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The concept of evanescent microbial ecosystems in Earth's atmosphere

This essay presents the hypothesis that short-lived or evanescent microbial ecosystems exist in Earth’s lower troposphere (~ < 4 km). This hypothesis is supported by culture- and molecular-based studies that have shown diverse, viable, and metabolically active microbial communities within Earth’s atmospheric boundary layer. Surprisingly, microorganisms are routinely recovered in samples collected at extreme altitudes including those within the stratosphere (> 18 km). Volcanic eruptions, dust storms, fires, and sea spray are known to seed the atmosphere with microorganisms and to serve as potential nutrient sources while in the atmosphere and upon deposition. Recent research has demonstrated that microorganisms are metabolically active in clouds; for example, archaea capable of utilizing gases such as methane and hydrogen-nitrogen have been identified in clouds and in the atmosphere over natural and anthropogenic gas seeps. The only difference between this hypothesized ecosystem to more traditionally defined ecosystems is its evanescent characteristics where clouds or gas plumes eventually dissipate as they reside over and traverse Earth’s terrestrial and/or aquatic environments. The life cycle of these hypothesized evanescent airborne ecosystems would be short-lived relative to the classically defined biomes or ecosystems.

Book chapter↗

Science questions and knowledge gaps to study microbial transport and survival in Asian and African dust plumes reaching North America

The Sahara in North Africa and the Gobi and Taklamakan deserts in Asia are the primary sources of mobilized dust in the atmosphere, with regional or global airborne transport estimated at 2 to 5 billion tonnes per year. Annual Asian dust plumes take about 7 to 10 d to cross the Pacific Ocean, and often reach the northwest USA between late February and May. In contrast, the peak season for the movement of African dust storms to the southeastern USA is typically June to August, and dust plumes take about 5 to 7 d to reach Florida. Although studies have documented that a wide range of bacteria, fungi, archaea, and viruses in dust plumes reach the USA each year, little is known about temporal and spatial variability in the microbial biodiversity in transoceanic dust plumes, or the effect on the deposition environments. A scoping study (called the Transoceanic Aerobiology Biodiversity Study) was conducted to develop field-based campaigns centered on examining the abundance, diversity, survival, and impact of microorganisms in transoceanic dust plumes arriving in the continental USA from Asia and Africa. This effort identified Science Questions (SQs) and Knowledge Gaps (KGs) that are highly relevant toward an understanding of the microbial diversity, transport, survival, and dispersal in transoceanic dusts. Science Questions were defined as broad science topics in transoceanic dust plume microbiology that were underexplored by the aerobiology community. Knowledge Gaps were defined as specific project-level research questions for each SQ that represented important topics in the study of transoceanic aerobiology.

Aerobiologia↗

Moisture and temperature controls on nitrification differ among ammonia oxidizer communities from three alpine soil habitats

Climate change is altering the timing and magnitude of biogeochemical fluxes in many high elevation ecosystems. The consequent changes in alpine nitrification rates have the potential to influence ecosystem scale responses. In order to better understand how changing temperature and moisture conditions may influence ammonia oxidizers and nitrification activity, we conducted laboratory incubations on soils collected in a Colorado watershed from three alpine habitats (glacial outwash, talus, and meadow). We found that bacteria, not archaea, dominated all ammonia oxidizer communities. Nitrification increased with moisture in all soils and under all temperature treatments. However, temperature was not correlated with nitrification rates in all soils. Site-specific temperature trends suggest the development of generalist ammonia oxidizer communities in soils with greater in situ temperature fluctuations and specialists in soils with more steady temperature regimes. Rapidly increasing temperatures and changing soil moisture conditions could explain recent observations of increased nitrate production in some alpine soils.

