Marcell Experimental Forest Research Articles

Redox Gradient Shapes the Chemical Composition of Peatland Microbial Communities.

The response of soil carbon to climate change and anthropogenic forcing depends on the relationship between the physicochemical variables of the environment and microbial communities. In anoxic soils that store large amounts of organic carbon, it can be hypothesized that the low amount of catabolic energy available leads microbial organisms to minimize the energy costs of biosynthesis, which may shape the composition of microbial communities. To test this hypothesis, thermodynamic modeling was used to assess the link between redox gradients in the ombrotrophic peatland of the Marcell Experimental Forest (Minnesota, USA) and the chemical and taxonomic composition of microbial communities. The average amino acid composition of community-level proteins, called hereafter model proteins, was calculated from shotgun metagenomic sequencing. The carbon oxidation state of model proteins decreases linearly from -0.14 at 10 cm depth to -0.17 at 150 cm depth. Calculating equilibrium activities of model proteins for a wide range of chemical conditions allows identification of the redox potential of maximum chemical activity. Consistent with redox measurements across peat soils, this model Eh decreases logarithmically from an average value of 300 mV at 10 cm depth, close to the stability domain of goethite relative to Fe2+, to an average value of -200 mV at 150 cm, within the stability domain of CH4 relative to CO2. The correlation identified between the taxonomic abundance and the carbon oxidation state of model proteins enables predicting the evolution of taxonomic abundance as a function of model Eh. The model taxonomic abundance is consistent with the measured gene and taxonomic abundance, which evolves from aerobic bacteria at the surface including Acidobacteria, Proteobacteria, and Verrumicrobia, to anaerobes at depth dominated by Crenarchaeota. These results indicate that the thermodynamic forcing imposed by redox gradient across peat soils shapes both the chemical and taxonomic composition of microbial communities. By providing a mechanistic understanding of the relationship between microbial community and environmental conditions, this work sheds new light on the mechanisms that govern soil microbial life and opens up prospects for predicting geochemical and microbial evolution in changing environments.

Parsimonious streamflow forecasting system based on a dynamical systems approach

Operational hydrologic forecasting often relies on the use of physically-based computer models at the catchment scale. However, our understanding of streamflow generation is still evolving and incomplete, and oftentimes linearization assumptions must be made to make physically-based hydrologic and hydraulic models tractable. In addition to uncertainty in the physical runoff-generation mechanisms, many physically-based approaches do not formally assimilate observed streamflow data. Streamflow is their output, deterministically simulated from observed inputs (such as precipitation, temperature, etc.). Divergences between observed and simulated streamflow may be due to input uncertainty, calibration issues, or problems with the inherent physical process representation. Thus, data-driven approaches that are responsive to observed streamflow by including assimilation as an essential feature are promising in operational hydrology. In this study, we use a novel forcings-weighted k-Nearest Neighbor (kNN) nonlinear timeseries forecasting method developed in the context of the modeling of nonlinear dynamical systems. The rainfall-runoff transform is assumed to be a low-dimensional system of (unknown) nonlinear differential equations; the deterministic state space is reconstructed based on the streamflow timeseries; and a forecast is generated based on how “similar” the current hydrograph is to historical events encoded within the reconstructed state space, weighted by the similarity of forcings. We tested the model in a long-term test period with no real data assimilation, and in forecast mode with daily data assimilation. When used for forecasting, we also illustrate two different uncertainty quantification approaches: forecast postprocessing and internal-to-the-model ensemble generation. The method is conceptually and computationally simple, but offers promising results. We tested the model in three different scenarios. First, investigating a long-term test period with no real data assimilation in a rainfall-dominated research catchment at the Coweeta Hydrologic Lab in North Carolina, USA. The simulation exhibited an NSE of 0.73. Next, we included a conceptual snowmelt model to drive the long term simulation and studied boreal-temperate transition zone catchments in northern Minnesota, USA. The study area included a small (<0.1 mi2) reference research catchment at the Marcell Experimental Forest, and a large river basin (>19 k mi2) with incomplete forcings information. The kNN method yielded NSEs from 0.28 to 0.50. And finally, we conducted a pseudo-operational forecasting test for a large river basin in northern Minnesota, USA with a focus on performance forecasting a flood-of-record on the Rainy River. The test involved reforecasting, data assimilation, and implementation of ensemble and postprocessing bias correction uncertainty-quantification schemes. The technique demonstrated good performance capturing a flood-of-record on a large catchment (−5.7 to −3.3 % crest error with 1-day of lead time). For the entire test period in forecast mode, the 1-day lead time NSE was 0.99, going down to 0.79 for the 7-day lead time, showing its general utility as an operational forecasting tool.

