Given the substrate specificity of this MCR type in utilizing butane instead of methane, and amino acid divergence of this MCR type from its methane metabolizing related counterparts, it is possible that the MCR clusters in some members of Bathyarchaeota are responsible for butane oxidation instead of methane metabolism (Laso-Prezetal.2016). They also acquired some subunits of coenzyme F420 hydrogenase; this enzyme generates reduced ferredoxin, with hydrogen as the electron donor, as an alternative to MvhADG in many Methanomicrobiales (Thaueretal.2008; Lazaretal.2016; Sousaetal.2016). A recent study found that the refractory aromatic polymer lignin stimulated the growth of Bathyarchaeota (Subgroup-8) and they incorporated CO2 as a carbon source autotrophically and utilized lignin as an energy source (Yuetal.2018). In the two recent metagenomic bathyarchaeotal binning studies, nearly all the identified bins placed H4MPT as a C1-carrier in the WoodLjungdahl pathway, which is often used by the methanogenic archaea for carbon fixation (Heetal.2016; Lazaretal.2016). Bathyarchaeota The discovery of BchG of archaeal origin in the genomic content of Bathyarchaeota also suggests that an archaeon-based photosynthetic pathway might exist in nature, and that photosynthesis might have evolved before the divergence of bacteria and archaea (Mengetal.2009). In contrast, Subgroup-15 (Crenarchaeota group C3) organisms dominate cDNA libraries from all sediment layers, albeit with minor contribution to the corresponding DNA libraries; this indicates that this group is metabolically active in the benthic euxinic, organic-rich sediments of karstic lakes (Filloletal.2015). The product, acetate, would then be used by acetate-consuming SRB to benefit the thermodynamic efficiency of AOM. Following the four treatments, the viable bathyarchaeotal communities mainly comprised Subgroups-4 and -8, thus indicating that these two subgroups could tolerate the initial aerobic conditions (Gagenetal.2013). Bathyarchaeota are believed to have roles in the carbon cycle in marine systems. Kuboetal. BA1 (Subgroup-3) genome contains many genes of the reductive acetyl-CoA (WoodLjungdahl) pathway and key genes of the methane metabolism pathway. On the other hand, the proportion of bathyarchaeotal sequence in the total archaeal community sequence increases with depth, and they may favor anoxic benthic sediments with iron-reducing conditions. Within Bathyarchaeota, the sequences were classified into six subclades according to . All sequences were clustered at 90% identity using Usearch v10.0.240 (https://www.drive5.com/usearch/), then the 16S rRNA gene sequences from available bathyarchaeotal genomes in public database, the anchor sequences from Kuboetal. In addition, the catalyzed reporter deposition-fluorescent in situ hybridization (CARD-FISH) studies for the detection and quantification of bathyarchaeotal cells suggest that they are abundant in the center and marine invertebrate-inhabited layers in the Haakon Mosby Mud Volcano, and in the marine subsurface sediments in the Equatorial ODP site 1125 and Peru Basin ODP site 1231 (Kuboetal.2012). Subsequent heterologous expression of bathyarchaeotal Ack revealed that the enzyme can catalyze the biochemical reaction in the direction from acetyl phosphate to acetate, with a higher affinity for the substrates than the products (Heetal.2016). Archaea In a recent global evaluation of the archaeal clone libraries from various terrestrial environmental settings, permutational analysis that tested the relationship between Bathyarchaeota and environmental factors suggested that salinity, total organic carbon and temperature are the most influential factors impacting community distribution across different terrestrial habitats (Xiangetal.2017). It has been suggested that Bathyarchaeota is one of the cosmopolitan groups frequently detected in the freshwater and marine sediments (68% of all sediments analyzed), accounting for a large proportion of the sediment microbial communities (average 36 22%) (Filloletal.2016). A model based on the thermodynamic considerations of chemicals and temperatures may be used to offer a framework linking the distribution of microbial groups and energy landscapes (Amendetal.2011; LaRowe and Amend 2014; Dahleetal.2015). BATHYARCHAEOTA OCCURRENCE IN SHALLOW MARINE With respect to its function, the protein might be responsible for photosynthesis in archaea; this suggests that photosynthesis may have evolved before the divergence of the bacteria and archaea domains (Mengetal.2009; Lietal.2012). The assignment of bathyarchaeotal subgroups was made based on either having been formerly defined