Colorado↗

Relationships between microbial communities and environmental parameters at sites impacted by mining of volcanogenic massive sulfide deposits, Prince William Sound, Alaska

The relations among geochemical parameters and sediment microbial communities were examined at three shoreline sites in the Prince William Sound, Alaska, which display varying degrees of impact by acid-rock drainage (ARD) associated with historic mining of volcanogenic massive sulfide deposits. Microbial communities were examined using total fatty acid methyl esters (FAMEs), a class of compounds derived from lipids produced by eukaryotes and prokaryotes (bacteria and Archaea); standard extraction techniques detect FAMEs from both living (viable) and dead (non-viable) biomass, but do not detect Archaeal FAMEs. Biomass and diversity (as estimated by FAMEs) varied strongly as a function of position in the tidal zone, not by study site; subtidal muds, Fe oxyhydroxide undergoing biogenic reductive dissolution, and peat-rich intertidal sediment had the highest values. These estimates were lowest in acid-generating, intertidal zone sediment; if valid, the estimates suggest that only one or two bacterial species predominate in these communities, and/or that Archeal species are important members of the microbial community in this sediment. All samples were dominated by bacterial FAMEs (median value >90%). Samples with the highest absolute abundance of eukaryotic FAMEs were biogenic Fe oxyhydroxides from shallow freshwater pools (fungi) and subtidal muds (diatoms). Eukaryotic FAMEs were practically absent from low-pH, sulfide-rich intertidal zone sediments. The relative abundance of general microbial functional groups such as aerobes/anaerobes and gram(+)/gram(-) was not estimated due to severe inconsistency among the results obtained using several metrics reported in the literature. Principal component analyses (PCAs) were performed to investigate the relationship among samples as separate functions of water, sediment, and FAMEs data. PCAs based on water chemistry and FAMEs data resulted in similar relations among samples, whereas the PCA based on sediment chemistry produced a very different sample arrangement. Specifically, the sediment parameter PCA grouped samples with high bulk trace metal concentration regardless of whether the metals were incorporated into secondary precipitates or primary sulfides. The water chemistry PCA and FAMEs PCA appear to be less prone to this type of artifact. Signature lipids in sulfide-rich sediments could indicate the presence of acid-tolerant and/or acidophilic members of the genus Thiobacillus or they could indicate the presence of SO4-reducing bacteria. The microbial community documented in subtidal and offshore sediments is rich in SRB and/or facultative anaerobes of the Cytophaga-Flavobacterium group; both could reasonably be expected in PWS coastal environments. The results of this study provide evidence for substantial feedback between local (meter to centimeter-scale) geochemical variations, and sediment microbial community composition, and show that microbial community signatures in the intertidal zone are significantly altered at sites where ARD drainage is present relative to sites where it is not, even if the sediment geochemistry indicates net accumulation of ARD-generated trace metals in the intertidal zone. ?? 2007 Elsevier Ltd. All rights reserved.

Applied Geochemistry↗

Hydrogeochemical processes governing the origin, transport and fate of major and trace elements from mine wastes and mineralized rock to surface waters

The formation of acid mine drainage from metals extraction or natural acid rock drainage and its mixing with surface waters is a complex process that depends on petrology and mineralogy, structural geology, geomorphology, surface-water hydrology, hydrogeology, climatology, microbiology, chemistry, and mining and mineral processing history. The concentrations of metals, metalloids, acidity, alkalinity, Cl - , F - and SO 4 2- found in receiving streams, rivers, and lakes are affected by all of these factors and their interactions. Remediation of mine sites is an engineering concern but to design a remediation plan without understanding the hydrogeochemical processes of contaminant mobilization can lead to ineffective and excessively costly remediation. Furthermore, remediation needs a goal commensurate with natural background conditions rather than water-quality standards that might bear little relation to conditions of a highly mineralized terrain. This paper reviews hydrogeochemical generalizations, primarily from US Geological Survey research, that enhance our understanding of the origin, transport, and fate of contaminants released from mined and mineralized areas. Mobility of potential or actual contaminants from mining and mineral processing activities depends on (1) occurrence : is the mineral source of the contaminant actually present? (2) abundance : is the mineral present in sufficient quantity to make a difference? (3) reactivity : what are the energetics, rates, and mechanisms of sorption and mineral dissolution and precipitation relative to the flow rate of the water? and (4) hydrology : what are the main flow paths for contaminated water? Estimates of relative proportions of minerals dissolved and precipitated can be made with mass-balance calculations if minerals and water compositions along a flow path are known. Combined with discharge, these mass-balance estimates quantify the actual weathering rate of pyrite mineralization in the environment and compare reasonably well with laboratory rates of pyrite oxidation except when large quantities of soluble salts and evaporated mine waters have accumulated underground. Quantitative mineralogy with trace-element compositions can substantially improve the identification of source minerals for specific trace elements through mass balances. Post-dissolution sorption and precipitation (attenuation) reactions depend on the chemical behavior of each element, solution composition and pH, aqueous speciation, temperature, and contact-time with mineral surfaces. For example, little metal attenuation occurs in waters of low pH (<3.5) and metals tend to maintain element ratios indicative of the main mineral or group of minerals from which they dissolved, except Fe, SiO 2 , and redox-sensitive oxyanions (As, Sb, Se, Mo, Cr, V). Once dissolved, metal and metalloid concentrations are strongly affected by redox conditions and pH. Iron is the most reactive because it is rapidly oxidized by bacteria and archaea and Fe(III) hydrolyzes and precipitates at low pH (1&ndash;3) which is related directly to its first hydrolysis constant, pK 1 = 2.2. Several insoluble sulfate minerals precipitate at low pH including anglesite, barite, jarosite, alunite and basaluminite. Aluminum hydrolyzes near pH 5 (pK 1 = 5.0) and provides buffering and removal of Al by mineral precipitation from pH 4&ndash;5.5. Dissolved sulfate behaves conservatively because the amount removed from solution by precipitation is usually too small relative to the high concentrations in the water column and relative to the flow rate of the water.

