Patterns of Fungal and Bacterial Carbon Mineralization Across Northern European Peatlands

The fungal and bacterial activity was determined in 20 northern European peatlands ranging from ombrotrophic bogs to eutrophic fens with key differences in degree of humification, pH, dry bulk density, carbon (C) content and vegetation communities using the selective inhibition (SI) technique. These peatlands were partly disturbed and the respective water tables lowered below the surface layer. Basal respiration ranged from 24 to 128 µg CO₂-C g^?1 dry peat d^?1. Bacterial contributions to CO₂ production were high in most peatlands and showed the following pattern: eutrophic >> transitional ≥ mesotrophic >> ombrotrophic peatland types. The fungal-to-bacterial (F:B) ratios varied substantially within peatland type, and this was mainly attributed to differences in peat botanical compositions and chemistry. The computed mean Inhibitor Additivity Ratio (IAR) was quite close to 1 to suggest that the SI techniques can be used to partition eukaryotic and prokaryotic activity in wide range of peatlands. Overall, basal respiration, microbial biomass-C, fungal and bacterial activities varied across the studied peatland types, and such differences could have consequences for C- and nutrient-cycling as well as how bogs and fens will respond to environmental changes.     

Microbial activity across a boreal peatland nutrient gradient: the role of fungi and bacteria

Microbes involved in decomposition within peatlands and the conditions that influence their activities have implications for C and greenhouse gas exchange. The objectives of this research were to characterize the role of fungal and bacterial activities in peatlands using selective antibiotic inhibition techniques across a nutrient gradient (rich to poor fens) and to search for environmental controls on the activity of each group. Bacterial activities predominated across a range of rich to poor boreal peatlands in central Ontario, Canada, although fungal activity became increasingly important in the poor sites. Linkages between soil pH and nutrient status and fungal and bacterial activities were found. However, they did not confirm our initial hypotheses that bacterial activity would be low in poor sites due to proton stress and low nutrient (particularly N) availability, whereas, fungal activity would be low in rich sites due to increased competitive ability of bacteria under near neutral pH conditions and high nutrient availability. Further work across these sites aimed at characterizing the phylogeny of the rhizosphere fungi is needed to determine if increased presence of mycorrhizae in poor sites could have explained our observed patterns. However, regardless of fungal: bacterial activity ratio differences across sites and its associated controls, microbial CO₂ production rates across fen types did not vary significantly, suggesting that the proportion of bacteria and fungi may not matter to broader carbon cycling and greenhouse gas emissions in peat soils.     

Responses of aerobic microbial communities and soil respiration to water-level drawdown in a northern boreal fen

On a global basis, peatlands are a major reserve of carbon (C). Hydrological changes can affect the decomposition processes in peatlands and in turn can alter their C balance. Since 1959, a groundwater extraction plant has generated a water-level gradient at our study site that has gradually changed part of the wet fen into a dry peatland forest. The average water-level drawdown of the gradient (from a pristine 9 cm to 26 cm in the dry end) is close to an estimate predicted by an increase in mean global temperature of 3°C. We studied the total microbial community of the aerobic surface peat in four locations along the gradient through phospholipid fatty acid and PCR-DGGE methods. Additionally, field measurements of soil respiration showed a threefold increase in the C-emission rate at the driest location compared with the wettest one, indicating enhanced decomposition. Also, both fungal and bacterial biomass increased in the drier locations. At the species level, the fungal community changed due to water-level drawdown whereas actinobacteria were less sensitive to drying. The majority of fungal sequences were similar to ectomycorrhizal (ECM) fungi, which dominated throughout the gradient. Our results indicate that ECM fungi might act as important facultative decomposers in organic-rich environments such as peatlands.     

