Biological nitrogen fixation, diversity and community structure of diazotrophs in two mosses in 25 temperate forests.

Many moss species are associated with nitrogen (N)-fixing bacteria (diazotrophs) that support the N supply of mosses. Our knowledge relates primarily to pristine ecosystems with low atmospheric N input, but knowledge of biological N fixation (BNF) and diazotrophic communities in mosses in temperate forests with high N deposition is limited. We measured BNF rates using the direct stable isotope method and studied the total and potentially active diazotrophic communities in two abundant mosses, Brachythecium rutabulum and Hypnum cupressiforme, both growing on lying deadwood trunks in 25 temperate forest sites. BNF rates in both mosses were similar to those observed in moss species of pristine ecosystems. H. cupressiforme fixed three times more N2 and exhibited lower diazotrophic richness than B. rutabulum. Frankia was the most prominent diazotroph followed by cyanobacteria Nostoc. Manganese, iron, and molybdenum contents in mosses were positively correlated with BNF and diazotrophic communities. Frankia maintained high BNF rates in H. cupressiforme and B. rutabulum even under high chronic N deposition in Central European forests. Moss N concentration and 15 N abundance indicate a rather minor contribution of BNF to the N nutrition of these mosses.

Climate-induced hydrological fluctuations shape Arctic Alaskan peatland plant communities.

Rapidly increasing temperatures in high-latitude regions are causing major changes in wetland ecosystems. To assess the impact of concomitant hydroclimatic fluctuations, mineral deposition, and autogenous succession on the rate and direction of changing arctic plant communities in Arctic Alaska, we conducted detailed palaeoecological analyses using plant macrofossil, pollen, testate amoebae, elemental analyses, and radiocarbon and lead (210Pb) dating on two replicate monoliths from a peatland that developed in a river valley on the northern foothills of the Books Range. We observed an expansion of Sphagnum populations and vascular plants preferring dry habitats, such as Sphagnum warnstorfii, Sphagnum teres/squarrosum, Polytrichum strictum, Aulacomnium palustre and Salix sp., in recent decades between 2000 and 2015 CE, triggered by an increase in temperature and deepening water tables. Deepening peatland water tables became accentuated over the last two decades, when it reached its lowest point in the last 700 years. Conversely, a higher water-table between ca. 1500 and 1950 CE led to a recession of Sphagnum communities and an expansion of sedges. The almost continuous supply of mineral matter during this time led to a dominance of minerotrophic plant communities, although with varying species composition throughout the study period. The replicate cores show similar patterns, but nuanced differences are also visible, depicting fine spatial scale differences particularly in peat-forming plant distribution and the different timings of their presence. In conclusion, our study provides valuable insights into the impact of hydroclimatic fluctuations on peatland vegetation in Arctic Alaska, highlighting their tendency to dry out in recent decades. It also highlights the importance of river valley peatlands in paleoenvironmental reconstructions.

Permafrost thaw causes large carbon loss in boreal peatlands while changes to peat quality are limited.

Rapid, ongoing permafrost thaw of peatlands in the discontinuous permafrost zone is exposing a globally significant store of soil carbon (C) to microbial processes. Mineralization and release of this peat C to the atmosphere as greenhouse gases is a potentially important feedback to climate change. Here we investigated the effects of permafrost thaw on peat C at a peatland complex in western Canada. We collected 15 complete peat cores (between 2.7 and 4.5 m deep) along four chronosequences, from elevated permafrost peat plateaus to saturated thermokarst bogs that thawed up to 600 years ago. The peat cores were analysed for peat C storage and peat quality, as indicated by decomposition proxies (FTIR and C/N ratios) and potential decomposability using a 200-day aerobic laboratory incubation. Our results suggest net C loss following thaw, with average total peat C stocks decreasing by ~19.3 ± 7.2 kg C m-2 over <600 years (~13% loss). Average post-thaw accumulation of new peat at the surface over the same period was ~13.1 ± 2.5 kg C m-2 . We estimate ~19% (±5.8%) of deep peat (>40 cm below surface) C is lost following thaw (average 26 ± 7.9 kg C m-2 over <600 years). Our FTIR analysis shows peat below the thaw transition in thermokarst bogs is slightly more decomposed than peat of a similar type and age in permafrost plateaus, but we found no significant changes to the quality or lability of deeper peat across the chronosequences. Our incubation results also showed no increase in C mineralization of deep peat across the chronosequences. While these limited changes in peat quality in deeper peat following permafrost thaw highlight uncertainty in the exact mechanisms and processes for C loss, our analysis of peat C stocks shows large C losses following permafrost thaw in peatlands in western Canada.