Applied Geochemistry↗

Effect of an algal amendment on the microbial conversion of coal to methane at different sulfate concentrations from the Powder River Basin, USA

Biogenic methane is estimated to account for one-fifth of the natural gas worldwide and there is great interest in controlling methane from different sources. Biogenic coalbed methane (CBM) production relies on syntrophic associations between fermentative bacteria and methanogenic archaea to anaerobically degrade recalcitrant coal and produce methanogenic substrates. However, very little is known about how differences in geochemistry, hydrology, and microbial community composition influence subsurface carbon utilization and CBM production. The addition of an amendment consisting of microalgal biomass has previously been shown to increase CBM production while providing the possibility of a closed-loop fossil system where waste (production water) is used to grow algae to ultimately produce energy (methane). However, the efficiency of enhancing CBM production under different redox conditions remains unresolved. In this study, we focused on the U.S. Geological Survey's Birney test site (Montana, USA) that has nine wells vertically accessing four coal seams with varying geochemistry (low and high sulfate (SO 4 2− )) and methane production rates. We used organic matter (OM) in the form of algal biomass to discern the effect of this amendment on OM degradation and microbially enhanced CBM production potential under different geochemical constraints. We tracked changes in community composition, OM composition, organic carbon (OC) concentration, methane production, and nutrients in batch systems over six months. Methane production was detected only in microcosms from low SO 4 2− wells (168 to 800 μg methane per gram of coal). The OC consumption varied across time for all wells and the variation was greatest for the low SO 4 2− wells. Different groups of syntrophic bacteria were associated with net‑carbon consuming microcosms, and specifically Syntrophorhabdus was identified with several different statistical methods as a potentially important coal degrader. Results from this study provide insight into potential coal-degraders, the compositional changes in some of the different OM fractions, and trends in carbon consumption related to methane production across coal seams along the vertical SO 4 2− gradient.

Montana, Wyoming↗

Changes in microbial communities and associated water and gas geochemistry across a sulfate gradient in coal beds: Powder River Basin, USA

Competition between microbial sulfate reduction and methanogenesis drives cycling of fossil carbon and generation of CH 4 in sedimentary basins. However, little is understood about the fundamental relationship between subsurface aqueous geochemistry and microbiology that drives these processes. Here we relate elemental and isotopic geochemistry of coal-associated water and gas to the microbial community composition from wells in two different coal beds across CH 4 and SO 4 2− gradients (Powder River Basin, Montana, USA). Areas with high CH 4 concentrations generally have higher alkalinity and δ 13 C-DIC values, little to no SO 4 2− , and greater conversion of coal-biodegradable organics to CH 4 (based on δ 13 C-CH 4 and δ 13 C-CO 2 values). Wells with SO 4 2− concentrations from 2 to 10 mM had bacterial populations dominated by several different sulfate-reducing bacteria and archaea that were mostly novel and unclassified. In contrast, in wells with SO 4 2− concentrations <1 mM, the sequences were dominated by presumptive syntrophic bacteria as well as archaeal Methanosarcinales and Methanomicrobiales . The presence of sequences indicative of these bacteria in low SO 4 2− methanogenic wells may suggest a syntrophic role in coal biodegradation and/or the generation of methanogenic substrates from intermediate organic compounds. Archaeal sequences were observed in all sampled zones, with an enrichment of sequences indicative of methanogens in low SO 4 2− zones and unclassified sequences in high SO 4 2− zones. However, sequences indicative of Methanomassiliicoccales were enriched in intermediate SO 4 2− zones and suggest tolerance to SO 4 2− and/or alternative metabolisms in the presence of SO 4 2− . Moreover, sequences indicative of methylotrophic methanogens were more prevalent in an intermediate SO 4 2− and CH 4 well and results suggest an important role for methylotrophic methanogens in critical zone transitions. The presented results demonstrate in situ changes in bacterial and archaeal population distributions along a SO 4 2− gradient associated with recalcitrant, organic carbon that is biodegraded and converted to CO 2 and/or CH 4 .