Microbial community structure and activity linked to contrasting biogeochemical gradients in bog and fen environments of the glacial lake agassiz peatland

The abundances, compositions, and activities of microbial communities were investigated at bog and fen sites in the Glacial Lake Agassiz Peatland of northwestern Minnesota. These sites contrast in the reactivity of dissolved organic matter (DOM) and the presence or absence of groundwater inputs. Microbial community composition was characterized using pyrosequencing and clone library construction of phylogenetic marker genes. Microbial distribution patterns were linked to pH, concentrations of dissolved organic carbon and nitrogen, C/N ratios, optical properties of DOM, and activities of laccase and peroxidase enzymes. Both bacterial and archaeal richness and rRNA gene abundance were >2 times higher on average in the fen than in the bog, in agreement with a higher pH, labile DOM content, and enhanced enzyme activities in the fen. Fungi were equivalent to an average of 1.4% of total prokaryotes in gene abundance assayed by quantitative PCR. Results revealed statistically distinct spatial patterns between bacterial and fungal communities. Fungal distribution did not covary with pH and DOM optical properties and was vertically stratified, with a prevalence of Ascomycota and Basidiomycota near the surface and much higher representation of Zygomycota in the subsurface. In contrast, bacterial community composition largely varied between environments, with the bog dominated by Acidobacteria (61% of total sequences), while the Firmicutes (52%) dominated in the fen. Acetoclastic Methanosarcinales showed a much higher relative abundance in the bog, in contrast to the dominance of diverse hydrogenotrophic methanogens in the fen. This is the first quantitative and compositional analysis of three microbial domains in peatlands and demonstrates that the microbial abundance, diversity, and activity parallel with the pronounced differences in environmental variables between bog and fen sites.     

Comparison of carbon and nitrogen accumulation rate between bog and fen phases in a pristine peatland with the fen-bog transition

Long-term carbon and nitrogen dynamics in peatlands are affected by both vegetation production and decomposition processes. Here, we examined the carbon accumulation rate (CAR), nitrogen accumulation rate (NAR) and δ¹³C, δ¹⁵N of plant residuals in a peat core dated back to ~8500 cal year BP in a temperate peatland in Northeast China. Impacted by the tephra during 1160 and 789 cal year BP and climate change, the peatland changed from a fen dominated by vascular plants to a bog dominated by Sphagnum mosses. We used the Clymo model to quantify peat addition rate and decay constant for acrotelm and catotelm layers during both bog and fen phases. Our studied peatland was dominated by Sphagnum fuscum during the bog phase (789 to −59 cal year BP) and lower accumulation rates in the acrotelm layer was found during this phase, suggesting the dominant role of volcanic eruption in the CAR of the peat core. Both mean CAR and NAR were higher during the bog phase than during the fen phase in our study, consistent with the results of the only one similar study in the literature. Because the input rate of organic matter was considered to be lower during the bog phase, the decomposition process must have been much lower during the bog phase than during the fen phase and potentially controlled CAR and NAR. During the fen phase, CAR was also lower under higher temperature and summer insolation, conditions beneficial for decomposition. δ¹⁵N of Sphagnum hinted that nitrogen fixation had a positive effect on nitrogen accumulation, particular in recent decades. Our study suggested that decomposition is more important for carbon and nitrogen sequestration than production in peatlands in most conditions and if future climate changes or human disturbance increase decomposition rate, carbon sequestration in peatlands will be jeopardized.     

Histosol as an ecologically active constituent of peatland: a case study from Estonia

This study clarifies the area distribution of Estonian peat soils by three factors: main peat soil groups, peat thickness and peat decomposition degree. A digital soil map (1:10,000) and supplementary database were used for summarizing the distribution of peat soils. From the combined database with 859,701 polygons the soil mapping unit code, formula of soil texture (including peat) and formula of epipedon fabric were compiled using the MapInfo software. Peat soils form altogether 10,038 km² or 23.5% of the total Estonian soil cover. From the peat soils the fen soils form 59.0%, bog soils 21.7% and transitional bog soils 14.7%. 45% of peat soils are well, 26% moderately and 29% slightly decomposed, by the peat thickness 13% are very shallow, 21% shallow and 66% thick. The general ecological characterization of peat soils and their mutual relationship with plant cover are given for the main peat soil taxa. The dominant natural ecosystems formed on peatlands are: (1) mixed birch, alder, spruce and pine forests on thin (<100 cm) well decomposed eutrophic fen soils, and (2) a sparse pine forests and hummock-ridge-hollow raised bogs wooded sparsely by pine on thick (>100 cm) slightly decomposed oligotrophic bog soils. The accumulation of organic carbon in peatlands soil cover (0–50 cm) totals 269.4±12.7 Tg and in epipedon layer (as superficial part of soil cover; 0–30 cm) 129.9±5.8 Tg. The former is sequestrated into 543.7 Tg of peat, which forms 22.9% of the total Estonian peat resources (2.37 Pg).     