or being monophyletic, using both distance and maximum-likelihood estimations (Kuboetal.2012). This suggests that methane metabolism might have evolved before the divergence of the ancient archaeal lineages of Bathyarchaeota and Euryarchaeota, in agreement with the assumption that methanogenesis might represent one of the earliest metabolic transformations (Battistuzzi, Feijao and Hedges 2004; Ferry and House 2006; Evansetal.2015; Lloyd 2015). Further membrane lipid characterization of enriched or pure bathyarchaeotal cultures will help to validate this discovery. Schematic figure representing major eco-niches of Bathyarchaeota. It was proposed that the high diversity of Bathyarchaeota implies a high metabolic diversity among its subgroups (Kuboetal.2012). Archaebacteria Facts - Softschools.com Biddle JF, Fitz-Gibbon S, Schuster SC et al. The central product, acetyl-CoA, would either (i) be involved in substrate-level phosphorylation to generate acetate and ATP, catalyzed by an ADP-forming acetyl-CoA synthase as in other peptide-degrading archaea; (ii) be metabolized to generate acetate through the Pta-Ack pathway, similarly to bona fide bacterial homoacetogens; or (iii) be utilized for biosynthesis, e.g. According to the meta-analysis of archaeal sequences available in the ARB SILVA database (Kuboetal.2012), Bathyarchaeota was further recognized as a group of global generalists dwelling in various environments, including marine sediments, hydrothermal vents, tidal flat and estuary sediments, hypersaline sediments, terrestrial subsurface, biomats, limnic water and sediments, underground aquifers, hot springs, soils, municipal wastewaters, animal digestive tract, etc. The Distribution of Bathyarchaeota in Surface Sediments Bathyarchaeota, formerly known as the Miscellaneous Crenarchaeotal Group, is a phylum of global generalists that are widespread in anoxic sediments, which host relatively high abundance archaeal communities. Other archaeal groups are also commonly detected in estuaries worldwide. Based on the lineage distribution pattern analysis of the archaeal community of seven major eco-niches, it is also evident that the different evolutionary lineages are habitat-specific, and salinity rather than temperature is the primary driving force of the variation of global archaea distribution, with a similar pattern also evident for the global bacterial distribution (Lozupone and Knight 2007; Auguet, Barberan and Casamayor 2010). Bathyarchaeota, a recently proposed archaeal phylum, is globally distributed and highly abundant in anoxic sediments. WebEtymology: Gr. It is known that a methane microbiome can be established in methane seeps sites; however, they are still poorly characterised. The phylum Bathyarchaeota, which has high species and functional diversity, is abundant and widespread in marine sediments. Jacquemet A, Barbeau J, Lemiegre L et al. Microbial communities of deep marine subsurface sediments: molecular and cultivation surveys, Methanogenic archaea: ecologically relevant differences in energy conservation, Methylotrophic methanogenesis discovered in the archaeal phylum, Methanotrophic archaea possessing diverging methane-oxidizing and electron-transporting pathways, Prokaryotic community composition and biogeochemical processes in deep subseafloor sediments from the Peru Margin, Prokaryotic functional diversity in different biogeochemical depth zones in tidal sediments of ?the Severn Estuary, UK, revealed by stable-isotope probing, Enrichment and cultivation of prokaryotes associated with the sulphate-methane transition zone of diffusion-controlled sediments of Aarhus Bay, Denmark, under heterotrophic conditions, The physiology and habitat of the last universal common ancestor, Distribution of Bathyarchaeota communities across different terrestrial settings and their potential ecological functions, Uniting the classification of cultured and uncultured bacteria and archaea using 16S rRNA gene sequences, A large-scale evaluation of algorithms to calculate average nucleotide identity, High occurrence of Bathyarchaeota (MCG) in the deep-sea sediments of South China Sea quantified using newly designed PCR primers, Growth of sedimentary Bathyarchaeota on lignin as an energy source, Genomic and transcriptomic evidence for carbohydrate consumption among microorganisms in a cold seep brine pool, This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (, Illuminating the Oral Microbiome and its Host Interactions: Animal models of disease, Engineering lanthipeptides by introducing a large variety of RiPP modifications to obtain new-to-nature bioactive peptides, Meat fermentation at a crossroads: where