Temperature and organic carbon quality control the anaerobic carbon mineralization in peat profiles via modulating microbes: A case study of Changbai Mountain.

Peatlands account for a significant fraction of the global carbon stock. However, the complex interplay of abiotic and biotic factors governing anaerobic carbon mineralization in response to warming remains unclear. In this study, peat sediments were collected from a typical northern peatland-Changbai Mountain to investigate the behavior and mechanism of anaerobic carbon mineralization in response to depth (0-200 cm) and temperature (5 °C, 15 °C and 20 °C), by integrating geochemical and microbial analysis. Several indices including humification indexes (HI), aromaticity, and water extractable organic carbon (WEOC) components were applied to evaluate carbon quality, while 16S rRNA sequencing was used to measure microbial composition. Regardless of temperature, degradations of carbon quality and associated reduction in microbial abundance as well as diversity resulted in a decrease in anaerobic carbon mineralization (both CO2 and CH4) towards greater depth. Warming either from 5 °C to 15 °C or 20 °C significantly increased anaerobic carbon mineralization in all depth profiles by improving carbon availability. Enhanced carbon availabilities were mediated by the change in microbial composition (p < 0.01) and an increase in metabolic activities, which was particularly evident in the enhanced ß-glucosidase activity and microbial collaborations. A remarkable increase of over 10-fold in the relative abundance of the Geothrix genus was observed under warming. Overall, warming resulted in an enhanced contribution of CH4 emission and a higher ratio of hydrogenotrophic methanogenesis, as evidenced by carbon isotope fractionation factors. In addition, deep peat soils (>100 cm) with recalcitrant carbon demonstrated greater temperature sensitivity (Q10: ∼2.0) than shallow peat soils (Q10:∼1.2) when temperature increased from 15 °C to 20 °C. The findings of this study have significantly deepened our understanding for mechanisms of carbon quality and microbe-driven anaerobic carbon mineralization in peatlands under global warming.

Biogas digestate as a sustainable phytosterol source for biotechnological cascade valorization.

Every year, several million tonnes of anaerobic digestate are produced worldwide as a by-product of the biogas industry, most of which is applied as agricultural fertilizer. However, in the context of a circular bioeconomy, more sustainable uses of residual digestate biomass would be desirable. This study investigates the fate of the sterol lipids ß-sitosterol and cholesterol from the feedstocks to the final digestates of three agricultural and one biowaste biogas plants to assess if sterols are degraded during anaerobic digestion or if they remain in the digestate, which could provide a novel opportunity for digestate cascade valorization. Gas chromatographic analyses showed that feedstock sterols were not degraded during anaerobic digestion, resulting in their accumulation in the digestates to up to 0.15% of the dry weight. The highest concentrations of around 1440 mg ß-sitosterol and 185 mg cholesterol per kg dry weight were found in liquid digestate fractions, suggesting partial sterol solubilization. Methanogenic batch cultures spiked with ß-sitosterol, cholesterol, testosterone and ß-oestradiol confirmed that steroids persist during anaerobic digestion. Mycobacterium neoaurum was able to transform digestate sterols quantitatively into androstadienedione, a platform chemical for steroid hormones, without prior sterol extraction or purification. These results suggest that digestate from agricultural and municipal biowaste is an untapped resource for natural sterols for biotechnological applications, providing a new strategy for digestate cascade valorization beyond land application.

Variation in carbon and nitrogen concentrations among peatland categories at the global scale.

