Changes in Quinone Profiles of Hot Spring Microbial Mats with a Thermal Gradient

The respiratory and photosynthetic quinones of microbial mats which occurred in Japanese sulfide-containing neutral-pH hot springs at different temperatures were analyzed by spectrochromatography and mass spectrometry. All of the microbial mats that developed at high temperatures (temperatures above 68°C) were so-called sulfur-turf bacterial mats and produced methionaquinones (MTKs) as the major quinones. A 78°C hot spring sediment had a similar quinone profile. Chloroflexus-mixed mats occurred at temperatures of 61 to 65°C and contained menaquinone 10 (MK-10) as the major component together with significant amounts of either MTKs or plastoquinone 9 (PQ-9). The sunlight-exposed biomats growing at temperatures of 45 to 56°C were all cyanobacterial mats, in which the photosynthetic quinones (PQ-9 and phylloquinone) predominated and MK-10 was the next most abundant component in most cases. Ubiquinones (UQs) were not found or were detected in only small amounts in the biomats growing at temperatures of 50°C and above, whereas the majority of the quinones of a purple photosynthetic mat growing at 34°C were UQs. A numerical analysis of the quinone profiles was performed by using the following three parameters: dissimilarity index (D), microbial divergence index (MD_q), and bioenergetic divergence index (BD_q). A D matrix tree analysis showed that the hot spring mats consisting of the sulfur-turf bacteria, Chloroflexus spp., cyanobacteria, and purple phototrophic bacteria formed distinct clusters. Analyses of MD_q and BD_q values indicated that the microbial diversity of hot spring mats decreased as the temperature of the environment increased. The changes in quinone profiles and physiological types of microbial mats in hot springs with thermal gradients are discussed from evolutionary viewpoints.     

Biodiversity of the microbial mat of the Garga hot spring

Background

Microbial mats are a good model system for ecological and evolutionary analysis of microbial communities. There are more than 20 alkaline hot springs on the banks of the Barguzin river inflows. Water temperature reaches 75 °C and pH is usually 8.0–9.0. The formation of microbial mats is observed in all hot springs. Microbial communities of hot springs of the Baikal rift zone are poorly studied. Garga is the biggest hot spring in this area.

Results

In this study, we investigated bacterial and archaeal diversity of the Garga hot spring (Baikal rift zone, Russia) using 16S rRNA metagenomic sequencing. We studied two types of microbial communities: (i) small white biofilms on rocks in the points with the highest temperature (75 °C) and (ii) continuous thick phototrophic microbial mats observed at temperatures below 70 °C. Archaea (mainly Crenarchaeota; 19.8% of the total sequences) were detected only in the small biofilms. The high abundance of Archaea in the sample from hot springs of the Baikal rift zone supplemented our knowledge of the distribution of Archaea. Most archaeal sequences had low similarity to known Archaea. In the microbial mats, primary products were formed by cyanobacteria of the genus Leptolyngbya. Heterotrophic microorganisms were mostly represented by Actinobacteria and Proteobacteria in all studied samples of the microbial mats. Planctomycetes, Chloroflexi, and Chlorobi were abundant in the middle layer of the microbial mats, while heterotrophic microorganisms represented mostly by Firmicutes (Clostridia, strict anaerobes) dominated in the bottom part. Besides prokaryotes, we detect some species of Algae with help of detection their chloroplasts 16 s rRNA.

Conclusions

High abundance of Archaea in samples from hot springs of the Baikal rift zone supplemented our knowledge of the distribution of Archaea. Most archaeal sequences had low similarity to known Archaea. Metagenomic analysis of microbial communities of the microbial mat of Garga hot spring showed that the three studied points sampled at 70 °C, 55 °C, and 45 °C had similar species composition. Cyanobacteria of the genus Leptolyngbya dominated in the upper layer of the microbial mat. Chloroflexi and Chlorobi were less abundant and were mostly observed in the middle part of the microbial mat. We detected domains of heterotrophic organisms in high abundance (Proteobacteria, Firmicutes, Verrucomicrobia, Planctomicetes, Bacteroidetes, Actinobacteria, Thermi), according to metabolic properties of known relatives, which can form complete cycles of carbon, sulphur, and nitrogen in the microbial mat. The studied microbial mats evolved in early stages of biosphere formation. They can live autonomously, providing full cycles of substances and preventing live activity products poisoning.