Soil frost controls streamflow generation processes in headwater catchments

The relationship between snowmelt and spring streamflow is changing under warming temperatures and diminishing snowpack. At the same time, the hydrologic connectivity across catchment landscape elements, such as snowpack and surface wetlands, can play a critical role in controlling the routing of snowmelt to streams. The role of hydrologic connectivity is important in headwater regions of the continental northern latitudes, where catchments have low topographic relief and seasonally frozen ground. Nevertheless, the effects of soil frost on the sequence, timing, and magnitudes of hydrologic events that drive the movement of water from a snowpack to a stream are not fully understood. Therefore, we examine two questions: First, what is the flowpath that snow melt and precipitation from spring rain events takes to generate spring streamflow, and second, what hydrologic, climatic, or landscape variables exert the most control on the magnitude of streamflow? Here, we use long-term hydrological records from the two reference basins at the Marcell Experimental Forest in northern Minnesota to analyze the cascading effects across precipitation, snow, water table elevation, soil frost, and streamflow in peatland-dominated headwater catchments. We identify a sequence of fill-and-spill effects across the landscape that control the timing of spring streamflow generation. Then, we use stepwise regression to show that soil frost is a key supporting predictor for both the magnitude of streamflow in the spring as it adds significantly to the predictive power of precipitation and water table elevation. Our results highlight the importance of recognizing the role of soil frost, when present, on the partitioning of snowmelt between overland runoff and water table recharge during the critical snowmelt period, as well as the later partitioning between evapotranspiration and subsurface flows.

Effects of Clearcutting and Residual Biomass Harvesting on Hillslope Mercury Mobilization and Downgradient Mercury Accumulation

AbstractMercury (Hg) in forest runoff varies geographically and in relation to silvicultural practices. There is also considerable uncertainty about how forest management practices, such as residual biomass removal, affect Hg mobilization, and downgradient cumulative effects. In this study, total Hg, dissolved organic carbon (DOC), and sulfate (SO42−) mobilization in runoff were compared among unharvested and harvested hillslopes with and without residual biomass removal, and peat soil and invertebrate methylmercury (MeHg) concentrations were assessed in a down‐gradient peatland. Using a before‐after‐control‐impact design, runoff from three adjacent hillslopes was monitored pre‐harvest (2010–2011) and post‐harvest (2012–2013) from snowmelt to freeze‐up at the USDA Forest Service’s Marcell Experimental Forest in northern Minnesota. Dilution, due to increased available hillslope soil water, led to decreased THg and DOC concentrations after harvest at both harvested hillslopes. Sulfate concentrations did not statistically change following harvest. Compared to its removal, leaving residual biomass significantly increased yields of all solutes but only DOC yields significantly increased when the residual biomass was removed. In the adjacent down‐gradient peatland, peat MeHg concentrations decreased, whereas no change was observed in peat below the unharvested hillslope. There was no discernible change post‐harvest in MeHg levels among several peatland macroinvertebrate taxa. This study provides a much‐needed hillslope‐scale understanding of Hg response to forest management practices, highlighting that increased solute yields from hillslopes do not necessarily stimulate Hg methylation or invertebrate bioaccumulation in down‐gradient peatland systems.

The stable isotopes of natural waters at the Marcell Experimental Forest

AbstractWe present a new data set from the Marcell Experimental Forest (MEF) that compiles water isotope measurements from multiple research catchments, some of which have been studied since the 1960s. The MEF is located in northern Minnesota, USA, and is home to heavily studied and monitored forests, streams, bogs, and fens. Peat‐forming systems (bogs and fens) are an important component of the MEF landscape and have a profound impact on the water cycle in these catchments. Within the last decade, analysis of stable isotopes of water (expressed as δD and δ18O) has been implemented to characterize the different components of the water budget, and to allow researchers to look at catchment and peatland‐specific hydrologic effects in the watershed. This δD and δ18O data set of natural waters from MEF catchments is primarily composed of measurements from three peatlands (S1, S2, S6) during an 11‐year period. More recently collection and analysis were expanded to also include samples from the Spruce and Peatland Responses Under Changing Environments (SPRUCE) project in the S1 bog, peatlands S3, S4, S5, as well as nearby lakes. We establish a local meteoric water line by analyzing the isotopic composition of precipitation, which fills a void in regional meteoric water lines for Minnesota. Furthermore, we establish baseline isotopic composition for bog outlet streams, bog porewater, aquifer groundwater, overland flow, subsurface stormflow, and snowpack, as well as runoff from the SPRUCE experimental chambers. These data are publicly available and will be expanded upon in the future.