Peatlands account for 15 to 30% of the world's soil carbon (C) stock and are important controls over global nitrogen (N) cycles. However, C and N concentrations are known to vary among peatlands contributing to the uncertainty of global C inventories, but there are few global studies that relate peatland classification to peat chemistry. We analyzed 436 peat cores sampled in 24 countries across six continents and measured C, N, and organic matter (OM) content at three depths down to 70 cm. Sites were distinguished between northern (387) and tropical (49) peatlands and assigned to one of six distinct broadly recognized peatland categories that vary primarily along a pH gradient. Peat C and N concentrations, OM content, and C:N ratios differed significantly among peatland categories, but few differences in chemistry with depth were found within each category. Across all peatlands C and N concentrations in the 10-20 cm layer, were 440 ± 85.1 g kg-1 and 13.9 ± 7.4 g kg-1, with an average C:N ratio of 30.1 ± 20.8. Among peatland categories, median C concentrations were highest in bogs, poor fens and tropical swamps (446-532 g kg-1) and lowest in intermediate and extremely rich fens (375-414 g kg-1). The C:OM ratio in peat was similar across most peatland categories, except in deeper samples from ombrotrophic tropical peat swamps that were higher than other peatlands categories. Peat N concentrations and C:N ratios varied approximately two-fold among peatland categories and N concentrations tended to be higher (and C:N lower) in intermediate fens compared with other peatland types. This study reports on a unique data set and demonstrates that differences in peat C and OM concentrations among broadly classified peatland categories are predictable, which can aid future studies that use land cover assessments to refine global peatland C and N stocks.

A novel belowground in-situ gas labeling approach: CH4 oxidation in deep peat using passive diffusion chambers and 13C excess.

In-vitro incubation of environmental samples is a common approach to estimate CH4 oxidation potential. Here we developed and verified an in-situ method utilizing passive diffusion chambers (PDC, silicone tubes) to deliver 13C-labeled CH4 into peat for the determination of the CH4 oxidation potential based on 13C excess of CO2. To target CH4 oxidation under semi-aerobic and anaerobic conditions, we installed 20 PDCs (30 ml volume) below the water table in profiles from 35-cm to 2-m depths of a peatland in north-eastern Sweden in July 2017 using a peat auger. 13C-labeled CH4 was injected into PDCs through tubing twice during 12 days (day 0 and 6) and samples were collected at days 1, 3, 6, 8 and 11. Background (non-labeled) δ13C of CO2 ranged from -7.3 (35 cm) to +5.7‰ (200 cm) with depth. These δ13C values rose to +110 and + 204‰ after the CH4 injection. The estimated CH4-derived C in CO2 was the lowest at the bottom of the profile (0.3 µmol L-1), whereas the maximum was at 100 cm (6.1 µmol L-1) at five days after the second labeling. This corresponded to 1.5-7.2% of the total CH4 pool to be oxidized, depending on depth. This novel approach with belowground in-situ 13C labeling of gases demonstrated the suitability of tracing the transformations of these gases in soil depth by PDCs and for the first time verified the in-situ occurrence of a deep-peat CH4 oxidation.

Review on the interactions of arsenic, iron (oxy)(hydr)oxides, and dissolved organic matter in soils, sediments, and groundwater in a ternary system.

High concentrations of arsenic (As) in groundwater threaten the environment and public health. Geogenically, groundwater As contamination predominantly occurs via its mobilization from underground As-rich sediments. In an aquatic ecosystem, As is typically driven by several underlying processes, such as redox transitions, microbially driven reduction of iron (Fe) oxide minerals, and release of associated As. Notably, dissolved As mobilized from soils and sediments exhibits high affinity for dissolved organic matter (DOM). Thus, high DOM concentrations can increase As mobility. Therefore, it is crucial to understand the complex interactions and biogeochemical cycling of As, DOM, and Fe oxides. This review collates knowledge regarding the fate of As in multicomponent As-DOM-Fe systems, including ternary complexes involving both Fe and DOM. Additionally, the release mechanisms of As from sediments into groundwater in the presence of both Fe and DOM have been discussed. The mechanisms of As mobilization/sorption at the solid-water interface can be affected by negatively charged DOM competing for sorption sites with As on Fe (oxy)(hydr)oxides and may be further modified by other anionic ubiquitous species such as phosphate, silicic acid, or sulfur. This review emphasizes the need for a comprehensive understanding of the impact of DOM, Fe oxides, and related biogeochemical processes on As mobilization to aquifers. The review identifies important knowledge gaps that may aid in developing applicable practices for preventing the spread of As contamination in aquatic resources and traditional soil management practices.