     

Characterization of Microbial Mats from a Desert Wadi Ecosystem in the Sultanate of Oman

Desert wadis are widespread in the Arabian Peninsula and play a vital role in the ecology of the region; nevertheless, these ecosystems are among the least studied. Various types of microbial mats are predominant in wadis, but information on their bacterial diversity and spatial distribution is very scarce. We investigated bacterial diversity, pigments and lipid composition of ten mats located at the down-, mid- and upstream of a desert wadi in Oman. Direct microscopy revealed the existence of different unicellular and filamentous cyanobacteria, with the dominance of the heterocystous genera Calothrix and Scytonema. The majority of MiSeq 16S rRNA sequences (44-76%) were affiliated to Cyanobacteria and Proteobacteria. While Alphaproteobacteria was the most dominant proteobacterial class (10 to 48% of total sequences), Gamma- and Deltaproteobacteria were subdominant. Cluster analysis showed that the mats’ bacterial communities at the different locations along the wadi were different and shared less than 60% of their operational taxonomic units (OTUs). Chlorophyll a and scytonemin were the most predominant pigments in all mats. Different saturated, branched and mono- and poly-unsaturated fatty acids were detected in all mats, with C₁₆ and C₁₈ compounds as most dominant. The detected pigments and fatty acids indicate a major role of cyanobacteria in the wadi mats and the adaptation of microorganisms therein to the harsh wadi environment. Detection of diadinoxanthin and fucoxanthin confirmed the presence of diatoms. We conclude that microbial mats are important elements in wadi ecosystems and exist in a great variety of structure and community composition.     

Uptake and transformation of metals and metalloids by microbial mats and their use in bioremediation

Summary Constructed microbial mats, used for studies on the removal and transformation of metals and metalloids, are made by combining cyanobacteria inoculum with a sediment inoculum from a metal-contaminated site. These mats are a heterotrophic and autotrophic community dominated by cyanobacteria and held together by slimy secretions produced by various microbial groups. When contaminated water containing high concentrations of metals is passed over microbial mats immobilized on glass wool, there is rapid removal of the metals from the water. The mats are tolerant of high concentrations of toxic metals and metalloids, such as cadmium, lead, chromium, selenium and arsenic (up to 350 mg L^–1). This tolerance may be due to a number of mechanisms at the molecular, cellular and community levels. Management of toxic metals by the mats is related to deposition of metal compounds outside the cell surfaces as well as chemical modification of the aqueous environment surrounding the mats. The location of metal deposition is determined by factors such as redox gradients, cell surface micro-environments and secretion of extra-cellular bioflocculents. Metal-binding flocculents (polyanionic polysaccharides) are produced in large quantities by the cyanobacterial component of the mat. Steep gradients of redox and oxygen exist from the surface through the laminated strata of microbes. These are produced by photosynthetic oxygen production at the surface and heterotrophic consumption in the deeper regions. Additionally, sulfur-reducing bacteria colonize the lower strata, removing and utilizing the reducing H₂S, rather than water, for photosynthesis. Thus, depending on the chemical character of the microzone of the mat, the sequestered metals or metalloids can be oxidized, reduced and precipitated as sulfides or oxides. For example precipitates of red amorphous elemental selenium were identified in mats exposed to selenate (Se-VI) and insoluble precipitates of manganese, chromium, cadmium, cobalt, and lead were found in mats exposed to soluble salts of these metals. Constructed microbial mats offer several advantages for use in the bioremediation of metal-contaminated sites. These include low cost, durability, ability to function in both fresh and salt water, tolerance to high concentrations of metals and metalloids and the unique capacity of mats to form associations with new microbial species. Thus one or several desired microbial species might be integrated into mats in order to design the community for specific bioremediation applications.     