Analyzing Trends in Water Table Elevations at the Marcell Experimental Forest, Minnesota, U.S.A.

Water table fluctuations in peatlands are closely coupled with the local climate setting and drive critical ecosystem processes such as nutrient cycling. In Minnesota, USA, peatlands cover ten percent of the surface area, approximately 2.5 million hectares, some of which are actively managed for forest products. To explore the relationship between peatland water tables and precipitation, long-term data (1961 to 2019) were used from the Marcell Experimental Forest in northern Minnesota. Starting in 1961, water table data from seven peatlands, including two types of peatlands (bogs and fens), were measured. We used the Theil-Sen estimator to test for monotonic trends in mean monthly water table elevations for individual peatlands and monthly precipitation. Water levels in bogs were both more variable and had mean water table elevations that were closer to the surface. Individual trends of water table elevations differed among peatlands. Water table elevations increased over time in three of the bogs studied and decreased over time in two of the bogs studied. Trends within fens were notably nonlinear across time. No significant linear trend was found for mean monthly precipitation between 1961 and 2019. These results highlight differences in peatlands types, local physiography, and the importance of understanding how changes in long-term dynamics coupled with changing current conditions will influence the effects of water table fluctuations on ecosystem services. The variability of water table elevations in bogs poses potential difficulties in modeling these ecosystems or creating adaptive management plans. KEYWORDS: Peatlands; Hydrology; Water tables; Bogs; Fens; Monitoring; Minnesota; Climate Change

Hydrological and meteorological data from research catchments at the Marcell Experimental Forest, Minnesota, USA

AbstractThe Marcell Experimental Forest (MEF) in northern Minnesota, USA, with hydrological research and monitoring of peatland catchments in a low‐topographic relief landscape, contrasts with the mountainous terrain that typifies most research catchments. Six research catchments were instrumented and hydrological and meteorological monitoring was initiated during 1960. Paired‐catchment studies, which started during 1969, have been used to assess land management and environmental change effects on forests, water availability, and biogeochemistry. Over the decades, the research and collaborations have proliferated to include new monitoring and ecosystem experiments. We provide an overview of available datasets and access information for hydrological and meteorological data. Data on streamflow, water table elevation, precipitation, snow, ground frost, air temperature, soil moisture, upland runoff, and water chemistry are discoverable with associated metadata and are archived through several Web‐based, community repositories. The research programme is ongoing and we anticipate updates on an annual or more frequent basis. Additionally, we aim to release other physical, chemical, and isotopic measurements associated with long‐term catchment monitoring and studies at the MEF.

Nonstationary flood-frequency analysis to assess effects of harvest and cover type conversion on peak flows at the Marcell Experimental Forest, Minnesota, USA

Forest harvest, climate change, and their interaction can alter catchment peak discharges, which have ramifications for channel geomorphology and water quality. Catchments in the boreal-temperate transition zone may be especially vulnerable to these factors. We developed a new approach to peak flow analysis using two long-term, paired catchment experiments in that landscape at the Marcell Experimental Forest (MEF) in north-central Minnesota. We investigated how forest cover change affects the magnitude, occurrence probability, event decoupling, and seasonal decoupling of annual maximum peak flows. Event decoupling is when the annual maximum flows on control and treatment catchments occur in response to different events within a given year. Seasonal decoupling is when the annual maximum flows on control and treatment catchments occur in different seasons within a given year. Commonly used statistical methods (e.g., ANCOVA) support inferences only for effects on peak flow magnitude, disregarding effects on event occurrence probability. Statistical stationarity (i.e., time-invariant parameters) is generally assumed within the ANCOVA analysis, despite temporal changes in climatic and land cover controls. Further, use of these methods requires event coupling, which is not always valid for annual maximum flows in catchments with mixed precipitation regimes. To address these limitations, we developed new nonstationary flood-frequency analysis methods with Bayesian parameter estimation to analyze two harvesting experiments, including conversion from deciduous to coniferous species. We compared results from these models to traditional least-squares based methods (e.g., ANCOVA). We assessed event and seasonal decoupling using logistic regression with Bayesian parameter estimation. There was no effect of harvesting on the annual maximum flow according to ANCOVA and generalized least squares models, but these methods were not reliable due to the presence of outliers, and both event and seasonal decoupling. Flood-frequency results from one clearcutting experiment showed increases in the annual maximum flow across nearly all return intervals, with 80–85% confidence for peak discharges between the 10 and 50-year peak flow. The 50-year return interval peak increased from an expected 17 cfs to 34 cfs after harvest. Harvesting induced substantial event decoupling in the catchment pairs, and seasonal decoupling in one catchment pair. As forest cover regenerated, decoupling probability decreased. Our coupling analysis results indicated that control and treatment catchments generated and processed flood peaks differently after harvest, and explained the poor fit of the linear models for which we assumed event coupling. Our study calls into question several key assumptions of traditional linear models in the analysis of paired-catchments given certain conditions. For example, where forest harvesting can affect the season of annual maximum peaks, and thus the phase of the generating precipitation event (snowmelt versus rainfall), it changes the relationship between the catchment pair in a way not accounted for in traditional linear models. We advocate the use of probabilistic methods that can incorporate nonstationarity and are robust to different mechanisms of catchment hydrologic change in response to forest harvest.