Powder River Basin↗

Comparison of surficial CO2 efflux to other measures of subsurface crude oil degradation

At a spill site near Bemidji, Minnesota, crude oil at the water table has been undergoing anaerobic biodegradation for over 30 years. Previous work at this site has shown that methane produced from biodegradation of the oil migrates upward and is oxidized in a methanotrophic zone midway between the water table and the surface. To compare microbial activity measurement methods from multiple locations in the oil body, surficial carbon dioxide efflux, methanogen and methanotroph concentrations, and oil degradation state were collected. Carbon dioxide effluxes over the oil body averaged more than four times those at the background site. Methanotrophic bacteria concentrations measured using pmoA were over 10 5 times higher above the oil-contaminated sediments compared with the background site. Methanogenic archaea measured using mcrA ranged from 10 5 to over 10 7 in the oil and were below detection in the background. Methanogens correlated very well with methanotroph concentrations (r = 0.99), n-alkylcyclohexane losses as a proxy for degradation state (r = − 0.96), and somewhat less well with carbon dioxide efflux (r = 0.92). Carbon dioxide efflux similarly correlated to methanotroph concentrations (r = 0.90) and n-alkylcyclohexane losses (r = − 0.91).

Minnesota↗

Organic petrology and geochemistry of Eocene Suzak bituminous marl, north-central Afghanistan: Depositional environment and source rock potential

Organic geochemistry and petrology of Eocene Suzak bituminous marl outcrop samples from Madr village in north-central Afghanistan were characterized via an integrated analytical approach to evaluate depositional environment and source rock potential. Multiple proxies suggest the organic-rich (TOC ∼6 wt.%) bituminous marls are ‘immature’ for oil generation (e.g., vitrinite R o < 0.4%, T max < 425 °C, PI ≤ 0.05, C 29 ααα S/S + R ≤ 0.12, C 29 ββS/ββS+ααR ≤ 0.10, others), yet oil seeps are present at outcrop and live oil and abundant solid bitumen were observed via optical microscopy. Whole rock sulfur content is ∼2.3 wt.% whereas sulfur content is ∼5.0–5.6 wt.% in whole rock extracts with high polar components, consistent with extraction from S-rich Type IIs organic matter which could generate hydrocarbons at low thermal maturity. Low Fe-sulfide mineral abundance and comparison of Pr/Ph ratios between saturate and whole extracts suggest limited Fe concentration resulted in sulfurization of organic matter during early diagenesis. From these observations, we infer that a Type IIs kerogen in ‘immature’ bituminous marl at Madr could be generating high sulfur viscous oil which is seeping from outcrop. However, oil-seep samples were not collected for correlation studies. Aluminum-normalized trace element concentrations indicate enrichment of redox sensitive trace elements Mo, U and V and suggest anoxic-euxinic conditions during sediment deposition. The bulk of organic matter observed via optical microscopy is strongly fluorescent amorphous bituminite grading to lamalginite, possibly representing microbial mat facies. Short chain n- alkanes peak at C 14 –C 16 ( n- C 17 / n- C 29 > 1) indicating organic input from marine algae and/or bacterial biomass, and sterane/hopane ratios are low (0.12–0.14). Monoaromatic steroids are dominated by C 28 clearly indicating a marine setting. High gammacerane index values (∼0.9) are consistent with anoxia stratification and may indicate intermittent saline-hypersaline conditions. Stable C isotope ratios also suggest a marine depositional scenario for the Suzak samples, consistent with the presence of marine foraminifera including abundant planktic globigerinida (?) and rare benthic discocyclina (?) and nummulites (?). Biomarker 2α-methylhopane for photosynthetic cyanobacteria implies shallow photic zone deposition of Madr marls and 3β-methylhopane indicates presence of methanotrophic archaea in the microbial consortium. The data presented herein are consistent with deposition of Suzak bituminous marls in shallow stratified waters of a restricted marine basin associated with the southeastern incipient or proto-Paratethys. Geochemical proxies from Suzak rock extracts (S content, high polar content, C isotopes, normal (αααR) C 27–29 steranes, and C 29 /C 30 and C 26 /C 25 hopane ratios) are similar to extant data from Paleogene oils produced to the north in the Afghan-Tajik Basin. This observation may indicate laterally equivalent strata are effective source rocks as suggested by previous workers; however, further work is needed to strengthen oil-source correlations.