Structure and function of natural sulphide-oxidizing microbial mats under dynamic input of light and chemical energy

We studied the interaction between phototrophic and chemolithoautotrophic sulphide-oxidizing microorganisms in natural microbial mats forming in sulphidic streams. The structure of these mats varied between two end-members: one characterized by a layer dominated by large sulphur-oxidizing bacteria (SOB; mostly Beggiatoa-like) on top of a cyanobacterial layer (B/C mats) and the other with an inverted structure (C/B mats). C/B mats formed where the availability of oxygen from the water column was limited (<5 μm). Aerobic chemolithotrophic activity of the SOB depended entirely on oxygen produced locally by cyanobacteria during high light conditions. In contrast, B/C mats formed at locations where oxygen in the water column was comparatively abundant (>45 μM) and continuously present. Here SOB were independent of the photosynthetic activity of cyanobacteria and outcompeted the cyanobacteria in the uppermost layer of the mat where energy sources for both functional groups were concentrated. Outcompetition of photosynthetic microbes in the presence of light was facilitated by the decoupling of aerobic chemolithotrophy and oxygenic phototrophy. Remarkably, the B/C mats conserved much less energy than the C/B mats, although similar amounts of light and chemical energy were available. Thus ecosystems do not necessarily develop towards optimal energy usage. Our data suggest that, when two independent sources of energy are available, the structure and activity of microbial communities is primarily determined by the continuous rather than the intermittent energy source, even if the time-integrated energy flux of the intermittent energy source is greater.     

Spatial distribution of diatom and cyanobacterial mats in the Dead Sea is determined by response to rapid salinity fluctuations

Cyanobacteria and diatom mats are ubiquitous in hypersaline environments but have never been observed in the Dead Sea, one of the most hypersaline lakes on Earth. Here we report the discovery of phototrophic microbial mats at underwater freshwater seeps in the Dead Sea. These mats are either dominated by diatoms or unicellular cyanobacteria and are spatially separated. Using in situ and ex situ O₂ microsensor measurements we show that these organisms are photosynthetically active in their natural habitat. The diatoms, which are phylogenetically associated to the Navicula genus, grew in culture at salinities up to 40 % Dead Sea water (DSW) (14 % total dissolved salts, TDS). The unicellular cyanobacteria belong to the extremely halotolerant Euhalothece genus and grew at salinities up to 70 % DSW (24.5 % TDS). As suggested by a variable O₂ penetration depth measured in situ, the organisms are exposed to drastic salinity fluctuations ranging from brackish to DSW salinity within minutes to hours. We could demonstrate that both phototrophs are able to withstand such extreme short-term fluctuations. Nevertheless, while the diatoms recover better from rapid fluctuations, the cyanobacteria cope better with long-term exposure to DSW. We conclude that the main reason for the development of these microbial mats is a local dilution of the hypersaline Dead Sea to levels allowing growth. Their spatial distribution in the seeping areas is a result of different recovery rates from short or long-term fluctuation in salinity.     

The Structure and Biogeochemical Activity of the Phototrophic Communities from the Bol'sherechenskii Alkaline Hot Spring

Microbial communities growing in the bed of the alkaline, sulfide hot spring Bol'sherechenskii (the Baikal rift area) were studied over many years (1986–2001). The effluent water temperature ranged from 72 to 74°C, pH was from 9.25 to 9.8, and sulfide content was from 12 to 13.4 mg/ml. Simultaneous effects of several extreme factors restrict the spread of phototrophic microorganisms. Visible microbial mat appears with a decrease in the temperature to 62°C and in sulfide content to 5.9 mg/l. Cyanobacteria predominated in all biological zones of the microbial mat. The filamentous cyanobacteria of the genus Phormidium are the major mat-forming organisms, whereas unicellular cyanobacteria and the filamentous green bacterium Chloroflexus aurantiacus are minor components of the phototrophic communities. No cyanobacteria of the species Mastigocladus laminosus, typical of neutral and subacid springs, were identified. Seventeen species of both anoxygenic phototrophic bacteria and cyanobacteria were isolated from the microbial mats, most of which exhibited optimum growth at 20 to 45°C. The anoxygenic phototrophs were neutrophiles with pH optimum at about 7. The cyanobacteria were the most adapted to the alkaline conditions in the spring. Their optimum growth was observed at pH 8.5–9.0. As determined by the in situ radioisotope method, the optimal growth and decomposition rates were observed at 40–32°C, which is 10–15°C lower than the same parameter in the sulfide-deficient Octopus Spring (Yellowstone, United States). The maximum chlorophyll a concentration was 555 mg/m² at 40°C. The total rate of photosynthesis in the mats reached 1.3 g C/m² per day. The maximum rate of dark fixation of carbon dioxide in the microbial mats was 0.806 g C/m² per day. The maximum rate of sulfate reduction comprised 0.367 g S/m² per day at 40°C. The rate of methanogenesis did not exceed 1.188 g C/m² per day. The role of methanogenesis in the terminal decomposition of the organic matter was insignificant. Methane formation consumed 100 times less organic matter than sulfate reduction.     