Control of carbon and nitrogen accumulation by vegetation in pristine bogs of southern Patagonia.

Peatlands are long-term sinks of carbon (C) and nitrogen (N) that are exposed to anthropogenic pressure. This has often induced a vegetation shift from peat mosses towards increasing presence of vascular plants. However, the impact of this vegetation shift on the sink function of peatlands remains unclear. To address this research gap, we studied C and N accumulation in a Patagonian cushion bog where a shift to the predominance of vascular cushion plants is a natural phenomenon since millennia. For comparison, long-term accumulation and decomposition patterns in a pristine Patagonian Sphagnum bog were studied. Thereto, we determined recent and long-term rates of C and N accumulation, their within-site variability, and studied plant-macrofossils. These results were related to decomposition indicators (C/N ratio, humification index, stable isotopes) of the bog types. Despite differences in decomposition indicators, long-term rates of C accumulation were of similar magnitude in the Sphagnum (21.9 g C m-2 yr-1) and in the cushion bog (22.2 g C m-2 yr-1). N accumulation was significantly lower in the Sphagnum bog (0.35 g N m-2 yr-1) compared to the surprisingly high accumulation in the cushion bog (0.55 g N m-2 yr-1). Tephra depositions in the cushion bog about 1600 cal. Years ago presumably triggered the vegetation shift towards dominance of cushion plants by a fertilization effect. C accumulation rates during past decades in the upper decimeters of peat were four times higher in the cushion bog (245 g C m-2 yr-1) compared to the Sphagnum bog (64 g C m-2 yr-1), but substantially decreased since the appearance of cushion plants. High decomposition rates as indicated by decomposition indicators thus apparently offset the higher productivity of cushion plants in the long term. While cushion bogs appear to be effective N sinks, their C sink function may therefore be equal to Sphagnum bogs.

Can nutrient uptake by Carex counteract eutrophication in fen peatlands?

Ground- and surface-water-fed peatlands (i.e., fens) of temperate Europe face high anthropogenic nutrient loads from atmospheric deposition, agricultural catchment areas, and from peat decomposition, if drained. As a result, nitrogen loads may exceed a fen's natural nutrient removal capacity, leading to increased eutrophication of adjacent water bodies. Therefore, it is important to address possible means to decrease a fen's nutrient load, including nutrient uptake by fen plants. To assess how much fen plants can contribute to nutrient removal by uptake, nutrient stocks of above- and below-ground biomass need to be quantified. Therefore, we investigated nitrogen, phosphorous, and potassium uptake capacities of sedges (Carex species), which are common dominants in fen plant communities. We grew specimens of five Carex species with varying preferences in nutrient availability under controlled, different nutrient levels. We show that Carex above-ground biomass harvest can remove up to one third of a system's total nitrogen even at high loads of about 40 g nitrogen m-2. Species-specific differences in biomass production, rather than preferences in nutrient availability under natural conditions, were drivers of standing nutrient stocks: Highly productive species, i.e., C. acutiformis and C. rostrata, had highest nutrient standing stocks across all nutrient levels. Amounts of nutrients stored in shoots increased almost linearly with increasing nutrient levels, whereas below-ground nutrient stocks species-specifically increased, saturated, or decreased, with increasing nutrient levels. As a rough estimate, depending on the species, 6-16 cycles of annual above-ground harvest would suffice to decrease nitrogen concentrations from the highest to the lowest level used in this study. Overall, our results indicate that Carex biomass harvest can be an efficient means to counteract anthropogenic nitrogen eutrophication in fens.

Methane Production Rate during Anoxic Litter Decomposition Depends on Si Mass Fractions, Nutrient Stoichiometry, and Carbon Quality.