the age-old interplay of human, animal, and microbial diversity and contemporary markets meet, Incorporation, fate, and turnover of free fatty acids in cyanobacteria, Ruminococcus gnavus: friend or foe for human health, About the Federation of European Microbiological Societies, GLOBAL DISTRIBUTION AND HIGH DIVERSITY OF BATHYARCHAEOTA, DISTRIBUTION PATTERN AND MOLECULAR DETECTION, PHYSIOLOGICAL AND GENOMIC CHARACTERIZATION, ECOLOGICAL FUNCTIONS AND EVOLUTION OF BATHYARCHAEOTA, https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model, Receive exclusive offers and updates from Oxford Academic, Copyright 2023 Federation of European Microbiological Societies. The members of Bathyarchaeota were positively and strongly correlated especially with the acetoclastic Methanosaeta; however, the second most abundant archaeal group, MG-I (subordinate to Thaumarchaeota) is negatively correlated with other groups, probably indicating segregation corresponding to two distinct lifestyles in this case (Liuetal.2014). Furthermore, one new subgroup (Subgroup-23) was proposed in this study (Fig. Reconsideration of the potential methane-oxidizing contribution of Bathyarchaeota would refine the congruency between the predicted and observed microbial communities, i.e. Four genomes (Subgroups-1, -6, -7 and -15) were recovered from the sediment metagenome. 2. The knowledge of their physiological and genomic properties, as well as their adaptive strategies in various eco-niches, is nonetheless still rudimentary. Genomic and enzymatic evidence for acetogenesis among These archaeal groups are the phylogenetically closest ones to the protoeukaryote that served as the mitochondrion-acquiring host; this gave rise to a hydrogen hypothesis that explains their hydrogen-dependent metabolism to address the mitochondrion acquisition and subsequent endosymbiont processes. In summary, the most recent research advances have considerably expanded our knowledge of Bathyarchaeota, their distribution, ecology and physiological and genomic properties since their first discovery and definition about two decades ago. However, after allowing for a single nucleotide mismatch, the coverage efficiency markedly increased, to around 8090%. A meta-analysis of the distribution of sediment archaeal communities towards environmental eco-factors (7098 archaeal operational taxonomic units from 207 sediment sites worldwide) was performed and a multivariate regression tree was constructed to depict the relationship between archaeal lineages and the environmental origin matrix (Filloletal.2016). Bathyarchaeota is of great interest to microbial ecologists for its wide distribution, high abundance, and diversity, as well as its potential ability to degrade detrital organic matter in aquatic environments and drive global elements cycling . Methanogenesis and acetogenesis are considered to be the two most fundamental and ancient microbial biochemical energy conservation processes, and they both employ the WoodLjungdahl pathway for CO2 reduction and ATP generation (Weissetal.2016). Furthermore, the phylogeny of concatenated alignments constituting 12 ribosomal proteins obtained from currently available bathyarchaeotal genomes (from GenBank, 29 November 2017 updated) was also reconstructed, which showed a similar topology to those of 16S rRNA genes with a few exceptions in Subgroup-17 (Fig. For instance, a study into the stratification of the archaeal community from a shallow sediment in the Pearl River Estuary defined bathyarchaeotal subgroups from MCG-A to -F (Jiangetal.2011), including the NT-A3 group, which is predominantly isolated from the hydrate stability zone in the deep subsurface hydrate-bearing marine sediment core in the Nankai Trough (Reedetal.2002); meanwhile, an investigation of archaeal composition in ca 200 m deep sub-seafloor sediment cores at the offshore Peru Margin ODP sites 1228 and 1229 listed Bathyarchaeota subgroups PM-1 to -8 (Websteretal.2006). This group of lipids has not been found in natural environments or microorganism enrichments dominated by methanotrophic archaea before (Rosseletal.2008; Kellermannetal.2012), nor have they been detected after re-analyzing lipid extracts from the above two studies using the same method in the study (Meadoretal.2015). Genes responsible for the dissimilatory nitrite reduction to ammonium (nirB and nrfD) were identified in Subgroups-1, -17 (formally Subgroup-7/17), -6 and -15, respectively, suggesting the potential existence of a respiratory pathway involving nitrite reduction (Lazaretal.2016). Phylogenetic tree of bathyarchaeotal 16S rRNA genes. The three methods described above may be used for the