Cyanobacterial mats from hot springs produce antimicrobial compounds and quorum-sensing inhibitors under natural conditions

Polar (water) and non-polar (ethyl acetate) extracts from the cyanobacterial layer (top 1–3 mm) of four hot spring microbial mats in the Sultanate of Oman were tested for their antibacterial, antidiatom and quorum-sensing inhibitory activities under natural conditions. The chemical composition of the active extracts was analysed using gas chromatography–mass spectrometry (GC-MS). Cyanobacteria within these mats were identified by direct microscopy while the total bacterial community composition was compared using automated ribosomal intergenic spacer analysis (ARISA). Only the extracts from Bowshar and Nakhl mats showed antibacterial properties against Bacillus sp., Micrococcus luteus, Shigella sonnei, Salmonella enterica and Klebsiella pneumoniae. All tested extracts inhibited the growth of the benthic diatom Amphora coffeaeformis. Extracts from Bowshar, Rustaq and Nakhl inhibited quorum-sensing of the reporter strains Chromobacterium violaceum CV017 and Agrobacterium tumefaciens NTL4. The highest bioactivity was recorded for ethyl acetate extracts from Nakhl mats, which had the lowest number of operational taxonomic units (OTUs). Using GC-MS, 74 chemical compounds were obtained, however with different distribution among the four mat extracts (similarity < 43%). Various cyanobacteria, belonging mainly to Chroococcus, Phormidium, Leptolyngbya, Spirulina and Lyngbya were detected in the different mats, and each mat had its unique bacterial community, as confirmed by ARISA profiles. We conclude that antimicrobial and quorum-sensing inhibitory compounds can be produced by hot spring mat microorganisms under natural conditions and the differences in these compounds could be attributed to the differences in the mats’ bacterial composition as well as the physical–chemical conditions of the springs.     

Distribution of greenhouse gases in hyper-arid and arid areas of northern Chile and the contribution of the high altitude wetland microbiome (Salar de Huasco,Chile)

Northern Chile harbors different bioclimatic zones including hyper-arid and arid ecosystems and hotspots of microbial life, such as high altitude wetlands, which may contribute differentially to greenhouse gases (GHG) such as carbon dioxide (CO₂), methane (CH₄) and nitrous oxide (N₂O). In this study, we explored ground level GHG distribution and the potential role of a wetland situated at 3800 m.a.s.l, and characterized by high solar radiation <?1600 W m^?2, extreme temperature ranges (?12 to 24 °C) and wind stress (<?17 m s^?1). The water source of the wetland is mainly groundwater springs, which generates streams and ponds surrounded by peatlands. These sites support a rich microbial aquatic life including diverse bacteria and archaea communities, which transiently form more complex structures, such as microbial mats. In this study, GHG were measured in the water and above ground level air at the wetland site and along an elevation gradient in different bioclimatic areas from arid to hyper-arid zones. The microbiome from the water and sediments was described by high-throughput sequencing 16S rRNA and rDNA genes. The results indicate that GHG at ground level were variable along the elevation gradient potentially associated with different bioclimatic zones, reaching high values at the high Andean steppe and variable but lower values in the Atacama Desert and at the wetland. The water areas of the wetland presented high concentrations of CH₄ and CO₂, particularly at the spring areas and in air bubbles below microbial mats. The microbial community was rich (>?40 phyla), including archaea and bacteria potentially active in the different matrices studied (water, sediments and mats). Functional microbial groups associated with GHG recycling were detected at low frequency, i.e., <?2.5% of total sequences. Our results indicate that hyper-arid and arid areas of northern Chile are sites of GHG exchange associated with various bioclimatic zones and particularly in aquatic areas of the wetland where this ecosystem could represent a net sink of N₂O and a source for CH₄ and CO₂.     