Marine and Petroleum Geology↗

Bacterial and enchytraeid abundance accelerate soil carbon turnover along a lowland vegetation gradient in interior Alaska

Boreal wetlands are characterized by a mosaic of plant communities, including forests, shrublands, grasslands, and fens, which are structured largely by changes in topography and water table position. The soil associated with these plant communities contain quantitatively and qualitatively different forms of soil organic matter (SOM) and nutrient availability that drive changes in biogeochemical cycling rates. Therefore different boreal plant communities likely contain different soil biotic communities which in turn affect rates of organic matter decomposition. We examined relationships between plant communities, microbial communities, enchytraeids, and soil C turnover in near-surface soils along a shallow topographic soil moisture and vegetation gradient in interior Alaska. We tested the hypothesis that as soil moisture increases along the gradient, surface soils would become increasingly dominated by bacteria and mesofauna and have more rapid rates of C turnover. We utilized bomb radiocarbon techniques to infer rates of C turnover and the 13C isotopic composition of SOM and respired CO 2 to infer the degree of soil humification. Soil phenol oxidase and peroxidase enzyme activities were generally higher in the rich fen compared with the forest and bog birch sites. Results indicated greater C fluxes and more rapid C turnover in the surface soils of the fen sites compared to the wetland forest and shrub sites. Quantitative PCR analyses of soil bacteria and archaea, combined with enchytraeid counts, indicated that surface soils from the lowland fen ecosystems had higher abundances of these microbial and mesofaunal groups. Fungal abundance was highly variable and not significantly different among sites. Microbial data was utilized in a food web model that confirmed that rapidly cycling systems are dominated by bacterial activity and enchytraeid grazing. However, our results also suggest that oxidative enzymes play an important role in the C mineralization process in saturated systems, which has been often ignored.

Soil Biology and Biochemistry↗

Nitrifying microorganisms linked to biotransformation of perfluoroalkyl sulfonamido precursors from legacy aqueous film forming foams

Drinking water supplies across the United States have been contaminated by firefighting and fire-training activities that use aqueous film-forming foams (AFFF) containing per- and polyfluoroalkyl substances (PFAS). Much of the AFFF is manufactured using electrochemical fluorination by 3M. Precursors with six perfluorinated carbons (C6) and non-fluorinated amine substituents make up approximately one-third of the PFAS in 3M AFFF. C6 precursors can be transformed through nitrification (microbial oxidation) of amine moieties into perfluorohexane sulfonate (PFHxS), a compound of regulatory concern. Here, we report biotransformation of the most abundant C6 sulfonamido precursors in 3M AFFF with available commercial standards (FHxSA, PFHxSAm, and PFHxSAmS) in microcosms representative of the groundwater/surface water boundary. Results show rapid (<1 day) biosorption to living cells by precursors but slow biotransformation into PFHxS (1–100 pM day –1 ). The transformation pathway includes one or two nitrification steps and is supported by the detection of key intermediates using high-resolution mass spectrometry. Increasing nitrate concentrations and total abundance of nitrifying taxa occur in parallel with precursor biotransformation. Together, these data provide multiple lines of evidence supporting microbially limited biotransformation of C6 sulfonamido precursors involving ammonia-oxidizing archaea ( Nitrososphaeria ) and nitrite-oxidizing bacteria ( Nitrospina ). Further elucidation of interrelationships between precursor biotransformation and nitrogen cycling in ecosystems would help inform site remediation efforts.