Vertical profiles of methanogenesis and methanogens in two contrasting acidic peatlands in central New York State, USA

Northern acidic peatlands are important sources of atmospheric methane, yet the methanogens in them are poorly characterized. We examined methanogenic activities and methanogen populations at different depths in two peatlands, McLean bog (MB) and Chicago bog (CB). Both have acidic (pH 3.5-4.5) peat soils, but the pH of the deeper layers of CB is near-neutral, reflecting its previous existence as a neutral-pH fen. Acetotrophic and hydrogenotrophic methanogenesis could be stimulated in upper samples from both bogs, and phylotypes of methanogens using H2/CO2 (Methanomicrobiales) or acetate (Methanosarcinales) were identified in 16S rRNA gene clone libraries and by terminal restriction fragment length polymorphism (T-RFLP) analyses using a novel primer/restriction enzyme set that we developed. Particularly dominant in the upper layers was a clade in the Methanomicrobiales, called E2 here and the R10 or fen group elsewhere, estimated by quantitative polymerase chain reaction to be present at approximately 10(8) cells per gram of dry peat. Methanogenic activity was considerably lower in deeper samples from both bogs. The methanogen populations detected by T-RFLP in deeper portions of MB were mainly E2 and the uncultured euryarchaeal rice cluster (RC)-II group, whereas populations in the less acidic CB deep layers were considerably different, and included a Methanomicrobiales clade we call E1-E1', as well as RC-I, RC-II, marine benthic group D, and a new cluster that we call the subaqueous cluster. E2 was barely detectable in the deeper samples from CB, further evidence for the associations of most organisms in this group with acidic habitats.     

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.     

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.     

Bacterial populations and environmental factors controlling cellulose degradation in an acidic Sphagnum peat

Northern peatlands represent a major global carbon store harbouring approximately one-third of the global reserves of soil organic carbon. A large proportion of these peatlands consists of acidic Sphagnum-dominated ombrotrophic bogs, which are characterized by extremely low rates of plant debris decomposition. The degradation of cellulose, the major component of Sphagnum-derived litter, was monitored in long-term incubation experiments with acidic (pH 4.0) peat extracts. This process was almost undetectable at 10°C and occurred at low rates at 20°C, while it was significantly accelerated at both temperature regimes by the addition of available nitrogen. Cellulose breakdown was only partially inhibited in the presence of cycloheximide, suggesting that bacteria participated in this process. We aimed to identify these bacteria by a combination of molecular and cultivation approaches and to determine the factors that limit their activity in situ. The indigenous bacterial community in peat was dominated by Alphaproteobacteria and Acidobacteria. The addition of cellulose induced a clear shift in the community structure towards an increase in the relative abundance of the Bacteroidetes. Increasing temperature and nitrogen availability resulted in a selective development of bacteria phylogenetically related to Cytophaga hutchinsonii (94-95% 16S rRNA gene sequence similarity), which densely colonized microfibrils of cellulose. Among isolates obtained from this community only some subdivision 1 Acidobacteria were capable of degrading cellulose, albeit at a very slow rate. These Acidobacteria represent indigenous cellulolytic members of the microbial community in acidic peat and are easily out-competed by Cytophaga-like bacteria under conditions of increased nitrogen availability. Members of the phylum Firmicutes, known to be key players in cellulose degradation in neutral habitats, were not detected in the cellulolytic community enriched at low pH.     

Cotton-Grass and Blueberry have Opposite Effect on Peat Characteristics and Nutrient Transformation in Peatland