Sources and biodegradability of dissolved organic matter in two headwater peatland catchments at the Marcell Experimental Forest, northern Minnesota,USA

AbstractWhere they are present in catchments, peatlands are a dominant source of dissolved organic matter (DOM) to surrounding waterways due, in part, to high production rates. Despite the preponderance of peatlands in northern latitudes and expected peatland vulnerability to climate change, little is known about peatland DOM degradation relative to a more comprehensive understanding of degradation when DOM is sourced from upland‐dominated catchments. We compared DOM biodegradability of various sources of stream water in two catchments having peatlands (22%–33% of the area) surrounded by upland forests (70%–90% of the area, either deciduous or coniferous). We measured total organic carbon (TOC), and biodegradable dissolved organic carbon concentrations; bacterial respiration rates; streamflow; and upland runoff during and after snowmelt (March to June, 2009–2011). We also explored if DOM in upland runoff stimulated biodegradation of peatland‐derived DOM (i.e., a priming effect), and if forest cover type affected DOM biodegradability. As expected, the peatlands were the largest sources of both water (72%–80%) and TOC (92%–96%) to the streams although more area in each catchment was in uplands (70%–90%). Several results were unexpected, yet revealing: (1) DOM from peatlands sometimes had the same biodegradability as DOM from uplands, (2) upland sources of DOM had negligible effects on biodegradability in the peatland and downstream, and (3) upland deciduous cover did not yield more degradable DOM than conifer cover. The most pronounced effect of upland runoff was dilution of downstream TOC concentrations when there was upland runoff. Overall, the effects of upland DOM may have been negligible due to the overriding effect of the large amount of biodegradable DOM that originated in bogs. This research highlights that peatland‐sourced DOM has important effects on downstream DOM biodegradability even in catchments where upland area is substantially larger than peatland area.

Changes in hillslope hydrology in a perched, shallow soil system due to clearcutting and residual biomass removal

AbstractSustainable fuels legislation and volatility in energy prices have put additional pressures on the forestry sector to intensify the harvesting of biomass for “advanced biofuel” production. To better understand how residual biomass removal after harvest affects forest hydrology in relatively low slope terrain, a Before‐After‐Control‐Impact (BACI) study was conducted in the USDA Forest Service's Marcell Experimental Forest, Minnesota, USA. Hydrological measurements were made from 2010–2013 on a forested hillslope that was divided into three treatment blocks, where one block was harvested and residual biomass removed (Biomass Removed), the second was harvested and residual biomass left (Biomass Left), and the last block was left as an Unharvested Control. The pre‐harvest period (2 years) was 2010–11 and post‐harvest (2 years) was 2012–13. Water table elevation at the upslope and downslope position, subsurface runoff, and soil moisture were measured between May–November. Mixed effect statistical models were used to compare both the before‐after and “control” treatment ratios (ratios between harvested hillslopes and the Unharvested Control hillslope). Subsurface runoff significantly increased (p &lt; .05) at both harvested hillslopes but to a greater degree on the Biomass Left hillslope. Greater subsurface runoff volumes at both harvested hillslopes were driven by substantial increases during fall, with additional significant increases during summer on the Biomass Left hillslope. The hydrological connectivity, inferred from event runoff ratios, increased due to harvesting at both hillslopes but only significantly on the Biomass Left hillslope. The winter harvest minimized soil disturbance, resulting in no change to the effective hydraulic conductivity distribution with depth. Thus, the observed hydrological changes were driven by increased effective precipitation and decreased evapotranspiration, increasing the duration that both harvested hillslopes were hydrologically active. The harvesting of residual biomass appears to lessen hydrological connectivity relative to leaving residual biomass on the hillslope, potentially decreasing downstream hydrological impacts of similar forestry operations.

