Juan De Fuca Ridge Flank Research Articles

Investigating carbon metabolisms in the marine subsurface environment of the Juan de Fuca Ridge Flank

Microorganisms are the most diverse and abundant life on earth. They are responsible for recycling the elements of our planet and keeping our world habitable. Despite their importance, genome‐based analyses estimate that 90% of the microorganisms on our planet have not been cultured for laboratory investigations. The crustal biosphere of the Juan de Fuca Ridge Flank is a warm, subsurface environment, which has been shown to contain many uncultured organisms. Some of these uncultured groups harbor functional genes of evolutionary and biogeochemical significance, suggesting that further study will increase our knowledge of microbial survival on early Earth, as well as in this hydrothermal, nutrient‐limited crustal environment. In previous work the recovery of mono‐ and bifunctional carbon monoxide dehydrogenase genes suggested that carboxydotrophy may be a viable lifestyle in this environment. In May of 2019, the RV Atlantis traveled to the Juan de Fuca Ridge Flank to test this hypothesis. Equipped with custom fluid samplers the ROV Jason‐II was used to collect pristine crustal fluids from subsurface borehole observatories for chemical analyses and microbial culturing efforts. Our main objective was to measure potential carbon substrates, including carbon monoxide and methane, and to cultivate unique organisms from this environment to better understand carbon‐related metabolisms in this important but understudied aquifer.

Dissolved organic carbon in basalt-hosted deep subseafloor fluids of the Juan de Fuca Ridge flank

Marine dissolved organic carbon (DOC) is highly depleted in radiocarbon and thus inferred to be largely refractory to removal processes that operate on less than millennial timescales. However, a growing number of reports have shown that a large fraction of marine DOC can be effectively removed during circulation through submarine hydrothermal systems. What is not clear, however, is whether the DOC that remains in hydrothermal fluids is remnant non-reactive DOC from recharged seawater, or DOC that has been largely produced in the subsurface. We collected and characterized warm (∼65°C) hydrothermal fluids from deep (18, 40, 73, 200 m) within the basalt-hosted basement of the Juan de Fuca Ridge flank in the Northeast Pacific Ocean. DOC concentrations in hydrothermal fluids were 9 to 18 μM, much lower than measured in local deep seawater (37.5 μM). DOC Δ14C values of −683‰ to −856‰ were much lower than the Δ14C-dissolved inorganic carbon (DIC) values of −880‰ to −918‰, while DOC δ13C values of −23.6‰ to −27.0‰ were much heavier than that of the particulate organic carbon (POC) pool (∼−34‰), suggesting that biological production in the subsurface is not a primary source of DOC. Rather, our data suggest that isotopically enriched DOC are selectively removed from recharged seawater, leaving DOC that is very isotopically depleted in the basaltic basement fluids. Despite the removal of 50–75% of DOC in the subsurface, nuclear magnetic resonance (NMR) functional group analyses indicate that aromatic compounds were added to basaltic basement fluids during passage through the deep subseafloor and may partly contribute to the depleted 14C DOC in the ridge-flank basement fluids.

Carboxydotrophy potential of uncultivated Hydrothermarchaeota from the subseafloor crustal biosphere

The exploration of Earth’s terrestrial subsurface biosphere has led to the discovery of several new archaeal lineages of evolutionary significance. Similarly, the deep subseafloor crustal biosphere also harbors many unique, uncultured archaeal taxa, including those belonging to Candidatus Hydrothermarchaeota, formerly known as Marine Benthic Group-E. Recently, Hydrothermarchaeota was identified as an abundant lineage of Juan de Fuca Ridge flank crustal fluids, suggesting its adaptation to this extreme environment. Through the investigation of single-cell and metagenome-assembled genomes, we provide insight into the lineage’s evolutionary history and metabolic potential. Phylogenomic analysis reveals the Hydrothermarchaeota to be an early-branching archaeal phylum, branching between the superphylum DPANN, Euryarchaeota, and Asgard lineages. Hydrothermarchaeota genomes suggest a potential for dissimilative and assimilative carbon monoxide oxidation (carboxydotrophy), as well as sulfate and nitrate reduction. There is also a prevalence of chemotaxis and motility genes, indicating adaptive strategies for this nutrient-limited fluid-rock environment. These findings provide the first genomic interpretations of the Hydrothermarchaeota phylum and highlight the anoxic, hot, deep marine crustal biosphere as an important habitat for understanding the evolution of early life.