Environmental Science and Technology↗

A hydrogen-based subsurface microbial community dominated by methanogens

The search for extraterrestrial life may be facilitated if ecosystems can be found on Earth that exist under conditions analogous to those present on other planets or moons. It has been proposed, on the basis of geochemical and thermodynamic considerations, that geologically derived hydrogen might support subsurface microbial communities on Mars and Europa in which methanogens form the base of the ecosystem1-5. Here we describe a unique subsurface microbial community in which hydrogen-consuming, methane-producing Archaea far outnumber the Bacteria. More than 90% of the 16s ribosomal DNA sequences recovered from hydrothermal waters circulating through deeply buried igneous rocks in Idaho are related to hydrogen-using methanogenic microorganisms. Geochemical characterization indicates that geothermal hydrogen, not organic carbon, is the primary energy source for this methanogen-dominated microbial community. These results demonstrate that hydrogen-based methanogenic communities do occur in Earth's subsurface, providing an analogue for possible subsurface microbial ecosystems on other planets.

Nature↗

Expanding the repertoire of electron acceptors for the anaerobic oxidation of methane in carbonates in the Atlantic and Pacific Ocean

Authigenic carbonates represent a significant microbial sink for methane, yet little is known about the microbiome responsible for the methane removal. We identify carbonate microbiomes distributed over 21 locations hosted by seven different cold seeps in the Pacific and Atlantic Oceans by carrying out a gene-based survey using 16S rRNA- and mcr A gene sequencing coupled with metagenomic analyses. Based on 16S rRNA gene amplicon analyses, these sites were dominated by bacteria affiliated to the Firmicutes, Alpha- and Gammaproteobacteria. ANME-1 and -2 archaeal clades were abundant in the carbonates yet their typical syntrophic partners, sulfate-reducing bacteria, were not significantly present. Based on mcrA amplicon analyses, the Candidatus Methanoperedens clades were also highly abundant. Our metagenome analysis indicated that methane oxidizers affiliated to the ANME-1 and -2, may be capable of performing complete methane- and potentially short-chain alkane oxidation independently using oxidized sulfur and nitrogen compounds as terminal electron acceptors. Gammaproteobacteria are hypothetically capable of utilizing oxidized nitrogen compounds and may be involved in syntrophy with methane-oxidizing archaea. Carbonate structures represent a window for a more diverse utilization of electron acceptors for anaerobic methane oxidation along the Atlantic and Pacific Margin.

ISME Journal↗

Metabolically diverse microorganisms mediate methylmercury formation under nitrate-reducing conditions in a dynamic hydroelectric reservoir

Brownlee Reservoir is a mercury (Hg)-impaired hydroelectric reservoir that exhibits dynamic hydrological and geochemical conditions and is located within the Hells Canyon Complex in Idaho, USA. Methylmercury (MeHg) contamination in fish is a concern in the reservoir. While MeHg production has historically been attributed to sulfate-reducing bacteria and methanogenic archaea, microorganisms carrying the hgcA gene are taxonomically and metabolically diverse and the major biogeochemical cycles driving mercury (Hg) methylation are not well understood. In this study, Hg speciation and redox-active compounds were measured throughout Brownlee Reservoir across the stratified period in four consecutive years (2016–2019) to identify the location where and redox conditions under which MeHg is produced. Metagenomic sequencing was performed on a subset of samples to characterize the microbial community with hgcA and identify possible links between biogeochemical cycles and MeHg production. Biogeochemical profiles suggested in situ water column Hg methylation was the major source of MeHg. These profiles, combined with genome-resolved metagenomics focused on hgcA -carrying microbes, indicated that MeHg production occurs in this system under nitrate- or manganese-reducing conditions, which were previously thought to preclude Hg-methylation. Using this multidisciplinary approach, we identified the cascading effects of interannual variability in hydrology on the redox status, microbial metabolic strategies, abundance and metabolic diversity of Hg methylators, and ultimately MeHg concentrations throughout the reservoir. This work expands the known conditions conducive to producing MeHg and suggests that the Hg-methylation mitigation efforts by nitrate or manganese amendment may be unsuccessful in some locations.