Climatic controls on peatland black spruce growth in relation to water table variation and precipitation

AbstractPrior research has demonstrated the importance of water limitations and increasing temperatures on upland black spruce (Picea mariana [Mill.] B.S.P.) tree growth, which is a dominant component of the North American boreal forest. However, little work has been done to investigate the connectivity between growth and hydro‐climate in peatland black spruce systems. The boreal forest is the largest global terrestrial biome and is highly threatened due to current and projected increases in temperatures for the northern latitudes. Here we explore the dynamics among annual black spruce growth, climate, and water table elevations using 45 years of in situ precipitation, temperature, and water table elevation coupled with dendrochronological analysis from six research peatlands at the Marcell Experimental Forest, MN, USA. From 1963 to 2010, we found weak relationships between water table elevation and black spruce growth at the six study sites. Instead, annual black spruce growth was most favourable during three climatic periods: (a) cool, moist conditions in mid‐summer; (b) warm mid‐spring temperatures; and (c) cool temperatures in the fall prior to the current growing season. The disconnect between black spruce growth and water table dynamics was surprising and suggests that either annual black spruce growth is minimally responsive to hydrological fluctuations at the timescale we analysed or there is great elasticity of black spruce growth to peatland water table and evapotranspiration dynamics under the range of hydrological fluctuations contained in our record.

High Genetic Potential for Proteolytic Decomposition in Northern Peatland Ecosystems.

Nitrogen (N) is a scarce nutrient commonly limiting primary productivity. Microbial decomposition of complex carbon (C) into small organic molecules (e.g., free amino acids) has been suggested to supplement biologically fixed N in northern peatlands. We evaluated the microbial (fungal, bacterial, and archaeal) genetic potential for organic N depolymerization in peatlands at Marcell Experimental Forest (MEF) in northern Minnesota. We used guided gene assembly to examine the abundance and diversity of protease genes and further compared them to those of N fixation (nifH) genes in shotgun metagenomic data collected across depths and in two distinct peatland environments (bogs and fens). Microbial protease genes greatly outnumbered nifH genes, with the most abundant genes (archaeal M1 and bacterial trypsin [S01]) each containing more sequences than all sequences attributed to nifH Bacterial protease gene assemblies were diverse and abundant across depth profiles, indicating a role for bacteria in releasing free amino acids from peptides through depolymerization of older organic material and contrasting with the paradigm of fungal dominance in depolymerization in forest soils. Although protease gene assemblies for fungi were much less abundant overall than those for bacteria, fungi were prevalent in surface samples and therefore may be vital in degrading large soil polymers from fresh plant inputs during the early stage of depolymerization. In total, we demonstrate that depolymerization enzymes from a diverse suite of microorganisms, including understudied bacterial and archaeal lineages, are prevalent within northern peatlands and likely to influence C and N cycling.IMPORTANCE Nitrogen (N) is a common limitation on primary productivity, and its source remains unresolved in northern peatlands that are vulnerable to environmental change. Decomposition of complex organic matter into free amino acids has been proposed as an important N source, but the genetic potential of microorganisms mediating this process has not been examined. Such information can inform possible responses of northern peatlands to environmental change. We show high genetic potential for microbial production of free amino acids across a range of microbial guilds in northern peatlands. In particular, the abundance and diversity of bacterial genes encoding proteolytic activity suggest a predominant role for bacteria in regulating productivity and contrasts with a paradigm of fungal dominance of organic N decomposition. Our results expand our current understanding of coupled carbon and nitrogen cycles in northern peatlands and indicate that understudied bacterial and archaeal lineages may be central in this ecosystem's response to environmental change.