Peatlands are large repositories of carbon (C). Sphagnum mosses play a key role in C sequestration, whereas the presence of vascular plants is generally thought to stimulate peat decomposition. Recent studies stress the importance of plant species for peat quality and soil microbial activity. Thus, learning about specific plant–microbe–soil relations and their potential feedbacks for C and nutrient cycling are important for a correct understanding of C sequestration in peatlands and its potential shift associated with vegetation change. We studied how the long-term presence of blueberry and cotton-grass, the main vascular dominants of spruce swamp forests, is reflected in the peat characteristics, soil microbial biomass and activities, and the possible implications of their spread for nutrient cycling and C storage in these systems. We showed that the potential effect of vascular plants on ecosystem functioning is species specific and need not necessarily result in increased organic matter decomposition. Although the presence of blueberry enhanced phosphorus availability, soil microbial biomass and the activities of C-acquiring enzymes, cotton-grass strongly depleted phosphorus and nitrogen from the peat. The harsh conditions and prevailing anoxia retarded the decomposition of cotton-grass litter and caused no significant enhancement in microbial biomass and exoenzymatic activity. Therefore, the spread of blueberry in peatlands may stimulate organic matter decomposition and negatively affect the C sequestration process, whereas the potential spread of cotton-grass would not likely change the functioning of peatlands as C sinks.     

Acidobacteria in Fens: Phylogenetic Diversity and Genome Analysis of the Key Representatives

Microbiology - Acidobacteriota are one of the major prokaryotic groups in soils and peatlands. They are especially abundant in acidic peat bogs, where representatives of the well-characterized...     

Abundance, biomass, structure, and activity of the microbial complexes of minerotrophic and ombrotrophic peatlands

Very large microbial biomass was revealed in peat bogs by means of fluorescence microscopy. In ombrotrophic peatlands, the pool of the dry-weight microbial biomass in the 1.5-m layer constituted 3-4 t/ha and was twice as high as in the minerotrophic peat bogs. Fungal biomass was predominant (55-99%) in ombrotrophic peatlands, while bacterial biomass predominated in minerotrophic peatlands (55-86%). In ombrotrophic peatlands, the microbial biomass was concentrated in the upper layers, while in minerotrophic peatlands, it was uniformly distributed in the bulk. After drainage, the microbial pool in the ombrotrophic peatlands increased twofold; that in the minerotrophic peatlands remained at the same level. The potential activity of nitrogen fixation and denitrification was revealed across the whole profile of the peatlands. The average values of these potential activities were five times higher in the minerotrophic peatlands, where bacterial biomass predominated.     

Functional-Structural Analysis of Nitrogen-Cycle Bacteria in a Hypersaline Mat from the Omani Desert

Anaerobic microbial activity in northern peat soils most often results in more carbon dioxide (CO ₂ ) production than methane (CH₄) production. This study examined why methanogenic conditions (i.e., equal molar amounts of CH₄ production and CO₂ production) prevail so infrequently. We used peat soils from two ombrotrophic bogs and from two rheotrophic fens. The former two represented a relatively dry bog hummock and a wet bog hollow, and the latter two represented a forested fen and a sedge-dominated fen. We quantified gas production rates in soil samples incubated in vitro with and without added metabolic substrates (glucose, ethanol, H₂/CO₂). None of the peat soils exhibited methanogenic conditions when incubated in vitro for a short time (< 5 days) and without added substrates. Incubating some samples > 50 days without added substrates led to methanogenic conditions in only one of four experiments. The anaerobic CO₂:CH₄ production ratio ranged from 5:1 to 40:1 in peat soil without additions and was larger in samples from the dry bog hummock and forested fen than the wet bog hollow and sedge fen. Adding ethanol or glucose separately to peat soils led to methanogenic conditions within 5 days after the addition by stimulating rates of CH₄ production, suggesting CH₄ production from both hydrogenotrophic and acetoclastic methanogenesis. Our results suggest that methanogenic conditions in peat soils rely on a constant supply of easily decomposable metabolic substrates. Sample handling and incubation procedures might obscure methanogenic conditions in peat soil incubated in vitro.     

Abundance,biomass, structure,and activity of the microbial complexes of minerotrophic and ombrotrophic peatlands

A very large microbial biomass was revealed in peat bogs by means of fluorescence microscopy. In ombrotrophic peatlands, the pool of the dry-weight microbial biomass in the 1.5-m layer constituted 3–4 t/ha and was twice as high as in the minerotrophic peat bogs. Fungal biomass was predominant (55–99%) in ombrotrophic peatlands, while bacterial biomass predominated (55–86%) in minerotrophic peatlands. In ombrotrophic peatlands, the microbial biomass was concentrated in the upper layers, while in minerotrophic peatlands, it was uniformly distributed in the bulk. After drainage, the microbial pool in the ombrotrophic peatlands increased twofold; that in the minerotrophic peatlands remained at the same level. The potential activity of nitrogen fixation and denitrification was revealed across the whole profile of the peatlands. The average values of these potential activities were five times higher in the minerotrophic peatlands, where bacterial biomass predominated.     