Divergent methyl-coenzyme M reductase genes in a deep-subseafloor Archaeoglobi

The methyl-coenzyme M reductase (MCR) complex is a key enzyme in archaeal methane generation and has recently been proposed to also be involved in the oxidation of short-chain hydrocarbons including methane, butane, and potentially propane. The number of archaeal clades encoding the MCR continues to grow, suggesting that this complex was inherited from an ancient ancestor, or has undergone extensive horizontal gene transfer. Expanding the representation of MCR-encoding lineages through metagenomic approaches will help resolve the evolutionary history of this complex. Here, a near-complete Archaeoglobi metagenome-assembled genome (MAG; Ca. Polytropus marinifundus gen. nov. sp. nov.) was recovered from the deep subseafloor along the Juan de Fuca Ridge flank that encodes two divergent McrABG operons similar to those found in Ca. Bathyarchaeota and Ca. Syntrophoarchaeum MAGs. Ca. P. marinifundus is basal to members of the class Archaeoglobi, and encodes the genes for β-oxidation, potentially allowing an alkanotrophic metabolism similar to that proposed for Ca. Syntrophoarchaeum. Ca. P. marinifundus also encodes a respiratory electron transport chain that can potentially utilize nitrate, iron, and sulfur compounds as electron acceptors. Phylogenetic analysis suggests that the Ca. P. marinifundus MCR operons were horizontally transferred, changing our understanding of the evolution and distribution of this complex in the Archaea.

Strong impact of ion filtration on the isotopic composition of chlorine in young clay-rich oceanic sediment pore fluids

A collection of 34 pore fluids from young clay-rich sedimentary piles of the Shikoku basin on the Philippine oceanic plate at the front of the Nankai subduction zone, southeast of Japan, were analyzed for their chloride content and chlorine stable isotope composition (35Cl and 37Cl, reported as δ37Cl). These pore fluids are samples from IODP Sites C0011 and C0012, which penetrate deeper than 500 meters below the seafloor. At both sites, the δ37Cl values display a regular decrease with depth from about 0‰ (seawater value) at the seafloor to −8.5 and −6.5‰ respectively in hydrogeologically isolated, low permeability, sediment levels. At some depths, local deviations of up to 6‰ in δ37Cl interrupt the overall decreasing trends. They may be interpreted as the injection of external fluids into the sedimentary piles as a result of sedimentary levels with higher permeability. At the bottom of the sedimentary piles, the injected fluids, probably originating from the altered oceanic crust, systematically have δ37Cl values closer to seawater. Literature data indicate the display of downward trends of decreasing δ37Cl values of the pore fluids in many other young clay-rich sedimentary piles (Nankai Trough and the Japan Trench accretionary prisms, Black Ridge and Juan de Fuca Ridge flank). Here, we use conservation equations to demonstrate that these δ37Cl depth profiles can be modeled using the effect of the compaction of growing clay–rich sedimentary piles on the water and chlorides (including 35Cl and 37Cl isotopes). This modeling shows that a small isotope fractionation of chlorine isotopes, α37Cl/35Clexpulsed fluid-residual fluid, in the range of 1.006 to 1.001, may explain the observed negative δ37Cl chlorides at the bottom of the sedimentary piles. The complementary positive δ37Cl chlorides must accumulate at the top but are very likely erased by dilution with seawater chlorides across the seafloor interface. This range of chlorine isotope fractionation values is in agreement with the theory of ion filtration of Phillips and Bentley (1987) in which the mobility of chlorides through semi-permeable clay membrane is determined by ion repulsion. Based on this study, we show that young pore fluids in clay-rich sediments of the oceanic crust are 37Cl-depleted chloride reservoirs. They might be the source of 37Cl-depleted chloride fluids advected at mud volcanoes on the seafloor but do not seem to contribute largely to subduction zone magmatic, tectonic, metamorphic products where no such 37Cl-depleted chlorine is documented. This in turn is a strong argument to propose that most of the clay-rich sediment’s pore waters are released back into the ocean rather than subducted.

Influence of Igneous Basement on Deep Sediment Microbial Diversity on the Eastern Juan de Fuca Ridge Flank.

Microbial communities living in deeply buried sediment may be adapted to long-term energy limitation as they are removed from new detrital energy inputs for thousands to millions of years. However, sediment layers near the underlying oceanic crust may receive inputs from below that influence microbial community structure and/or activity. As part of the Census of Deep Life, we used 16S rRNA gene tag pyrosequencing on DNA extracted from a spectrum of deep sediment-basement interface samples from the subsurface of the Juan de Fuca Ridge flank (collected on IODP Expedition 327) to examine this possible basement influence on deep sediment communities. This area experiences rapid sedimentation, with an underlying basaltic crust that hosts a dynamic flux of hydrothermal fluids that diffuse into the sediment. Chloroflexi sequences dominated tag libraries in all sediment samples, with variation in the abundance of other bacterial groups (e.g., Actinobacteria, Aerophobetes, Atribacteria, Planctomycetes, and Nitrospirae). These variations occur in relation to the type of sediment (clays versus carbonate-rich) and the depth of sample origin, and show no clear connection to the distance from the discharge outcrop or to basement fluid microbial communities. Actinobacteria-related sequences dominated the basalt libraries, but these should be viewed cautiously due to possibilities for imprinting from contamination. Our results indicate that proximity to basement or areas of seawater recharge is not a primary driver of microbial community composition in basal sediment, even though fluids diffusing from basement into sediment may stimulate microbial activity.