Temperature sensitivity of extracellular enzymes differs with peat depth but not with season in an ombrotrophic bog

Peatlands contain a large portion of Earth's terrestrial soil organic matter in part due to a reduction in decomposition rates. Organic matter decomposition is initially mediated by extracellular enzyme activity, which is in turn controlled by temperature, moisture, and substrate availability; and all are subject to seasonal variation. As depth increases in peatlands, temperature variability and labile carbon inputs decrease. We hypothesized that the more stable recalcitrant subsurface would contain a smaller less diverse enzyme pool, that is better adapted to a narrow temperature range. Thus temperature dependence would be diminished at depth compared to superficial peat. Potential enzyme activity rates were determined across seasons and with depth in peat samples collected from the Marcell Experimental Forest in northern Minnesota, USA. The temperature dependence, assessed by activation energy, was quantified for three hydrolytic enzymes involved in nutrient cycling at up to 15 temperature points ranging from 2 °C to 65 °C. Potential enzyme activity decreased with peat depth as expected and corresponded with changes in peat composition and microbial biomass from the acrotelm to the catotelm. In an environmentally relevant temperature range (2–23 °C), activation energy decreased with depth for β-glucosidase as predicted and leucine amino peptidase activation energy was the lowest of all enzymes. Stable temperatures at depth appear to result in a microbial community containing enzymes that have lower sensitivity to temperature increases. Surprisingly, there was no significant seasonal effect on enzyme temperature dependence observed in our study. Based on these results, and without shifts in microbial community composition, warming of peat could result in increased carbon and phosphorus cycling at the surface but little change at depth. Additionally differences in enzyme temperature sensitivity suggest nitrogen cycling could remain constant with warming, potentially resulting in proteolytic nitrogen cycling being decoupled from carbon and phosphorus cycling.

Topographic, edaphic, and vegetative controls on plant‐available water

AbstractSoil moisture varies within landscapes in response to vegetative, physiographic, and climatic drivers, which makes quantifying soil moisture over time and space difficult. Nevertheless, understanding soil moisture dynamics for different ecosystems is critical, as the amount of water in a soil determines a myriad ecosystem services and processes such as net primary productivity, runoff, microbial decomposition, and soil fertility. We investigated the patterns and variability in in situ soil moisture measurements converted to plant‐available water across time and space under different vegetative cover types and topographic positions at the Marcell Experimental Forest (Minnesota, USA). From 0‐ to 228.6‐cm soil depth, plant‐available water was significantly higher under the hardwoods (12%), followed by the aspen (8%) and red pine (5%) cover types. Across the same soil depth, toeslopes were wetter (mean plant‐available water = 10%) than ridges and backslopes (mean plant‐available water was 8%), although these differences were not statistically significant (p &lt; .05). Using a mixed model of fixed and random effects, we found that cover type, soil texture, and time were related to plant‐available water and that topography was not significantly related to plant‐available water within this low‐relief landscape. Additionally, during the 3‐year monitoring period, red pine and quaking aspen sites experienced plant‐available water levels that may be considered limiting to plant growth and function. Given that increasing temperatures and more erratic precipitation patterns associated with climate change may result in decreased soil moisture in this region, these species may be sensitive and vulnerable to future shifts in climate.

Molybdenum-Based Diazotrophy in a Sphagnum Peatland in Northern Minnesota.

Microbial N2 fixation (diazotrophy) represents an important nitrogen source to oligotrophic peatland ecosystems, which are important sinks for atmospheric CO2 and are susceptible to the changing climate. The objectives of this study were (i) to determine the active microbial group and type of nitrogenase mediating diazotrophy in an ombrotrophic Sphagnum-dominated peat bog (the S1 peat bog, Marcell Experimental Forest, Minnesota, USA); and (ii) to determine the effect of environmental parameters (light, O2, CO2, and CH4) on potential rates of diazotrophy measured by acetylene (C2H2) reduction and 15N2 incorporation. A molecular analysis of metabolically active microbial communities suggested that diazotrophy in surface peat was primarily mediated by Alphaproteobacteria (Bradyrhizobiaceae and Beijerinckiaceae). Despite higher concentrations of dissolved vanadium ([V] 11 nM) than molybdenum ([Mo] 3 nM) in surface peat, a combination of metagenomic, amplicon sequencing, and activity measurements indicated that Mo-containing nitrogenases dominate over the V-containing form. Acetylene reduction was only detected in surface peat exposed to light, with the highest rates observed in peat collected from hollows with the highest water contents. Incorporation of 15N2 was suppressed 90% by O2 and 55% by C2H2 and was unaffected by CH4 and CO2 amendments. These results suggest that peatland diazotrophy is mediated by a combination of C2H2-sensitive and C2H2-insensitive microbes that are more active at low concentrations of O2 and show similar activity at high and low concentrations of CH4 IMPORTANCE Previous studies indicate that diazotrophy provides an important nitrogen source and is linked to methanotrophy in Sphagnum-dominated peatlands. However, the environmental controls and enzymatic pathways of peatland diazotrophy, as well as the metabolically active microbial populations that catalyze this process, remain in question. Our findings indicate that oxygen levels and photosynthetic activity override low nutrient availability in limiting diazotrophy and that members of the Alphaproteobacteria (Rhizobiales) catalyze this process at the bog surface using the molybdenum-based form of the nitrogenase enzyme.

