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.

Monitoring of carbon-water fluxes at Eurasian meteorological stations using random forest and remote sensing.

Simulating the carbon-water fluxes at more widely distributed meteorological stations based on the sparsely and unevenly distributed eddy covariance flux stations is needed to accurately understand the carbon-water cycle of terrestrial ecosystems. We established a new framework consisting of machine learning, determination coefficient (R2), Euclidean distance, and remote sensing (RS), to simulate the daily net ecosystem carbon dioxide exchange (NEE) and water flux (WF) of the Eurasian meteorological stations using a random forest model or/and RS. The daily NEE and WF datasets with RS-based information (NEE-RS and WF-RS) for 3774 and 4427 meteorological stations during 2002-2020 were produced, respectively. And the daily NEE and WF datasets without RS-based information (NEE-WRS and WF-WRS) for 4667 and 6763 meteorological stations during 1983-2018 were generated, respectively. For each meteorological station, the carbon-water fluxes meet accuracy requirements and have quasi-observational properties. These four carbon-water flux datasets have great potential to improve the assessments of the ecosystem carbon-water dynamics.

Water level drawdown makes boreal peatland vegetation more responsive to weather conditions.

Climate warming and projected increase in summer droughts puts northern peatlands under pressure by subjecting them to a combination of gradual drying and extreme weather events. The combined effect of those on peatland functions is poorly known. Here, we studied the impact of long-term water level drawdown (WLD) and contrasting weather conditions on leaf phenology and biomass production of ground level vegetation in boreal peatlands. Data were collected during two contrasting growing seasons from a WLD experiment including a rich and a poor fen and an ombrotrophic bog. Results showed that WLD had a strong effect on both leaf area development and biomass production, and these responses differed between peatland types. In the poor fen and the bog, WLD increased plant growth, while in the rich fen, WLD reduced the growth of ground level vegetation. Plant groups differed in their response, as WLD reduced the growth of graminoids, while shrubs and tree seedlings benefited from it. In addition, the vegetation adjusted to the lower WTs, was more responsive to short-term climatic variations. The warmer summer resulted in a greater maximum and earlier peaking of leaf area index, and greater biomass production by vascular plants and Sphagnum mosses at WLD sites. In particular, graminoids benefitted from the warmer conditions. The change towards greater production in the WLD sites in general and during the warmer weather in particular, was related to the observed transition in plant functional type composition towards arboreal vegetation.

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.

Emissions of biogenic volatile organic compounds from adjacent boreal fen and bog as impacted by vegetation composition.

Peatland ecosystems emit biogenic volatile organic compounds (BVOC), which have a net cooling impact on the climate. However, the quality and quantity of BVOC emissions, and how they are regulated by vegetation and peatland type remain poorly understood. Here we measured BVOC emissions with dynamic enclosures from two major boreal peatland types, a minerotrophic fen and an ombrotrophic bog situated in Siikaneva, southern Finland and experimentally assessed the role of vegetation by removing vascular vegetation with or without the moss layer. Our measurements from four campaigns during growing seasons in 2017 and 2018 identified emissions of 59 compounds from nine different chemical groups. Isoprene accounted for 81 % of BVOC emissions. Measurements also revealed uptake of dichloromethane. Total BVOC emissions and the emissions of isoprene, monoterpenoids, sesquiterpenes, homoterpenes, and green leaf volatiles were tightly connected to vascular plants. Isoprene and sesquiterpene emissions were associated with sedges, whereas monoterpenoids and homoterpenes were associated with shrubs. Additionally, isoprene and alkane emissions were higher in the fen than in the bog and they significantly contributed to the higher BVOC emissions from intact vegetation in the fen. During an extreme drought event in 2018, emissions of organic halides were absent. Our results indicate that climate change with an increase in shrub cover and increased frequency of extreme weather events may have a negative impact on total BVOC emissions that otherwise are predicted to increase in warmer temperatures. However, these changes also accompanied a change in BVOC emission quality. As different compounds differ in their capacity to form secondary organic aerosols, the ultimate climate impact of peatland BVOC emissions may be altered.

Plant functional traits play the second fiddle to plant functional types in explaining peatland CO2 and CH4 gas exchange.