While Si influences nutrient stoichiometry and decomposition of graminoid litter, it is still unclear how Si influences anoxic litter decomposition and CH4 formation in graminoid dominated fen peatlands. First, Eriophorum vaginatum plants were grown under different Si and P availabilities, then shoots and roots were characterized regarding their proportions of C, Si, N and P and regarding C quality. Subsequently the Eriophorum shoots were subjected to anoxic decomposition. We hypothesized; that (I) litter grown under high Si availability would show a higher Si but lower nutrient mass fractions and a lower share of recalcitrant carbon moieties; (II) high-Si litter would show higher CH4 and CO2 production rates during anoxic decomposition; (III) methanogenesis would occur earlier in less recalcitrant high-Si litter, compared to low-Si litter. We found a higher Si mass fraction that coincides with a general decrease in C and N mass fractions and decreased share of recalcitrant organic moieties. For high-Si litter, the CH4 production rate was higher, but there was no long-term influence on the CO2 production rate. More labile high-Si litter and a differential response in nutrient stoichiometry led to faster onset of methanogenesis. This may have important implications for our understanding of anaerobic carbon turnover in graminoid-rich fens.

Global CO2 fertilization of Sphagnum peat mosses via suppression of photorespiration during the twentieth century.

Natural peatlands contribute significantly to global carbon sequestration and storage of biomass, most of which derives from Sphagnum peat mosses. Atmospheric CO2 levels have increased dramatically during the twentieth century, from 280 to > 400 ppm, which has affected plant carbon dynamics. Net carbon assimilation is strongly reduced by photorespiration, a process that depends on the CO2 to O2 ratio. Here we investigate the response of the photorespiration to photosynthesis ratio in Sphagnum mosses to recent CO2 increases by comparing deuterium isotopomers of historical and contemporary Sphagnum tissues collected from 36 peat cores from five continents. Rising CO2 levels generally suppressed photorespiration relative to photosynthesis but the magnitude of suppression depended on the current water table depth. By estimating the changes in water table depth, temperature, and precipitation during the twentieth century, we excluded potential effects of these climate parameters on the observed isotopomer responses. Further, we showed that the photorespiration to photosynthesis ratio varied between Sphagnum subgenera, indicating differences in their photosynthetic capacity. The global suppression of photorespiration in Sphagnum suggests an increased net primary production potential in response to the ongoing rise in atmospheric CO2, in particular for mire structures with intermediate water table depths.

Plant succession and geochemical indices in immature peatlands in the Changbai Mountains, northeastern region of China: Implications for climate change and peatland development.

Peatlands cover a small portion of the Earth's land surface but hold ~30% of soil carbon (C) globally. However, few studies have focused on the early stage of peatland development, which is a key stage in the initial C sink function of peatlands. An immature peatland is vulnerable to changes in environmental conditions, e.g., temperature and water conditions, as the peat accumulation process can be easily interrupted by such changes. It is important to understand how immature peatlands develop, what conditions are beneficial to this process, and the present status of these important peatlands. Plant macrofossil analysis and geochemical characteristics of peat were used to determine the plant succession and the degree of decomposition at two peatlands developing in the Changbai Mountain region of northeastern China. The results show that during the entire plant community succession, plants in the two studied peatlands are mainly characterized by sedges (Cyperaceae) and mosses (mainly Sphagnum). Plant macrofossil analysis reveals a wetter trend in the Yuan Lake (YL) peatland in the most upper part of peat layer, which provides favorable conditions for peat accumulation and peatland development. The C/N ratios of core Chi Lake (CL) show a steady peat decomposition and accumulation process in the CL peatland. Additionally, there was a clear impact of presence of Sphagnum on the variations in the C/N ratio. In the YL peatland, macro-charcoal pieces indicated that fire events during dry hydrological conditions had great effects on biogeochemical processes within the peatland, affecting peat decomposition and the succession of the local plant community. An increase in major and trace elements suggests only weak disturbance due to the considerable distance to human settlements. This study determines the characteristics of pristine mountainous peatlands and highlights the importance of understanding the regular plant community in the early stage of peatland formation, as well as its potential effects on C sinks.

A 1-year greenhouse gas budget of a peatland exposed to long-term nutrient infiltration and altered hydrology: high carbon uptake and methane emission.