Alpha- and Gammaproteobacterial Methanotrophs Codominate the Active Methane-Oxidizing Communities in an Acidic Boreal Peat Bog.

The objective of this study was to characterize metabolically active, aerobic methanotrophs in an ombrotrophic peatland in the Marcell Experimental Forest, in Minnesota. Methanotrophs were investigated in the field and in laboratory incubations using DNA-stable isotope probing (SIP), expression studies on particulate methane monooxygenase (pmoA) genes, and amplicon sequencing of 16S rRNA genes. Potential rates of oxidation ranged from 14 to 17 μmol of CH4g dry weight soil(-1)day(-1) Within DNA-SIP incubations, the relative abundance of methanotrophs increased from 4% in situ to 25 to 36% after 8 to 14 days. Phylogenetic analysis of the(13)C-enriched DNA fractions revealed that the active methanotrophs were dominated by the genera Methylocystis(type II;Alphaproteobacteria),Methylomonas, and Methylovulum(both, type I;Gammaproteobacteria). In field samples, a transcript-to-gene ratio of 1 to 2 was observed for pmoA in surface peat layers, which attenuated rapidly with depth, indicating that the highest methane consumption was associated with a depth of 0 to 10 cm. Metagenomes and sequencing of cDNA pmoA amplicons from field samples confirmed that the dominant active methanotrophs were Methylocystis and Methylomonas Although type II methanotrophs have long been shown to mediate methane consumption in peatlands, our results indicate that members of the genera Methylomonas and Methylovulum(type I) can significantly contribute to aerobic methane oxidation in these ecosystems.

Growth–climate relationships across topographic gradients in the northern Great Lakes

AbstractClimatic conditions exert important control over the growth, productivity, and distribution of forests, and characterizing these relationships is essential for understanding how forest ecosystems will respond to climate change. We used dendrochronological methods to develop climate–growth relationships for two dominant species, Populus tremuloides (quaking aspen) and Pinus resinosa (red pine), in the upper Great Lakes region to understand how climate and water availability influence annual forest productivity. Trees were sampled along a topographic gradient at the Marcell Experimental Forest (Minnesota, USA) to assess growth response to variations in temperature and different water availability metrics (precipitation, potential evapotranspiration (PET), cumulative moisture index (CMI), and soil water storage). Climatic variables were able to explain 33–58% of the variation in annual growth (as measured by ring‐width increment) for quaking aspen and 37–74% of the variation for red pine. Climate–growth relationships were influenced by topography for quaking aspen but not for red pine. Annual ring growth for quaking aspen decreased with June CMI on ridges, decreased with temperature in the November prior to the growing season on sideslopes, and decreased with June PET on toeslopes. Red pine growth increased with increasing July PET across all topographic positions. These results indicate the sensitivity of both quaking aspen and red pine to local climate and show several vulnerabilities of these species to shifts in water supply and temperature because of climate change. Copyright © 2015 John Wiley &amp; Sons, Ltd.

Ecoenzymatic stoichiometry and microbial processing of organic matter in northern bogs and fens reveals a common P-limitation between peatland types

We compared carbon (C), nitrogen (N), and phosphorus (P) concentrations in atmospheric deposition, runoff, and soils with microbial respiration [dehydrogenase (DHA)] and ecoenzyme activity (EEA) in an ombrotrophic bog and a minerotrophic fen to investigate the environmental drivers of biogeochemical cycling in peatlands at the Marcell Experimental Forest in northern Minnesota, USA. Ecoenzymatic stoichiometry was used to construct models for C use efficiency (CUE) and decomposition (M), and these were used to model respiration (Rm). Our goals were to determine the relative C, N, and P limitations on microbial processes and organic matter decomposition, and to identify environmental constraints on ecoenzymatic processes. Mean annual water, C, and P yields were greater in the fen, while N yields were similar in both the bog and fen. Soil chemistry differed between the bog and fen, and both watersheds exhibited significant differences among soil horizons. DHA and EEA differed by watersheds and soil horizons, CUE, M, and Rm differed only by soil horizons. C, N, or P limitations indicated by EEA stoichiometry were confirmed with orthogonal regressions of ecoenzyme pairs and enzyme vector analyses, and indicated greater N and P limitation in the bog than in the fen, with an overall tendency toward P-limitation in both the bog and fen. Ecoenzymatic stoichiometry, microbial respiration, and organic matter decomposition were responsive to resource availability and the environmental drivers of microbial metabolism, including those related to global climate changes.

