Plant phenology and species-specific traits control plant CH4 emissions in a northern boreal fen.

Aerenchymatic transport is an important mechanism through which plants affect methane (CH4 ) emissions from peatlands. Controlling environmental factors and the effects of plant phenology remain, however, uncertain. We identified factors controlling seasonal CH4 flux rate and investigated transport efficiency (flux rate per unit of rhizospheric porewater CH4 concentration). We measured CH4 fluxes through individual shoots of Carex rostrata, Menyanthes trifoliata, Betula nana and Salix lapponum throughout growing seasons in 2020 and 2021 and Equisetum fluviatile and Comarum palustre in high summer 2021 along with water-table level, peat temperature and porewater CH4 concentration. CH4 flux rate of C. rostrata was related to plant phenology and peat temperature. Flux rates of M. trifoliata and shrubs B. nana and S. lapponum were insensitive to the investigated environmental variables. In high summer, flux rate and efficiency were highest for C. rostrata (6.86 mg m-2  h-1 and 0.36 mg m-2  h-1 (µmol l-1 )-1 , respectively). Menyanthes trifoliata showed a high flux rate, but limited efficiency. Low flux rates and efficiency were detected for the remaining species. Knowledge of the species-specific CH4 flux rate and their different responses to plant phenology and environmental factors can significantly improve the estimation of ecosystem-scale CH4 dynamics in boreal peatlands.

Site fertility and soil water-table level affect fungal biomass production and community composition in boreal peatland forests.

A substantial amount of below-ground carbon (C) is suggested to be associated with fungi, which may significantly affect the soil C balance in forested ecosystems. Ergosterol from in-growth mesh bags and litterbags was used to estimate fungal biomass production and community composition in drained peatland forests with differing fertility. Extramatrical mycelia (EMM) biomass production was generally higher in the nutrient-poor site, increased with deeper water table level and decreased along the length of the recovery time. EMM biomass production was of the same magnitude as in mineral-soil forests. Saprotrophic fungal biomass production was higher in the nutrient-rich site. Both ectomycorrhizal (ECM) and saprotrophic fungal community composition changed according to site fertility and water table level. ECM fungal community composition with different exploration types may explain the differences in fungal biomass production between peatland forests. Melanin-rich Hyaloscypha may indicate decreased turnover of biomass in nutrient-rich young peatland forest. Genera Lactarius and Laccaria may be important in nutrient rich and Piloderma in the nutrient-poor conditions, respectively. Furthermore, Paxillus involutus and Cortinarius sp. may be important generalists in all sites and responsible for EMM biomass production during the first summer months. Saprotrophs showed a functionally more diverse fungal community in the nutrient-rich site.

Warming impacts on boreal fen CO2 exchange under wet and dry conditions.

Northern peatlands form a major soil carbon (C) stock. With climate change, peatland C mineralization is expected to increase, which in turn would accelerate climate change. A particularity of peatlands is the importance of soil aeration, which regulates peatland functioning and likely modulates the responses to warming climate. Our aim is to assess the impacts of warming on a southern boreal and a sub-arctic sedge fen carbon dioxide (CO2 ) exchange under two plausible water table regimes: wet and moderately dry. We focused this study on minerotrophic treeless sedge fens, as they are common peatland types at boreal and (sub)arctic areas, which are expected to face the highest rates of climate warming. In addition, fens are expected to respond to environmental changes faster than the nutrient poor bogs. Our study confirmed that CO2 exchange is more strongly affected by drying than warming. Experimental water level draw-down (WLD) significantly increased gross photosynthesis and ecosystem respiration. Warming alone had insignificant impacts on the CO2 exchange components, but when combined with WLD it further increased ecosystem respiration. In the southern fen, CO2 uptake decreased due to WLD, which was amplified by warming, while at northern fen it remained stable. As a conclusion, our results suggest that a very small difference in the WLD may be decisive, whether the C sink of a fen decreases, or whether the system is able to adapt within its regime and maintain its functions. Moreover, the water table has a role in determining how much the increased temperature impacts the CO2 exchange.

Responses of phenology and biomass production of boreal fens to climate warming under different water-table level regimes.

Climate change affects peatlands directly through increased air temperatures and indirectly through changes in water-table level (WL). The interactions of these two still remain poorly known. We determined experimentally the separate and interactive effects of temperature and WL regime on factors of relevance for the inputs to the carbon cycle: plant community composition, phenology, biomass production, and shoot:root allocation in two wet boreal sedge-dominated fens, "southern" at 62°N and "northern" at 68°Ν. Warming (1.5°C higher average daily air temperature) was induced with open-top chambers and WL drawdown (WLD; 3-7 cm on average) by shallow ditches. Total biomass production varied from 250 to 520 g/m2 , with belowground production comprising 25%-63%. Warming was associated with minor effects on phenology and negligible effects on community composition, biomass production, and allocation. WLD clearly affected the contribution of different plant functional types (PFTs) in the community and the biomass they produced: shrubs benefited while forbs and mosses suffered. These responses did not depend on the warming treatment. Following WLD, aboveground biomass production decreased mainly due to reduced growth of mosses in the southern fen. Aboveground vascular plant biomass production remained unchanged but the contribution of different PFTs changed. The observed changes were also reflected in plant phenology, with different PFTs showing different responses. Belowground production increased following WLD in the northern fen only, but an increase in the contributions of shrubs and forbs was observed in both sites, while sedge contribution decreased. Moderate warming alone seems not able to drive significant changes in plant productivity or community composition in these wet ecosystems. However, if warming is accompanied by even modest WL drawdown, changes should be expected in the relative contribution of PFTs, which could lead to profound changes in the function of fens. Consequently, hydrological scenarios are of utmost importance when estimating their future function.