Long-term increased nutrient influx into normally nutrient-limited peatlands in combination with altered hydrological conditions may threaten a peatland's carbon storage function and affect its greenhouse gas (GHG) budget. However, in situ studies on the effects of long-term altered conditions on peatland functioning and GHG budgets are scarce. We thus quantified GHG fluxes in a peatland exposed to enhanced water level fluctuations and long-term nutrient infiltration in Ontario, Canada, via eddy-covariance and flux chamber measurements. The peatland was a prominent sink of - 680 ± 202 g carbon dioxide (CO2) and a source of 22 ± 8 g methane (CH4) m-2 year-1, resulting in a negative radiative forcing of - 80 g CO2 eq. m-2 y-1. During the growing season CH4 fluxes were constantly high (0.1 g m-2 s-1). Further, on three dates, we measured nitrous oxide (N2O) fluxes and observed a small flux of 2.2 mg m-2 day-1 occurring during the thawing period. Taking the studied ecosystem as a model system for other peatlands exposed to long-term increased nutrient infiltration and enhanced water level fluctuations, our data suggest that such peatlands can maintain their carbon storage function and CO2 sequestration may outweigh emissions of CH4.

Plant communities control long term carbon accumulation and biogeochemical gradients in a Patagonian bog.

Peat carbon accumulation is controlled by both large scale factors, such as climate and hydrological setting, and small scale factors, such as microtopography and plant community. These small scale factors commonly vary within peatlands and can cause variation in biogeochemical traits and carbon accumulation within the same site. To understand these within-site variations, we investigated long term carbon accumulation, peat decomposition, biogeochemistry of pore water and plant macrofossils along a transect in an ombrotrophic bog in southern Patagonia. An additional question we addressed is how historical deposition of volcanic ash on the peatland has affected its carbon balance. Variability in plant community and water table led to differences in long term peat and carbon accumulation (peat moss > cushion plant), organic matter decomposition (cushion plant > peat moss), and methane production (peat moss > cushion plant). Macrofossil analysis and radiocarbon dating indicated a relationship between plant community and carbon accumulation or decomposition during the historical succession of vegetation in the peatland. C/N ratio and isotopic signatures reflected variability in plant community as litter source, and DOC concentrations were controlled by humification level. Volcanic ash deposition had only limited effect on plant composition, but it was associated with increased decomposition in overlying peat layers. This study highlights the importance of understanding how plant communities develop, as changes in communities could significantly affect the potential of ombrotrophic peatlands as C sink.

Occurrence and fate of colloids and colloid-associated metals in a mining-impacted agricultural soil upon prolonged flooding.

Colloids formed during soil flooding can potentially facilitate the mobilization of metal contaminants. Here, laboratory batch incubations with a contaminated soil were performed to monitor temporal changes in the porewater dynamics of metals, the morphology and composition of colloids, and the speciation of colloids-associated metals during 30 days of flooding. The concentrations of colloidal and dissolved metals increased initially and peaked at a certain time, but then decreased with the on-going sulfate reduction. The combined analysis of spectrometric, spectroscopic, and size-fractionation results revealed that the dynamics of Cu were dominated by microbe-associated colloids and were mediated largely by Cu(0) biomineralization and subsequent sulfidation, while the microbe-associated and freely dispersed colloids were equally relevant for governing the dynamics of Cd and Pb. Mobilization of Zn, on the other hand, was dominated by its dissolved form, probably due to the low thermodynamic stability of Zn-sulfide. Additionally, adsorption via organic functional groups was another mechanism for metal incorporation into colloids. We also provided direct spectroscopic evidence for the formation and persistence of dispersed heterocolloids consisting of CuxS and CdS during flooding. Our findings suggest that colloids-induced metal mobilization should be considered in assessing bioavailability and risks of metals in contaminated soils upon flooding.

Enhanced silicon availability leads to increased methane production, nutrient and toxicant mobility in peatlands.

Peatlands perform important ecosystem functions, such as carbon storage and nutrient retention, which are affected, among other factors, by vegetation and peat decomposition. The availability of silicon (Si) in peatlands differs strongly, ranging from <1 to >25 mg L-1. Since decomposition of organic material was recently shown to be accelerated by Si, the aim of this study was to examine how Si influences decomposition of carbon and nutrient and toxicant mobilization in peatlands. We selected a fen site in Northern Bavaria with naturally bioavailable Si pore water concentrations of 5 mg/L and conducted a Si addition experiment. At a fourfold higher Si availability, dissolved organic carbon, carbon dioxide, and methane concentrations increased significantly. Furthermore, dissolved nitrogen, phosphorus, iron, manganese, cobalt, zinc, and arsenic concentrations were significantly higher under high Si availability. This enhanced mobilization may result from Si competing for binding sites but also from stronger reducing conditions, caused by accelerated respiration. The stronger reducing conditions also increased reduction of arsenate to arsenite and thus the mobility of this toxicant. Hence, higher Si availability is suggested to decrease carbon storage and increase nutrient and toxicant mobility in peatland ecosystems.