quantification of bathyarchaeotal abundance based on DNA and RNA targets. Combined with the large amount of carbon deposited in the subseafloor (ca 15 1021 g) (Fryetal.2008), the high abundance of MCG archaea in marine sediments (10100% of total archaeal abundance) (Parkesetal.2005; Biddleetal.2006; Fryetal.2008; Kuboetal.2012; Lloydetal.2013) and their heterotrophic properties on detrital proteins, acetate, aromatic compounds and/or other organic substrates (Biddleetal.2006; Websteretal.2010; Websteretal.2011; Lloydetal.2013; Naetal.2015), naturally led to the proposal that this group of archaea may play an important role in global carbon biogeochemical cycling (Kuboetal.2012; Lloydetal.2013; Filloletal.2016; Heetal.2016). More importantly, the first-ever bacteriochlorophyll a synthase (BchG) of archaeal origin was identified in the archaeal portion of the genomic fragment, and its function confirmed by producing BchG in a heterologous expression system (Mengetal.2009). These indicative subgroups are the dominant ones in the environment, as evaluated by relatively abundant fraction of Bathyarchaeota in corresponding archaeal communities (on average 44% among all studies). adj. Species abundance distribution analysis indicates that Bathyarchaeota is one of the persistent and abundant core lineages of the sediment archaeal communities, showing, to some extent, habitat-specific distribution (Filloletal.2016). 4), although these might not necessarily exist in all bathyarchaeotal subgroups (Fig. This could be explained by the versatile pathways of organic matter assimilation present in the majority of Bathyarchaeota, reflected by inferences from genomic data. Frontiers | The Distribution of Bathyarchaeota in Surface 1) (Heetal.2016; Lazaretal.2016). No methane metabolism genes were recovered from bathyarchaeotal genomic bins or any contigs from the WOR estuarine sediments, in contrast to an earlier study (Evansetal.2015). Further, a close co-occurrence of Bathyarchaeota and Methanomicrobia hinted at a syntrophic association between them; the acetate production/consumption relationship between the two might be responsible for such a scenario, as proposed by metabolic predictions (Heetal.2016; Xiangetal.2017). The Subgroups-1, -6 and -15 genomes also encoded the methyl glyoxylate pathway, which is typically activated when slow-growing cells are exposed to an increased supply of sugar phosphates (Weber, Kayser and Rinas 2005). This will have a profound impact not only on deciphering the metabolic properties of Bathyarchaeota, by using butanetriol dibiphytanyl glycerol tetraethers as biomarkers to trace carbon acquisition by isotopic labeling, but also by representing their pivotal contribution, associated with their global abundance, to biogeochemical carbon cycling on a large ecological scale. The diversity of bathyarchaeotal community turns out to be similar in the four cultivation treatments (basal medium, addition of an amino acid mix, H2-CO2 headspace and initial aerobic treatment). Both Bathyarchaeota and the recently identified more basally branched Lokiarchaeota acquired the H4MPT-dependent WoodLjungdahl pathway and the hydrogen-dependent electron bifurcating system MvhADG-HdrABC, viewed as typical for the anaerobic and hydrogen-dependent archaeal lifestyle (Lazaretal.2016; Sousaetal.2016). Draft Genome Sequence of " Candidatus Results In the current study, nine To alleviate the nomenclature confusion, we constructed an updated RAxML tree (Fig. To avoid the confusion, Subgroups-18 and -19 were named to be consistent with subgroups MCG-18 and MCG-19 as proposed in two previous reports (respectively Lazaretal.2015; Filloletal.2016), while Subgroup-20 was renamed to replace the subgroup MCG-19 in Fillol et al.s tree (Filloletal.2016). the potential AOM metabolism of Bathyarchaeota in the flange of the hydrothermal vent would be consistent with the aforementioned genomic inferences (Evansetal.2015). S. butanivorans protein extracts; they are probably responsible for the initial step of butane activation to generate butyl-CoM. The primer pair MCG242dF/MCG528R may potentially be used for the determination of the bathyarchaeotal community abundance, with relatively high subgroup coverage and specificity in silico; however, experimental tests are needed to confirm this. WebHost. The indicator subgroups in saline and freshwater sediments were depicted accordingly. 