Phylogenetic Evidence for the Existence of Novel Thermophilic Bacteria in Hot Spring Sulfur-Turf Microbial Mats in Japan

So-called sulfur-turf microbial mats, which are macroscopic white filaments or bundles consisting of large sausage-shaped bacteria and elemental sulfur particles, occur in sulfide-containing hot springs in Japan. However, no thermophiles from sulfur-turf mats have yet been isolated as cultivable strains. This study was undertaken to determine the phylogenetic positions of the sausage-shaped bacteria in sulfur-turf mats by direct cloning and sequencing of 16S rRNA genes amplified from the bulk DNAs of the mats. Common clones with 16S rDNA sequences with similarity levels of 94.8 to 99% were isolated from sulfur-turf mat samples from two geographically remote hot springs. Phylogenetic analysis showed that the phylotypes of the common clones formed a major cluster with members of the Aquifex-Hydrogenobacter complex, which represents the most deeply branching lineage of the domain bacteria. Furthermore, the bacteria of the sulfur-turf mat phylotypes formed a clade distinguishable from that of other members of the Aquifex-Hydrogenobacter complex at the order or subclass level. In situ hybridization with clone-specific probes for 16S rRNA revealed that the common phylotype of sulfur-turf mat bacteria is that of the predominant sausage-shaped bacteria.Microbial mats develop in a wide variety of aquatic environments, including geothermal hot springs and hydrothermal vents. There are several types of thermophilic microbial mats, e.g., those of cyanobacteria, anoxygenic phototrophic bacteria, and chemotrophic sulfur bacteria, which differ according to the physical and chemical conditions they favor and other environmental factors (10, 38). These microbial mats in thermal habitats have been studied extensively as a peculiar microbial community of the ecosystem, in relation to the phylogeny and evolution of thermophilic prokaryotes, or as a source of new functional enzymes.So-called sulfur-turf microbial mats are macroscopic bundles of white filaments consisting of colorless sulfur bacteria and elemental sulfur particles that form in shallow streams of sulfide-containing high-temperature hot springs. Since first reported by Miyoshi in 1897 (33), this kind of microbial mat has been recorded for several geographically remote hot springs in Japan, although there have been only scattered reports of sulfur-turf microbial mats or chemotrophic sulfur streamers in geothermal springs in other countries (9, 13, 14). The sulfur-turf mats generally develop within a temperature range of 45 to 73°C, within a pH range of 6 to 9, and at discrete sulfide-oxygen interfaces in geothermal springs. These characteristics suggest that the major constituents of the sulfur-turf prokaryotic community are (hyper)thermophilic, neutrophilic, microaerophilic, and chemolithotrophic bacteria. Early studies of these sulfur-turf mats distinguished microscopically three morphotypes of bacteria, two of which were tentatively named Thiovibrio miyoshi and Thiothrix miyoshi (15). Moreover, in situ ecophysiological and microscopic studies have shown that one of these bacteria, the large sausage-shaped “Thiovibrio miyoshi,” predominates in sulfur-turf mats and oxidizes environmental sulfide to elemental sulfur and then to sulfate via thiosulfate (27–31). So far, however, it has not been possible to isolate and cultivate any thermophilic prokaryotes from the sulfur-turf mats predominated by these sausage-shaped bacteria with artificial media, and no attempt has been made to clarify their taxonomic and phylogenetic positions.Determination of 16S rRNA genes is a useful research strategy for identifying uncultivated prokaryotes and is now commonly performed in ecological studies. This technique, involving PCR amplification of 16S rRNA genes or synthesis of cDNAs from bulk 16S rRNAs of natural mixed microbial populations, has been used successfully for the phylogenetic characterization of prokaryotes in hydrothermal environments (6, 7, 34, 40, 41, 47, 48). In the present study, this approach was applied to characterize the sausage-shaped bacteria in sulfur-turf mats without isolating and cultivating them. Here we report that sulfur-turf mats contain novel thermophilic bacteria belonging to the earliest-branching lineage of the domain bacteria.     

