in the presence of oxygen during the second period. In surface peat (5–10‐cm depth), with an initially high level of CH₄ production, oxic conditions during period 2 did not result in a lower potential CH₄ production rate during period 3, although production was delayed ～1 week. In permanently anoxic, deep peat (50–55‐cm depth) with a comparatively low initial production of CH₄, oxic conditions during period 2 resulted in zero production of CH₄ during period 3. Thus, the methanogens in surface peal—but not in deep peat—remained viable after several weeks of oxic conditions. In contrast to CH₄ production, the oxic period had a negligible effect on anoxic CO₂ production during period 3, in surface as well as deep peat. In both surface and deep peat, CO₂ production was several times higher under oxic than under anoxic conditions. However, for the first 2 weeks of oxic conditions, CO₂ production in the deep peat was very low. Still, deep peat obviously contained facultative microorganisms that, after a relatively short period, were able to maintain a considerably higher rate of organic matter mineralization under oxic than under anoxic conditions.     

Partitioning pathways of CO2 production in peatlands with stable carbon isotopes

Although methanogenic pathways generally produce equimolar amounts of carbon dioxide and methane, CO₂ concentrations are often reported to be higher than CH₄ concentrations in both field and laboratory incubation studies of peat decomposition. In field settings, higher pore water concentrations of CO₂ may result from the loss of methane by: (1) ebullition due to the low solubility of methane in pore water and (2) vascular-plant transport. Higher CO₂ concentrations may also be caused by: (1) production of additional CO₂ by high-molecular weight (HMW) organic matter (OM) fermentation and/or (2) respiration from non-methanogenic pathways. In this study of a peatland where advection and transverse dispersion were the dominant pore water solute transport mechanisms, an isotope-mass balance approach was used to determine the proportions of CO₂ formed from non-fractionating OM respiration and HMW fermentation relative to CO₂ production from methanogenesis. This approach also allowed us to estimate the loss of CH₄ from the belowground system. The pathways of CO₂ production varied with depth and surface vegetation type. In a Carex-dominated fen, methane production initially produced 40 % of the total CO₂ and then increased to 90–100 % with increasing depth. In a Sphagnum-dominated bog, methanogenesis resulted in 60 % of total CO₂ production which increased to 100 % at depth. Both bogs and fens showed 85–100 % of methane loss from pore waters. Our results indicate that the isotopic composition of dissolved CO₂ is a powerful indicator to allow partitioning of the processes affecting peat remineralization and methane production.