Atribacteria from the Subseafloor Sedimentary Biosphere Disperse to the Hydrosphere through Submarine Mud Volcanoes.

Submarine mud volcanoes (SMVs) are formed by muddy sediments and breccias extruded to the seafloor from a source in the deep subseafloor and are characterized by the discharge of methane and other hydrocarbon gasses and deep-sourced fluids into the overlying seawater. Although SMVs act as a natural pipeline connecting the Earth’s surface and subsurface biospheres, the dispersal of deep-biosphere microorganisms and their ecological roles remain largely unknown. In this study, we investigated the microbial communities in sediment and overlying seawater at two SMVs located on the Ryukyu Trench off Tanegashima Island, southern Japan. The microbial communities in mud volcano sediments were generally distinct from those in the overlying seawaters and in the well-stratified Pacific margin sediments collected at the Peru Margin, the Juan de Fuca Ridge flank off Oregon, and offshore of Shimokita Peninsula, northeastern Japan. Nevertheless, in-depth analysis of different taxonomic groups at the sub-species level revealed that the taxon affiliated with Atribacteria, heterotrophic anaerobic bacteria that typically occur in organic-rich anoxic subseafloor sediments, were commonly found not only in SMV sediments but also in the overlying seawater. We designed a new oligonucleotide probe for detecting Atribacteria using the catalyzed reporter deposition-fluorescence in situ hybridization (CARD-FISH). CARD-FISH, digital PCR and sequencing analysis of 16S rRNA genes consistently showed that Atribacteria are abundant in the methane plumes of the two SMVs (0.58 and 1.5 × 104 cells/mL, respectively) but not in surrounding waters, suggesting that microbial cells in subseafloor sediments are dispersed as “deep-biosphere seeds” into the ocean. These findings may have important implications for the microbial transmigration between the deep subseafloor biosphere and the hydrosphere.

Genomic comparisons of a bacterial lineage that inhabits both marine and terrestrial deep subsurface systems

It is generally accepted that diverse, poorly characterized microorganisms reside deep within Earth’s crust. One such lineage of deep subsurface-dwelling bacteria is an uncultivated member of the Firmicutes phylum that can dominate molecular surveys from both marine and continental rock fracture fluids, sometimes forming the sole member of a single-species microbiome. Here, we reconstructed a genome from basalt-hosted fluids of the deep subseafloor along the eastern Juan de Fuca Ridge flank and used a phylogenomic analysis to show that, despite vast differences in geographic origin and habitat, it forms a monophyletic clade with the terrestrial deep subsurface genome of “Candidatus Desulforudis audaxviator” MP104C. While a limited number of differences were observed between the marine genome of “Candidatus Desulfopertinax cowenii” modA32 and its terrestrial relative that may be of potential adaptive importance, here it is revealed that the two are remarkably similar thermophiles possessing the genetic capacity for motility, sporulation, hydrogenotrophy, chemoorganotrophy, dissimilatory sulfate reduction, and the ability to fix inorganic carbon via the Wood-Ljungdahl pathway for chemoautotrophic growth. Our results provide insights into the genetic repertoire within marine and terrestrial members of a bacterial lineage that is widespread in the global deep subsurface biosphere, and provides a natural means to investigate adaptations specific to these two environments.

Seafloor tilt induced by ocean tidal loading inferred from broadband seismometer data from the Cascadia subduction zone and Juan de Fuca Ridge

Mass-balancing voltages from four buried broadband seismometers connected to the NEPTUNE Canada seafloor cable are being recorded at 24-bit resolution. Sites are located on the Vancouver Island continental shelf, the nearby Cascadia accretionary prism, the eastern flank of the Juan de Fuca Ridge, and the western flank close to the Juan de Fuca Ridge axis. Tidal variations are present throughout the records. Variations in vertical acceleration at three of the sites match predicted gravitational attraction variations very well; those at the fourth site show a small residual that is probably caused by sensitivity to tilt resulting from sensor inclination. Horizontal accelerations, which at tidal periods are sensitive primarily to tilt, are anomalously large relative to standard-earth model results. After removal of predicted tidal body and ocean attraction and loading terms, the residuals are seen to follow ocean pressure variations. Responses range from 0.4 μrad dbar−1 (0.04 μrad kPa−1) at 10° true (down under positive load) at the continental shelf site, to 0.6 μrad dbar−1 at 243° at the Cascadia prism, 0.4 μrad dbar−1 at 90° at the eastern Juan de Fuca Ridge flank, and 0.2 μrad dbar−1 at 116° true on the western ridge flank. Except at the continental shelf site, tilts are roughly perpendicular to structural strike. The tilt observations can be explained by loading-induced deformation in the presence of local lithologic gradients or by the influence of faults or structurally controlled anisotropic elastic properties. The observations highlight the utility of using mass position data from force-feedback broad-band seismometers for geodynamic studies.