Seasonal shifts in community composition and proteome expression in a sulphur-cycling cyanobacterial mat

Seasonal changes in light and physicochemical conditions have strong impacts on cyanobacteria, but how they affect community structure, metabolism, and biogeochemistry of cyanobacterial mats remains unclear. Light may be particularly influential for cyanobacterial mats exposed to sulphide by altering the balance of oxygenic photosynthesis and sulphide-driven anoxygenic photosynthesis. We studied temporal shifts in irradiance, water chemistry, and community structure and function of microbial mats in the Middle Island Sinkhole (MIS), where anoxic and sulphate-rich groundwater provides habitat for cyanobacteria that conduct both oxygenic and anoxygenic photosynthesis. Seasonal changes in light and groundwater chemistry were accompanied by shifts in bacterial community composition, with a succession of dominant cyanobacteria from Phormidium to Planktothrix, and an increase in diatoms, sulphur-oxidizing bacteria, and sulphate-reducing bacteria from summer to autumn. Differential abundance of cyanobacterial light-harvesting proteins likely reflects a physiological response of cyanobacteria to light level. Beggiatoa sulphur oxidation proteins were more abundant in autumn. Correlated abundances of taxa through time suggest interactions between sulphur oxidizers and sulphate reducers, sulphate reducers and heterotrophs, and cyanobacteria and heterotrophs. These results support the conclusion that seasonal change, including light availability, has a strong influence on community composition and biogeochemical cycling of sulphur and O₂ in cyanobacterial mats.     

Adaptation to Hydrogen Sulfide of Oxygenic and Anoxygenic Photosynthesis among Cyanobacteria

Four different types of adaptation to sulfide among cyanobacteria are described based on the differential toxicity to sulfide of photosystems I and II and the capacity for the induction of anoxygenic photosynthesis. Most cyanobacteria are highly sensitive to sulfide toxicity, and brief exposures to low concentrations cause complete and irreversible cessation of CO₂ photoassimilation. Resistance of photosystem II to sulfide toxicity, allowing for oxygenic photosynthesis under sulfide, is found in cyanobacteria exposed to low H₂S concentrations in various hot springs. When H₂S levels exceed 200 μM another type of adaptation involving partial induction of anoxygenic photosynthesis, operating in concert with partially inhibited oxygenic photosynthesis, is found in cyanobacterial strains isolated from both hot springs and hypersaline cyanobacterial mats. The fourth type of adaptation to sulfide is found at H₂S concentrations higher than 1 mM and involves a complete replacement of oxygenic photosynthesis by an effective sulfide-dependent, photosystem II-independent anoxygenic photosynthesis. The ecophysiology of the various sulfide-adapted cyanobacteria may point to their uniqueness within the division of cyanobacteria.     

Archaeal and bacterial diversity in two hot spring microbial mats from a geothermal region in Romania

The diversity of archaea and bacteria was investigated in two slightly alkaline, mesophilic hot springs from the Western Plain of Romania. Phylogenetic analysis showed a low diversity of Archaea, only three Euryarchaeota taxa being detected: Methanomethylovorans thermophila, Methanomassiliicoccus luminyensis and Methanococcus aeolicus. Twelve major bacterial groups were identified, both springs being dominated by Cyanobacteria, Chloroflexi and Proteobacteria. While at the phylum/class-level the microbial mats share a similar biodiversity; at the species level the geothermal springs investigated seem to be colonized by specific consortia. The dominant taxa were filamentous heterocyst-containing Fischerella, at 45 °C and non-heterocyst Leptolyngbya and Geitlerinema, at 55 °C. Other bacterial taxa (Thauera sp., Methyloversatilis universalis, Pannonibacter phragmitetus, Polymorphum gilvum, Metallibacterium sp. and Spartobacteria) were observed for the first time in association with a geothermal habitat. Based on their bacterial diversity the two mats were clustered together with other similar habitats from Europe and part of Asia, most likely the water temperature playing a major role in the formation of specific microbial communities that colonize the investigated thermal springs.     