Idaho↗

Large carbon isotope fractionation associated with oxidation of methyl halides by methylotrophic bacteria

The largest biological fractionations of stable carbon isotopes observed in nature occur during production of methane by methanogenic archaea. These fractionations result in substantial (as much as ≈70‰) shifts in δ 13 C relative to the initial substrate. We now report that a stable carbon isotopic fractionation of comparable magnitude (up to 70‰) occurs during oxidation of methyl halides by methylotrophic bacteria. We have demonstrated biological fractionation with whole cells of three methylotrophs (strain IMB-1, strain CC495, and strain MB2) and, to a lesser extent, with the purified cobalamin-dependent methyltransferase enzyme obtained from strain CC495. Thus, the genetic similarities recently reported between methylotrophs, and methanogens with respect to their pathways for C 1 -unit metabolism are also reflected in the carbon isotopic fractionations achieved by these organisms. We found that only part of the observed fractionation of carbon isotopes could be accounted for by the activity of the corrinoid methyltransferase enzyme, suggesting fractionation by enzymes further along the degradation pathway. These observations are of potential biogeochemical significance in the application of stable carbon isotope ratios to constrain the tropospheric budgets for the ozone-depleting halocarbons, methyl bromide and methyl chloride. Methyl bromide (MeBr) and methyl chloride (MeCl) are, respectively, the most abundant volatile bromo- and chlorocarbons in the troposphere and are major contributors to stratospheric ozone destruction (1). Both compounds have natural and human-influenced sources and a predominant sink by reaction with OH in the troposphere (2–4). MeBr also has a bacterial soil sink (5) that represents about 20% of the estimated total removal from the troposphere, and it is likely that a soil sink of similar magnitude exists for MeCl (6). Hence, if an isotopic fractionation is associated with the soil sink, it will influence the isotopic compositions of MeBr and MeCl in the lower atmosphere (7). The δ 13 C value of industrially produced MeBr ranges between −43.5‰ and −66.4‰ (7), but δ 13 C values of tropospheric MeBr and natural sources are not yet known. The δ 13 C of atmospheric MeCl has been measured from −22‰ to −45‰ (8, 9). If carbon isotope ratios are to be used to constrain the budgets of these methyl halides, it is essential to determine the extent of carbon isotope fractionation that occurs during biological degradation of these compounds. Methylotrophic bacteria use C 1 compounds, which are simple organic molecules that contain no carbon–carbon bonds. Strains IMB-1, CC495, and MB2 are as-yet-unnamed facultative methylotrophs isolated from agricultural soil, woodland leaf litter, and coastal seawater, respectively (10–13), environments where methyl halides are produced. They are members of the α subgroup of the Proteobacteria . On the basis of 16S rRNA gene sequences, strains IMB-1 and CC495 show some phylogenetic alignment with the genus Rhizobium (10, 11) and are very closely related to the new genus Pseudoaminobacte r (I. McDonald, personal communication). Strain MB2 aligns within the Ruegeria clade [J. K. Schaefer, K. D. Goodwin, I. R. McDonald, J. C. Murrell and R.S.O., unpublished work]. All of these aerobic bacteria are methylotrophs in that they can grow by using MeBr or MeCl as their sole carbon source, but they do not metabolize methane. They oxidize MeBr, MeCl, and methyl iodide (MeI) to CO 2 . Soil bacteria are known to consume MeBr at the ambient tropospheric mixing ratio of around 10 parts per trillion by volume (5). Preliminary experiments with strain IMB-1 indicate that it can oxidize MeBr at these mixing ratios** and is therefore likely to be characteristic of bacteria associated with MeBr uptake by soils. We examined δ 13 C of MeCl, MeBr, and MeI during oxidation by whole-cell suspensions of IMB-1 and CC495 and also the change in δ 13 C values of the three methyl halides during oxidation by the marine strain MB2. In addition, we measured the fractionation of carbon isotopes during formation of methane thiol (MeSH) from MeCl by the purified cobalamin-dependent enzyme, halomethane:bisulphide/halide ion methyltransferase (11) from CC495, to determine whether this initial step in MeCl degradation could account for the observed fractionation by whole cells. Finally, we determined the fractionation associated with the degradation of MeBr during field studies with agricultural soil by monitoring MeBr concentration and δ 13 C of MeBr in the headspace of flux chambers under fumigation conditions.