1 and Table S 5 ), and the average proportion of Bathyarchaeota in the mangrove sediments (43.32%, sd = 0.106) was significantly higher than that in the mud flat sediments (36.47%, sd = 0.084) ( p < It is evident that the phylogenetically diverse subgroups are heterotrophs with metabolism centralized around acetyl-CoA generation. Later on, members of Bathyarchaeota were also found to be abundant in deep marine subsurface sediments (Reedetal.2002; Inagakietal.2003), suggesting that this group of archaea is not restricted to terrestrial environments, and the name has been changed to MCG archaea (Inagakietal.2003). Archaea Definition & Meaning - Merriam-Webster In the case of Subgroup-15, which branched away from other groups, MCG242dF would be associated with a relatively low coverage efficiency in the absence of nucleotide mismatches, but high (above 80%) coverage efficiency with 1 or 2 nucleotide mismatches; similarly, MCG678R would be associated with a limited coverage efficiency in the absence of nucleotide mismatches, but the coverage efficiency increases considerably with 1 or 2 nucleotide mismatches. Barns SM, Delwiche CF, Palmer JD et al. They include Euryarchaeota, and members of the DPANN and Asgard archaea. A subsequent heterologous expression and activity assays of the bathyarchaeotal acetate kinase gene ack demonstrated the ability of these bathyarchaeotal members to grow as acetogens. Our results provide an overview of the archaeal population, Acetyl-CoA might be involved in acetate generation in a fermentative pathway; however, genomic evidence suggests that Subgroup-1 cells might rely on both fermentative and respiratory metabolism (a simple respiratory metabolism based on a membrane-bound hydrogenase). Hlne A, Mylne H, Christine D et al. Recent data point to the global occurrence of Bathyarchaeota and their potential impact on global carbon transformation, highlighting their important role as a group of global generalists participating in carbon cycling, similar to euryarchaeotal methanogens and Thaumarchaeota. Boetius A, Ravenschlag K, Schubert CJ et al. The distinct bathyarchaeotal subgroups diverged to adapt to marine and freshwater environments. However, in the above binning studies, none of the genomes encoded enzymes involved in the final methane production step (McrABG), suggesting that the WoodLjungdahl pathway is not used for methane production but for acetyl-CoA generation and further acetogenesis. Given that they are abundant, globally distributed and phylogenetically diverse, continued exploration of new potential bathyarchaeotal subgroups is encouraged. Evans PN, Parks DH, Chadwick GL et al. PubChem BioAssay. Similarly, rRNA slot blot hybridization indicates the existence of functionally active Bathyarchaeota not only in the surface and subsurface sediments from the Nyegga site 272-02, Cascadia Margin, Gulf of Mexico, Hydrate Ridge ODP site 1245 and Janssand (North Sea), but also in the oxic mats in the Arabian Gulf and subsurface White Oak River sediments (Kuboetal.2012). (C) The metabolic properties of 24 bathyarchaeotal genomes. In some flange subsamples, Bathyarchaeota were even more dominant than ANME; however, compared with the well-studied metabolism of ANME, the exact function of Bathyarchaeota in that ecological setting remains unknown. It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide, This PDF is available to Subscribers Only. Single amplified genomes (SAGs) of a Subgroup-15 bathyarchaeotal member from the Aarhus Bay sediments harbor genes for predicted extracellular protein degrading enzymes, such as clostripain (Lloydetal.2013). Genome labels are according to panel (B). It is one of the predominant groups in the marine subsurface archaeal community (Fryetal.2008; Teske and Srensen 2008; Lloydetal.2013). We also highlighted the unique genomic features and potential adaptation strategies of estuarine archaea, pointing out major unknowns in the field and scope for future research. It was proposed that reduced ferredoxin generated by peptide and/or glucose might be used for the reduction of methyl groups on methylated compounds to subsequently generate methane (Evansetal.2015). Taxonomy browser (Candidatus Bathyarchaeota) - National Community, Distribution, and Ecological Roles of Estuarine Archaea Vanwonterghem I, Evans PN, Parks DH et al. [43] (Figure 4). Lomstein BA, Langerhuus AT, DHondt S et al. Primers and probes for molecular detection and quantification of Bathyarchaeota subgroups. Open reading frames encoded by the three fosmid clones comprised genes related to lipid biosynthesis, energy metabolism and resistance to oxidants. Furthermore, evidence of fatty acid and aromatic compound utilization by Bathyarchaeota has been presented (Mengetal.2014; Evansetal.2015; Heetal.2016); these transformations would be supported by the beta-oxidation pathway and a potential anaerobic aromatic compound degradation