Organic matter transformation in the peat column at Marcell Experimental Forest: Humification and vertical stratification

We characterized peat decomposition at the Marcell Experimental Forest (MEF), Minnesota, USA, to a depth of 2 m to ascertain the underlying chemical changes using Fourier transform infrared (FT IR) and 13C nuclear magnetic resonance (NMR) spectroscopy) and related these changes to decomposition proxies C:N ratio, δ13C and δ15N, bulk density, and water content. FT IR determined that peat humification increased rapidly between 30 and 75 cm, indicating a highly reactive intermediate-depth zone consistent with changes in C:N ratio, δ13C and δ15N, bulk density, and water content. Peat decomposition at the MEF, especially in the intermediate-depth zone, is mainly characterized by preferential utilization of O-alkyl-C, carboxyl-C, and other oxygenated functionalities with a concomitant increase in the abundance of alkyl- and nitrogen-containing compounds. Below 75 cm, less change was observed but aromatic functionalities and lignin accumulated with depth. Significant correlations with humification indices, identified by FT IR spectroscopy, were found for C:N ratios. Incubation studies at 22°C revealed the highest methane production rates, greatest CH4:CO2 production ratios, and significant O-alkyl-C utilization within this 30 and 75 cm zone. Oxygen-containing functionalities, especially O-alkyl-C, appear to serve as excellent proxies for soil decomposition rate and should be a sensitive indicator of the response of the solid phase peat to increased temperatures caused by climate change and the field study manipulations that are planned to occur at this site. Radiocarbon signatures of microbial respiration products in deeper pore waters at the MEF resembled the signatures of more modern dissolved organic carbon rather than solid phase peat, indicating that recently photosynthesized organic matter fueled the bulk of subsurface microbial respiration. These results indicate that carbon cycling at depth at the MEF is not isolated from surface processes.

Microbial Community Stratification Linked to Utilization of Carbohydrates and Phosphorus Limitation in a Boreal Peatland at Marcell Experimental Forest, Minnesota, USA

This study investigated the abundance, distribution, and composition of microbial communities at the watershed scale in a boreal peatland within the Marcell Experimental Forest (MEF), Minnesota, USA. Through a close coupling of next-generation sequencing, biogeochemistry, and advanced analytical chemistry, a biogeochemical hot spot was revealed in the mesotelm (30- to 50-cm depth) as a pronounced shift in microbial community composition in parallel with elevated peat decomposition. The relative abundance of Acidobacteria and the Syntrophobacteraceae, including known hydrocarbon-utilizing genera, was positively correlated with carbohydrate and organic acid content, showing a maximum in the mesotelm. The abundance of Archaea (primarily crenarchaeal groups 1.1c and 1.3) increased with depth, reaching up to 60% of total small-subunit (SSU) rRNA gene sequences in the deep peat below the 75-cm depth. Stable isotope geochemistry and potential rates of methane production paralleled vertical changes in methanogen community composition to indicate a predominance of acetoclastic methanogenesis mediated by the Methanosarcinales in the mesotelm, while hydrogen-utilizing methanogens predominated in the deeper catotelm. RNA-derived pyrosequence libraries corroborated DNA sequence data to indicate that the above-mentioned microbial groups are metabolically active in the mid-depth zone. Fungi showed a maximum in rRNA gene abundance above the 30-cm depth, which comprised only an average of 0.1% of total bacterial and archaeal rRNA gene abundance, indicating prokaryotic dominance. Ratios of C to P enzyme activities approached 0.5 at the acrotelm and catotelm, indicating phosphorus limitation. In contrast, P limitation pressure appeared to be relieved in the mesotelm, likely due to P solubilization by microbial production of organic acids and C-P lyases. Based on path analysis and the modeling of community spatial turnover, we hypothesize that P limitation outweighs N limitation at MEF, and microbial communities are structured by the dominant shrub, Chamaedaphne calyculata, which may act as a carbon source for major consumers in the peatland.