Bacterial and Fungal Communities in a Degraded Ombrotrophic Peatland Undergoing Natural and Managed Re-Vegetation

The UK hosts 15–19% of global upland ombrotrophic (rain fed) peatlands that are estimated to store 3.2 billion tonnes of carbon and represent a critical upland habitat with regard to biodiversity and ecosystem services provision. Net production is dependent on an imbalance between growth of peat-forming Sphagnum mosses and microbial decomposition by microorganisms that are limited by cold, acidic, and anaerobic conditions. In the Southern Pennines, land-use change, drainage, and over 200 years of anthropogenic N and heavy metal deposition have contributed to severe peatland degradation manifested as a loss of vegetation leaving bare peat susceptible to erosion and deep gullying. A restoration programme designed to regain peat hydrology, stability and functionality has involved re-vegetation through nurse grass, dwarf shrub and Sphagnum re-introduction. Our aim was to characterise bacterial and fungal communities, via high-throughput rRNA gene sequencing, in the surface acrotelm/mesotelm of degraded bare peat, long-term stable vegetated peat, and natural and managed restorations. Compared to long-term vegetated areas the bare peat microbiome had significantly higher levels of oligotrophic marker phyla (Acidobacteria, Verrucomicrobia, TM6) and lower Bacteroidetes and Actinobacteria, together with much higher ligninolytic Basidiomycota. Fewer distinct microbial sequences and significantly fewer cultivable microbes were detected in bare peat compared to other areas. Microbial community structure was linked to restoration activity and correlated with soil edaphic variables (e.g. moisture and heavy metals). Although rapid community changes were evident following restoration activity, restored bare peat did not approach a similar microbial community structure to non-eroded areas even after 25 years, which may be related to the stabilisation of historic deposited heavy metals pollution in long-term stable areas. These primary findings are discussed in relation to bare peat oligotrophy, re-vegetation recalcitrance, rhizosphere-microbe-soil interactions, C, N and P cycling, trajectory of restoration, and ecosystem service implications for peatland restoration.     

Changes in Fungal Community Composition in Response to Vegetational Succession During the Natural Regeneration of Cutover Peatlands

Despite the importance of peatlands as a major store of sequestered carbon and the role of fungi in releasing sequestered C, we know little about the community structure of fungi in peatlands. We investigated these across a gradient of naturally regenerating peatland vegetation using denaturing gradient gel electrophoresis (DGGE) and clone libraries of fragments of the fungal rRNA internal transcribed spacer (ITS) region. Significant changes in the fungal community structure of peat samples at different stages of regeneration were observed, which relate to the composition of the vegetation recolonizing these sites. Cloning and sequence analysis also demonstrated a potential shift in the relative abundance of the main fungal phyla. Some of the clones identified to genus level were highly related to fungi known to play a role in the degradation of plant litter or wood in similar ecosystems and/or form mycorrhizal associations. In addition, several fungal isolates highly related to peat clones were obtained, and their enzymic capacity to degrade structural plant tissues was assessed. Together, these results suggest that the fungal community composition of peat may be an important indicator of the status of regeneration and potential carbon sequestration of cutover peatlands.     

北方泥炭地氮素有效性的变化及其对碳汇功能的影响

北方泥炭地是典型的氮限制性生态系统,对全球气候变化及人类活动响应敏感。气候变暖导致内源有效氮增加以及人类活动引起的大量外源氮输入,改变了北方泥炭地氮素有效性,对泥炭地碳氮循环过程及碳汇功能产生了深远影响。本文综述了北方泥炭地碳积累速率和碳汇功能的影响因素,分析了氮沉降、冻融、火烧等因素对北方泥炭地氮素有效性的影响,分别从碳固定和碳排放过程阐述了植物及土壤微生物对氮素有效性变化的响应,并对全球变化影响下泥炭生态系统碳汇功能相关研究进行了展望,以期助力“双碳”目标的实施。