Proceedings of the National Academy of Sciences of↗

Advances in the hydrogeochemistry and microbiology of acid mine waters

The last decade has witnessed a plethora of research related to the hydrogeochemistry and microbiology of acid mine waters and associated tailings and waste-rock waters. Numerous books, reviews, technical papers, and proceedings have been published that examine the complex bio-geochemical process of sulfide mineral oxidation, develop and apply geochemical models to site characterization, and characterize the microbial ecology of these environments. This review summarizes many of these recent works, and provides references for those investigating this field. Comparisons of measured versus calculated Eh and measured versus calculated pH for water samples from several field sites demonstrate the reliability of some current geochemical models for aqueous speciation and mass balances. Geochemical models are not, however, used to predict accurately time-dependent processes but to improve our understanding of these systems and to constrain possible processes that contribute to actual or potential water quality issues. Microbiological studies are demonstrating that there is much we have yet to learn about the types of different microorganisms and their function and ecology in mine-waste environments. A broad diversity of green algae, bacteria, archaea, yeasts, and fungi are encountered in acid mine waters, and a better understanding of their ecology and function may potentially enhance remediation possibilities as well as our understanding of the evolution of life.

International Geology Review↗

Relation between the relative abundance and collapse of Aphanizomenon flos-aquae and microbial antagonism in Upper Klamath Lake, Oregon

Aphanizomenon flos-aquae (AFA) is the dominant filamentous cyanobacterium that develops into blooms in Upper Klamath Lake, Oregon each year. During AFA bloom and collapse, ecosystem conditions for endangered Lost River and shortnose suckers deteriorate, thus motivating the need to identify processes that limit AFA abundance and decline. Here we investigate the relations between AFA and other members of the microbial community (photosynthetic and non-photosynthetic bacteria and archaea), how those relations impact abundance and collapse of AFA, and the types of microbial conditions that suppress AFA. We found significant spatial variation in AFA relative abundance during the 2016 bloom period using 16S rRNA sequencing. The Pelican Marina (PM) site had the lowest AFA relative abundance, and this was coincident with increased relative abundance of Candidatus Sericytochromatia , Flavobacterium , and Rheinheimera , some of which are known AFA antagonists. The AFA collapse coincided with phosphorus limitation relative to nitrogen and the increased relative abundance of Cyanobium and Candidatus Sericytochromatia, which outcompete AFA when dissolved inorganic nitrogen is available. The data collected in this study indicate the importance of dissolved inorganic nitrogen combined with microbial community structure in suppressing AFA abundance.

Oregon↗

Sulfide stress tolerance as a controller of methane production in temperate wetlands

Wetlands are a major source of methane emissions and contribute to the observed increase in atmospheric methane over the last 20 years. Methane production in wetlands is the final step of carbon decomposition performed by anaerobic archaea. Although hydrogen/carbon dioxide and acetate are the substrates most often attributed to methanogenesis, other substrates—such as methylated compounds—may additionally play important roles in driving methane production in wetland systems. Here we conducted mesocosm experiments combined with genome-resolved metatranscriptomics to investigate the impact of diverse methanogenic substrate amendment on methanogenesis in two high methane-emitting wetlands with distinct geochemistry, termed P7 and P8. Methanol amendment resulted in high methane production at both sites, whereas acetate and formate amendment only stimulated methanogenesis in P7 mesocosms, where aqueous sulfide concentrations were lower. In P7 sediments, formate amendment fueled acetogenic microbes that produced acetate, which was subsequently utilized by acetoclastic methanogens. In contrast to expression profiles in P7 mesocosms, active methylotrophic methanogen genomes from P8 showed increased expression of genes related to membrane remodeling and DNA damage repair, indicative of stress tolerance mechanisms to counter sulfide toxicity. Methylotrophic methanogenesis generates higher free energy yields than acetoclastic methanogenesis, which likely enables allocation of more energy toward stress responses. These findings contribute to the growing body of literature highlighting methylotrophic methanogenesis as an important methane production pathway in wetlands. By using less competitive substrates like methanol that provide greater energy yields, methylotrophic methanogens may invest in physiological strategies that provide competitive advantages across a range of environmental stresses.

North Dakota↗