Does iron reduction control the release of dissolved organic carbon and phosphate at catchment scales? Need for a joint research effort.

Increasing concentrations of dissolved iron and DOC are likely linked to decreasing nitrogen depositon.

Associative nitrogen fixation in nodules of the conifer Lepidothamnus fonkii (Podocarpaceae) inhabiting ombrotrophic bogs in southern Patagonia.

Biological N2 fixation (BNF) in the rhizosphere of Podocarpaceae is currently attributed to unspecific diazotrophs with negligible impact on N acquisition. Here, we report specific and high associative BNF in dead cells of root nodules of Lepidothamnus fonkii distributed in ombrotrophic peatlands of Patagonia. BNF of nodulated roots, intact plants of L. fonkii and rhizospheric peat was assessed by 15N2 and acetylene reduction. Diazotrophs were identified by electron microscopy, analysis of nitrogenase encoding genes (nifH) and transcripts, and 16S rRNA. Nitrogenase encoding nifH transcripts from root nodules point to Beijerinckiaceae (Rhizobiales), known as free-living diazotrophs. Electron microscopy and 16S rRNA analysis likewise identified active Beijerinckiaceae in outer dead cells of root nodules. NifH transcripts from the rhizopshere peat revealed diverse active diazotrophs including Beijerinckiaceae. Both methods revealed high activity of nitrogenase rates in cut roots of L. fonkii (2.5 µmol N g-1 d.w. d-1 based on 15N2 assay; 2.4 µmol C2H4 g-1 d.w. d-1 based on acetylene reduction assay). The data suggest that (i) nodules recruit diazotrophic Beijerinckiaceae from peat, (ii) dead nodule cells provide an exclusive habitat for Beijerinckiaceae, and (iii) BNF in L. fonkii is one potent pathway to overcome N deficiency in ombrotrophic peatlands of Patagonia.

Consortia of low-abundance bacteria drive sulfate reduction-dependent degradation of fermentation products in peat soil microcosms.

Dissimilatory sulfate reduction in peatlands is sustained by a cryptic sulfur cycle and effectively competes with methanogenic degradation pathways. In a series of peat soil microcosms incubated over 50 days, we identified bacterial consortia that responded to small, periodic additions of individual fermentation products (formate, acetate, propionate, lactate or butyrate) in the presence or absence of sulfate. Under sulfate supplementation, net sulfate turnover (ST) steadily increased to 16-174 nmol cm(-3) per day and almost completely blocked methanogenesis. 16S rRNA gene and cDNA amplicon sequencing identified microorganisms whose increases in ribosome numbers strongly correlated to ST. Natively abundant (⩾0.1% estimated genome abundance) species-level operational taxonomic units (OTUs) showed no significant response to sulfate. In contrast, low-abundance OTUs responded significantly to sulfate in incubations with propionate, lactate and butyrate. These OTUs included members of recognized sulfate-reducing taxa (Desulfosporosinus, Desulfopila, Desulfomonile, Desulfovibrio) and also members of taxa that are either yet unknown sulfate reducers or metabolic interaction partners thereof. Most responsive OTUs markedly increased their ribosome content but only weakly increased in abundance. Responsive Desulfosporosinus OTUs even maintained a constantly low population size throughout 50 days, which suggests a novel strategy of rare biosphere members to display activity. Interestingly, two OTUs of the non-sulfate-reducing genus Telmatospirillum (Alphaproteobacteria) showed strongly contrasting preferences towards sulfate in butyrate-amended microcosms, corroborating that closely related microorganisms are not necessarily ecologically coherent. We show that diverse consortia of low-abundance microorganisms can perform peat soil sulfate reduction, a process that exerts control on methane production in these climate-relevant ecosystems.