Plant-mediated CH4 exchange in wetlands: A review of mechanisms and measurement methods with implications for modelling.

Plant-mediated CH4 transport (PMT) is the dominant pathway through which soil-produced CH4 can escape into the atmosphere and thus plays an important role in controlling ecosystem CH4 emission. PMT is affected by abiotic and biotic factors simultaneously, and the effects of biotic factors, such as the dominant plant species and their traits, can override the effects of abiotic factors. Increasing evidence shows that plant-mediated CH4 fluxes include not only PMT, but also within-plant CH4 production and oxidation due to the detection of methanogens and methanotrophs attached to the shoots. Despite the inter-species and seasonal differences, and the probable contribution of within-plant microbes to total plant-mediated CH4 exchange (PME), current process-based ecosystem models only estimate PMT based on the bulk biomass or leaf area index of aerenchymatous plants. We highlight five knowledge gaps to which more research efforts should be devoted. First, large between-species variation, even within the same family, complicates general estimation of PMT, and calls for further work on the key dominant species in different types of wetlands. Second, the interface (rhizosphere-root, root-shoot, or leaf-atmosphere) and plant traits controlling PMT remain poorly documented, but would be required for generalizations from species to relevant functional groups. Third, the main environmental controls of PMT across species remain uncertain. Fourth, the role of within-plant CH4 production and oxidation is poorly quantified. Fifth, the simplistic description of PMT in current process models results in uncertainty and potentially high errors in predictions of the ecosystem CH4 flux. Our review suggest that flux measurements should be conducted over multiple growing seasons and be paired with trait assessment and microbial analysis, and that trait-based models should be developed. Only then we are capable to accurately estimate plant-mediated CH4 emissions, and eventually ecosystem total CH4 emissions at both regional and global scales.

Total Soil CO2 Efflux from Drained Terric Histosols.

Histosols cover about 8-10% of Lithuania's territory and most of this area is covered with nutrient-rich organic soils (Terric Histosols). Greenhouse gas (GHG) emissions from drained Histosols contribute more than 25% of emissions from the Land Use, Land Use Change and Forestry (LULUCF) sector. In this study, as the first step of examining the carbon dioxide (CO2) fluxes in these soils, total soil CO2 efflux and several environmental parameters (temperature of air and topsoil, soil chemical composition, soil moisture, and water table level) were measured in drained Terric Histosols under three native forest stands and perennial grasslands in the growing seasons of 2020 and 2021. The drained nutrient-rich organic soils differed in terms of concentrations of soil organic carbon and total nitrogen, as well as soil organic carbon and total nitrogen ratio. The highest rate of total soil CO2 efflux was found in the summer months. Overall, the rate was statistically significant and strongly correlated only with soil and air temperature. A trend emerged that total soil CO2 efflux was 30% higher in perennial grassland than in forested land. Additional work is still needed to estimate the net CO2 balance of these soils.

Fine-root biomass production and its contribution to organic matter accumulation in sedge fens under changing climate.

Climate change may affect the carbon sink function of peatlands through warming and drying. Fine-root biomass production (FRBP) of sedge fens, a widespread peatland habitat, is important in this context, since most of the biomass is below ground in these ecosystems. We examined the response of fine-root biomass production, depth distribution (10 cm intervals down to 60 cm), chemical characteristics, and decomposition along with other main litter types (sedge leaves, Sphagnum moss shoots) to an average May-to-October warming of 1.7 °C above ambient daily mean temperature and drying of 2-8 cm below ambient soil water-table level (WL) in two sedge fens situated in Northern and Southern Boreal zones. Warming was induced with open top chambers and drying with shallow ditching. Finally, we simulated short-term organic matter (OM) accumulation using net primary production and mass loss data. Total FRBP, and FRBP in deeper layers, was clearly higher in southern than northern fen. Drying significantly increased, and warming marginally increased, total FRBP, while warming significantly increased, and drying marginally increased, the proportional share of FRBP in deeper layers. Drying, especially, modified root chemistry as the relative proportions of fats, wax, lipids, lignin and other aromatics increased while the proportion of polysaccharides decreased. Warming did not affect the decomposition of any litter types, while drying reduced the decomposition of sedge leaf litter. Although drying increased OM accumulation from root litter at both fens, total OM accumulation decreased at the southern fen, while the northern fen with overall lower values showed no such pattern. Our results suggest that in warmer and/or modestly drier conditions, sedge fen FRBP will increase and/or be allocated to deeper soil layers. These changes along with the altered litter inputs may sustain the soil carbon sink function through OM accumulation, unless the WL falls below a tipping point.