Peatlands constitute a significant soil carbon (C) store, yet the C gas flux components show distinct spatial variation both between and within peatlands. Determining the controls on this variability could aid in our understanding of the response of peatlands to global changes. In this study, we assess the usefulness of different vegetation related parameters to explain spatial variation in peatland C gas flux components. We hypothesise that spatial variation is best explained by trait-based indices (similarly to other terrestrial ecosystems), and that the impact of soil physicochemical properties, such as nitrogen (N) content or water level, can be manifested through the traits. Furthermore, we expect that the spatial variability associated with each of the C gas flux components can be explained by a distinct set of traits. To address our aim, we used a successional peatland chronosequence from wet meadows to a bog, along which all variables were recorded with similar methods and under similar climatic conditions. We observed spatial variability with all measured gas fluxes, with carbon dioxide (CO2) fluxes showing significant variability between sites, while within site variability was more important for methane (CH4) fluxes. As expected, our results show that the impacts of physicochemical conditions were directed via vegetation. However, the cover of functional plant types that capture multiple traits proved to be more powerful in explaining gas flux variability compared to functional trait-based indices. Our findings imply that for future gas flux modelling purposes, rather than attempting to use individual traits - as is the ongoing trend in ecology - it might be more useful to refine plant functional groupings and ensure they are based on a set of plant traits relevant for the studied ecosystem process. This could be facilitated by the collation of a large data set of traits measured from peatlands.

Methane production and oxidation potentials along a fen-bog gradient from southern boreal to subarctic peatlands in Finland.

Methane (CH4 ) emissions from northern peatlands are projected to increase due to climate change, primarily because of projected increases in soil temperature. Yet, the rates and temperature responses of the two CH4 emission-related microbial processes (CH4 production by methanogens and oxidation by methanotrophs) are poorly known. Further, peatland sites within a fen-bog gradient are known to differ in the variables that regulate these two mechanisms, yet the interaction between peatland type and temperature lacks quantitative understanding. Here, we investigated potential CH4 production and oxidation rates for 14 peatlands in Finland located between c. 60 and 70°N latitude, representing bogs, poor fens, and rich fens. Potentials were measured at three different temperatures (5, 17.5, and 30℃) using the laboratory incubation method. We linked CH4 production and oxidation patterns to their methanogen and methanotroph abundance, peat properties, and plant functional types. We found that the rich fen-bog gradient-related nutrient availability and methanogen abundance increased the temperature response of CH4 production, with rich fens exhibiting the greatest production potentials. Oxidation potential showed a steeper temperature response than production, which was explained by aerenchymous plant cover, peat water holding capacity, peat nitrogen, and sulfate content. The steeper temperature response of oxidation suggests that, at higher temperatures, CH4 oxidation might balance increased CH4 production. Predicting net CH4  fluxes as an outcome of the two mechanisms is complicated due to their different controls and temperature responses. The lack of correlation between field CH4  fluxes and production/oxidation potentials, and the positive correlation with aerenchymous plants points toward the essential role of CH4 transport for emissions. The scenario of drying peatlands under climate change, which is likely to promote Sphagnum establishment over brown mosses in many places, will potentially reduce the predicted warming-related increase in CH4 emissions by shifting rich fens to Sphagnum-dominated systems.

Refining the role of phenology in regulating gross ecosystem productivity across European peatlands.

The role of plant phenology as a regulator for gross ecosystem productivity (GEP) in peatlands is empirically not well constrained. This is because proxies to track vegetation development with daily coverage at the ecosystem scale have only recently become available and the lack of such data has hampered the disentangling of biotic and abiotic effects. This study aimed at unraveling the mechanisms that regulate the seasonal variation in GEP across a network of eight European peatlands. Therefore, we described phenology with canopy greenness derived from digital repeat photography and disentangled the effects of radiation, temperature and phenology on GEP with commonality analysis and structural equation modeling. The resulting relational network could not only delineate direct effects but also accounted for possible effect combinations such as interdependencies (mediation) and interactions (moderation). We found that peatland GEP was controlled by the same mechanisms across all sites: phenology constituted a key predictor for the seasonal variation in GEP and further acted as a distinct mediator for temperature and radiation effects on GEP. In particular, the effect of air temperature on GEP was fully mediated through phenology, implying that direct temperature effects representing the thermoregulation of photosynthesis were negligible. The tight coupling between temperature, phenology and GEP applied especially to high latitude and high altitude peatlands and during phenological transition phases. Our study highlights the importance of phenological effects when evaluating the future response of peatland GEP to climate change. Climate change will affect peatland GEP especially through changing temperature patterns during plant phenologically sensitive phases in high latitude and high altitude regions.

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.