Chemical reactivity of microbe and mineral surfaces in hydrous ferric oxide depositing hydrothermal springs

The hot springs in Yellowstone National Park, USA, provide concentrated microbial biomass and associated mineral crusts from which surface functional group (FG) concentrations and pK_a distributions can be determined. To evaluate the importance of substratum surface reactivity for solute adsorption in a natural setting, samples of iron‐rich sediment were collected from three different springs; two of the springs were acid‐sulfate‐chloride (ASC) in composition, while the third was neutral‐chloride (NC). At one of the ASC springs, mats of S^º‐rich Hydrogenobaculum‐like streamers and green Cyanidia algae were also collected for comparison to the sediment. All samples were then titrated over a pH range of 3–11, and comparisons were made between the overall FG availability and the concentration of solutes bound to the samples under natural conditions. Sediments from ASC springs were composed of hydrous ferric oxides (HFO) that displayed surface FGs typical of synthetic HFO, while sediments from the NC spring were characterized by a lower functional group density, reflected by decreased excess charge over the titration range (i.e., lower surface reactivity). The latter also showed a lower apparent point of zero charge (PZC), likely due the presence of silica (up to 78 wt. %) in association with HFO. Variations in the overall HFO surface charge are manifest in the quantities and types of solutes complexed; the NC sediments bound more cations, while the ASC sediments retained significantly more arsenic, presumably in the form of arsenate (H₂AsO₄^?). When the microbial biomass samples were analyzed, FG concentrations summed over the titratable range were found to be an order of magnitude lower for the Sº‐rich mats, relative to the algal and HFO samples that displayed similar FG concentrations on a dry weight basis. A diffuse‐layer surface complexation model was employed to further illustrate the importance of surface chemical parameters on adsorption reactions in complex natural systems.     

Jelly-like Microbial Mats over Subsurface Fields of Gas Hydrates at the St. Petersburg Methane Seep (Central Baikal)

Jelly-like microbial mat samples were collected from benthic surfaces at the St. Petersburg methane seep located in Central Baikal. The concentrations of certain ions, specifically chloride, bromide, sulphate, acetate, iron, calcium, and magnesium, were 2–40 times higher in the microbial mats than those in the pore and bottom water. A large number of diatom valves, cyanobacteria, and filamentous, rod-shaped and coccal microorganisms were found in the samples of bacterial mats using light, epifluorescence and scanning microscopy.Comparative analysis of a 16S rRNA gene fragment demonstrated the presence of bacteria and archaea belonging to the following classes and phyla: Betaproteobacteria, Gammaproteobacteria, Deltaproteobacteria, Verrucomicrobia, Cytophaga-Flavobacteria-Bacteroidetes, Cyanobacteria, and Euryarchaeota. The chemical composition and phylogenetic structure of the microbial community showed that the life activity of the mat occurs due to methane and its derivatives involved. Values of δ¹³C for the microbial mats varied from ?73.6‰ to ?65.8‰ and for animals from ?68.9‰ to ?36.6‰. Functional genes of the sequential methane oxidation (pmoA and mxaF) and different species of methanotrophic bacteria inhabiting cold ecosystems were recorded in the total DNA. Like in other psychroactive communities, the destruction of organic substances forming formed as a result of methanotrophy, terminates at the stage of acetate formation in the microbial mats of Lake Baikal (1,400 m depth). Its further transformation is limited by hydrogen content and carried out in the subsurface layers of sediments.     