pathway. Uncultured archaea in deep marine subsurface sediments: have we caught them all? Furthermore, another study demonstrated that the archaeal communities of the sulfatemethane transition zone at diffusion-controlled sediments of Aarhus Bay (Denmark) contain considerable amounts of Bathyarchaeota; the overall archaeal community structure did not change greatly during the experimentits diversity was lower after 6 months of incubation under heterotrophic conditions, with periodic modest sulfate and acetate additions (Websteretal.2011). As suggested by the classification of uncultured archaea based on nearly full-length 16S rRNA gene sequences, the bathyarchaeotal sequence boundary falls into the minimum sequence identity range of phylum level (74.9579.9%), and each subgroup generally falls into the median sequence identity range of family and order levels (91.6592.9% and 88.2590.1%, respectively) (Yarzaetal.2014). The marine/freshwater segregation is a distribution pattern widely shared by diverse microorganisms, including archaea, bacteria, viruses and eukaryotes (Logaresetal.2009). Lineage (full): cellular organisms; Archaea; TACK group. neut. Bathyarchaeota, a recently proposed archaeal phylum, is globally distributed and highly abundant in anoxic sediments. Core Metabolic Features and Hot Origin of Bathyarchaeota Bathyarchaeota, formerly known as the Miscellaneous Crenarchaeotal Group, is a phylum of global generalists that are widespread in anoxic sediments, which host relatively high abundance archaeal communities. Characteristics of the Bathyarchaeota community in During the enriching process with lignin addition, the Subgroup-8 abundance climbed over 10 times compared with the initial stage and became the most dominant archaeal species. A complete set of active sites and signal sequences for extracellular transport is also encoded by bathyarchaeotal SAGs (Lloydetal.2013). The in silico tests revealed that primers MCG528, MCG493, MCG528 and MCG732 cover 87, 79, 44 and 27% of sequences of Subgroups-1 to -12 on average, respectively. It has been proposed that the deduced last common ancestor was most likely a saline-adapted organism, and the evolutionary progression occurred most likely in the saline-to-freshwater direction, with few environmental transitional events. Rossel PE, Lipp JS, Fredricks HF et al. Given the diverse and complex phylogeny of the Bathyarchaeota (Kuboetal.2012; Filloletal.2016), the occurrence of commonly shared physiological and metabolic properties in different lineages seems unlikely, with the evolutionary diversification of bathyarchaeotal lineages largely driven by the adaptation to various environmental conditions and available carbon and energy sources, etc. The deduced last common ancestor of Bathyarchaeota might be a saline-adapted organism, which evolved from saline to freshwater habitats during the diversification process, with the occurrence of few environmental transitional events. Metabolic pathways of the Genomic characterization and metabolic potentials of Bathyarchaeota. In this process, methane is not assimilated by Bathyarchaeota but serves as an energy source. Diversity, metabolism and cultivation of archaea In summary, there are a total of 25 subgroups of Bathyarchaeota based on all available 16S rRNA gene sequences at this moment, and the former names for each subgroup are also labeled in the tree (Fig. Taxonomic classification revealed that between 0.1 and 2% of all classified sequences were assigned to Bathyarchaeota. Until now, 2). Third, only limited reports on the distribution patterns of bathyarchaeotal subgroups and the associated environmental factors are available. Individual metagenome assemblies The concatenated ribosomal protein (RP) alignment contained 12 RPs, and those genomes with <25% RPs were excluded from tree construction. Among the presently recognized 25 bathyarchaeotal subgroups, eight are delineated as significantly niche-specific based on their marine/freshwater segregation. Because of the high diversity of Bathyarchaeota and various independent analyses of samples from diverse environments, the nomenclature for this archaeal group in previous reports was very complex. Obtaining direct physiological evidence for the generation or oxidization of methane by Bathyarchaeota in the future is also important. Bathyarchaeia occurrence in rich methane sediments On the other hand, because of the bidirectionality of these enzymes in methane metabolism (Boetiusetal.2000; Knittel and Boetius 2009), it is still possible that some members of Bathyarchaeota are involved in anaerobic methane oxidation.
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