Soil GHG dynamics after water level rise - Impacts of selection harvesting in peatland forests.

Managed boreal peatlands are widespread and economically important, but they are a large source of greenhouse gases (GHGs). Peatland GHG emissions are related to soil water-table level (WT), which controls the vertical distribution of aerobic and anaerobic processes and, consequently, sinks and sources of GHGs in soils. On forested peatlands, selection harvesting reduces stand evapotranspiration and it has been suggested that the resulting WT rise decreases soil net emissions, while the tree growth is maintained. We monitored soil concentrations of CO2, CH4, N2O and O2 by depth down to 80 cm, and CO2 and CH4 fluxes from soil in two nutrient-rich Norway spruce dominated peatlands in Southern Finland to examine the responses of soil GHG dynamics to WT rise. Selection harvesting raised WT by 14 cm on both sites, on average, mean WTs of the monitoring period being 73 cm for unharvested control and 59 cm for selection harvest. All soil gas concentrations were associated with proximity to WT. Both CH4 and CO2 showed remarkable vertical concentration gradients, with high values in the deepest layer, likely due to slow gas transfer in wet peat. CH4 was efficiently consumed in peat layers near and above WT where it reached sub-atmospheric concentrations, indicating sustained oxidation of CH4 from both atmospheric and deeper soil origins also after harvesting. Based on soil gas concentration data, surface peat (top 25/30 cm layer) contributed most to the soil-atmosphere CO2 fluxes and harvesting slightly increased the CO2 source in deeper soil (below 45/50 cm), which could explain the small CO2 flux differences between treatments. N2O production occurred above WT, and it was unaffected by harvesting. Overall, the WT rise obtained with selection harvesting was not sufficient to reduce soil GHG emissions, but additional hydrological regulation would have been needed.

Quantification of Plant Root Species Composition in Peatlands Using FTIR Spectroscopy.

Evidence of plant root biomass and production in peatlands at the level of species or plant functional type (PFT) is needed for defining ecosystem functioning and predicting its future development. However, such data are limited due to methodological difficulties and the toilsomeness of separating roots from peat. We developed Fourier transform infrared (FTIR) spectroscopy based calibration models for quantifying the mass proportions of several common peatland species, and alternatively, the PFTs that these species represented, in composite root samples. We further tested whether woody roots could be classified into diameter classes, and whether dead and living roots could be separated. We aimed to solve whether general models applicable in different studies can be developed, and what would be the best way to build such models. FTIR spectra were measured from dried and powdered roots: both "pure roots", original samples of 25 species collected in the field, and "root mixtures", artificial composite samples prepared by mixing known amounts of pure roots of different species. Partial least squares regression was used to build the calibration models. The general applicability of the models was tested using roots collected in different sites or times. Our main finding is that pure roots can replace complex mixtures as calibration data. Using pure roots, we constructed generally applicable models for quantification of roots of the main PFTs of northern peatlands. The models provided accurate estimates even for far distant sites, with root mean square error (RMSE) 1.4-6.6% for graminoids, forbs and ferns. For shrubs and trees the estimates were less accurate due to higher within-species heterogeneity, partly related to variation in root diameter. Still, we obtained RMSE 3.9-10.8% for total woody roots, but up to 20.1% for different woody-root types. Species-level and dead-root models performed well within the calibration dataset but provided unacceptable estimates for independent samples, limiting their routine application in field conditions. Our PFT-level models can be applied on roots separated from soil for biomass determination or from ingrowth cores for estimating root production. We present possibilities for further development of species-level or dead-root models using the pure-root approach.

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 degrees 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 ecology in a future climate: effects of temperature and moisture on microbial communities of two boreal fens.

Impacts of warming with open-top chambers on microbial communities in wet conditions and in conditions resulting from moderate water-level drawdown (WLD) were studied across 0-50 cm depth in northern and southern boreal sedge fens. Warming alone decreased microbial biomass especially in the northern fen. Impact of warming on microbial PLFA and fungal ITS composition was more obvious in the northern fen and linked to moisture regime and sample depth. Fungal-specific PLFA increased in the surface peat in the drier regime and decreased in layers below 10 cm in the wet regime after warming. OTUs representing Tomentella and Lactarius were observed in drier regime and Mortierella in wet regime after warming in the northern fen. The ectomycorrhizal fungi responded only to WLD. Interestingly, warming together with WLD decreased archaeal 16S rRNA copy numbers in general, and fungal ITS copy numbers in the northern fen. Expectedly, many results indicated that microbial response on warming may be linked to the moisture regime. Results indicated that microbial community in the northern fen representing Arctic soils would be more sensitive to environmental changes. The response to future climate change clearly may vary even within a habitat type, exemplified here by boreal sedge fen.