Calcium dynamics in microbialite‐forming exopolymer‐rich mats on the atoll of Kiritimati,Republic of Kiribati,Central Pacific

Microbialite‐forming microbial mats in a hypersaline lake on the atoll of Kiritimati were investigated with respect to microgradients, bulk water chemistry, and microbial community composition. O₂, H₂S, and pH microgradients show patterns as commonly observed for phototrophic mats with cyanobacteria‐dominated primary production in upper layers, an intermediate purple layer with sulfide oxidation, and anaerobic bottom layers with sulfate reduction. Ca²⁺ profiles, however, measured in daylight showed an increase of Ca²⁺ with depth in the oxic zone, followed by a sharp decline and low concentrations in anaerobic mat layers. In contrast, dark measurements show a constant Ca²⁺ concentration throughout the entire measured depth. This is explained by an oxygen‐dependent heterotrophic decomposition of Ca²⁺‐binding exopolymers. Strikingly, the daylight maximum in Ca²⁺ and subsequent drop coincides with a major zone of aragonite and gypsum precipitation at the transition from the cyanobacterial layer to the purple sulfur bacterial layer. Therefore, we suggest that Ca²⁺ binding exopolymers function as Ca²⁺ shuttle by their passive downward transport through compression, triggering aragonite precipitation in the mats upon their aerobic microbial decomposition and secondary Ca²⁺ release. This precipitation is mediated by phototrophic sulfide oxidizers whose action additionally leads to the precipitation of part of the available Ca²⁺ as gypsum.     

Timing of morphological and ecological innovations in the cyanobacteria – a key to understanding the rise in atmospheric oxygen

When cyanobacteria originated and diversified, and what their ancient traits were, remain critical unresolved problems. Here, we used a phylogenomic approach to construct a well‐resolved ‘core’ cyanobacterial tree. The branching positions of four lineages (Thermosynechococcus elongatus, Synechococcus elongatus, Synechococcus PCC 7335 and Acaryochloris marina) were problematic, probably due to long branch attraction artifacts. A consensus genomic tree was used to study trait evolution using ancestral state reconstruction (ASR). The early cyanobacteria were probably unicellular, freshwater, had small cell diameters, and lacked the traits to form thick microbial mats. Relaxed molecular clock analyses suggested that early cyanobacterial lineages were restricted to freshwater ecosystems until at least 2.4 Ga, before diversifying into coastal brackish and marine environments. The resultant increases in niche space and nutrient availability, and consequent sedimentation of organic carbon into the deep oceans, would have generated large pulses of oxygen into the biosphere, possibly explaining why oxygen rose so rapidly. Rapid atmospheric oxidation could have destroyed the methane‐driven greenhouse with simultaneous drawdown in pCO₂, precipitating ‘Snowball Earth’ conditions. The traits associated with the formation of thick, laminated microbial mats (large cell diameters, filamentous growth, sheaths, motility and nitrogen fixation) were not seen until after diversification of the LPP, SPM and PNT clades, after 2.32 Ga. The appearance of these traits overlaps with a global carbon isotopic excursion between 2.2 and 2.1 Ga. Thus, a massive re‐ordering of biogeochemical cycles caused by the appearance of complex laminated microbial communities in marine environments may have caused this excursion. Finally, we show that ASR may provide an explanation for why cyanobacterial microfossils have not been observed until after 2.0 Ga, and make suggestions for how future paleobiological searches for early cyanobacteria might proceed. In summary, key evolutionary events in the microbial world may have triggered some of the key geologic upheavals on the Paleoproterozoic Earth.     

Bacterial Diversity and Functional Activity of Microbial Communities in Hot Springs of the Baikal Rift Zone

In this study the bacterial diversity of thermophilic microbial mats (40 to 65°C) in three alkaline hot springs of the Baikal Rift Zone (BRZ) was determined through pyrosequencing of 16S rRNA gene libraries. Significant diversity of bacterial species was found in the biomats of the hot springs with total number of detected phylotypes of 607. The highest share of the microbial community was represented by the phyla Chloroflexi (Seya Spring, 76.4%), Deinococcus-Thermus (Alla Spring, 45.1%), Nitrospira (Alla Spring, 36.1%), Cyanobacteria (Tsenkher Spring, 33.1%), and Proteobacteria (Tsenkher Spring, 22.6%), but their ratio varied significantly in different springs. A comparison of the biodiversity and composition of microbial communities between hot springs showed a decrease in biodiversity with increasing temperature. A large number of sequences showed a low degree of similarity with cultivated representatives in public databases. Microbial communities showed intensive rates of production and destruction of organic compounds, as revealed by the quantitative assessment of their functional activity.