Temperature and Redox Effect on Mineral Colonization in Juan de Fuca Ridge Flank Subsurface Crustal Fluids.

To examine microbe-mineral interactions in subsurface oceanic crust, we evaluated microbial colonization on crustal minerals that were incubated in borehole fluids for 1 year at the seafloor wellhead of a crustal borehole observatory (IODP Hole U1301A, Juan de Fuca Ridge flank) as compared to an experiment that was not exposed to subsurface crustal fluids (at nearby IODP Hole U1301B). In comparison to previous studies at these same sites, this approach allowed assessment of the effects of temperature, fluid chemistry, and/or mineralogy on colonization patterns of different mineral substrates, and an opportunity to verify the approach of deploying colonization experiments at an observatory wellhead at the seafloor instead of within the borehole. The Hole U1301B deployment did not have biofilm growth, based on microscopy and DNA extraction, thereby confirming the integrity of the colonization design against bottom seawater intrusion. In contrast, the Hole U1301A deployment supported biofilms dominated by Epsilonproteobacteria (43.5% of 370 16S rRNA gene clone sequences) and Gammaproteobacteria (29.3%). Sequence analysis revealed overlap in microbial communities between different minerals incubated at the Hole U1301A wellhead, indicating that mineralogy did not separate biofilm structure within the 1-year colonization experiment. Differences in the Hole U1301A wellhead biofilm community composition relative to previous studies from within the borehole using similar mineral substrates suggest that temperature and the diffusion of dissolved oxygen through plastic components influenced the mineral colonization experiments positioned at the wellhead. This highlights the capacity of low abundance crustal fluid taxa to rapidly establish communities on diverse mineral substrates under changing environmental conditions such as from temperature and oxygen.

Novel microbial assemblages inhabiting crustal fluids within mid-ocean ridge flank subsurface basalt.

Although little is known regarding microbial life within our planet's rock-hosted deep subseafloor biosphere, boreholes drilled through deep ocean sediment and into the underlying basaltic crust provide invaluable windows of access that have been used previously to document the presence of microorganisms within fluids percolating through the deep ocean crust. In this study, the analysis of 1.7 million small subunit ribosomal RNA genes amplified and sequenced from marine sediment, bottom seawater and basalt-hosted deep subseafloor fluids that span multiple years and locations on the Juan de Fuca Ridge flank was used to quantitatively delineate a subseafloor microbiome comprised of distinct bacteria and archaea. Hot, anoxic crustal fluids tapped by newly installed seafloor sampling observatories at boreholes U1362A and U1362B contained abundant bacterial lineages of phylogenetically unique Nitrospirae, Aminicenantes, Calescamantes and Chloroflexi. Although less abundant, the domain Archaea was dominated by unique, uncultivated lineages of marine benthic group E, the Terrestrial Hot Spring Crenarchaeotic Group, the Bathyarchaeota and relatives of cultivated, sulfate-reducing Archaeoglobi. Consistent with recent geochemical measurements and bioenergetic predictions, the potential importance of methane cycling and sulfate reduction were imprinted within the basalt-hosted deep subseafloor crustal fluid microbial community. This unique window of access to the deep ocean subsurface basement reveals a microbial landscape that exhibits previously undetected spatial heterogeneity.

Activity and phylogenetic diversity of sulfate-reducing microorganisms in low-temperature subsurface fluids within the upper oceanic crust.

The basaltic ocean crust is the largest aquifer system on Earth, yet the rates of biological activity in this environment are unknown. Low-temperature (<100°C) fluid samples were investigated from two borehole observatories in the Juan de Fuca Ridge (JFR) flank, representing a range of upper oceanic basement thermal and geochemical properties. Microbial sulfate reduction rates (SRR) were measured in laboratory incubations with 35S-sulfate over a range of temperatures and the identity of the corresponding sulfate-reducing microorganisms (SRM) was studied by analyzing the sequence diversity of the functional marker dissimilatory (bi)sulfite reductase (dsrAB) gene. We found that microbial sulfate reduction was limited by the decreasing availability of organic electron donors in higher temperature, more altered fluids. Thermodynamic calculations indicate energetic constraints for metabolism, which together with relatively higher cell-specific SRR reveal increased maintenance requirements, consistent with novel species-level dsrAB phylotypes of thermophilic SRM. Our estimates suggest that microbially-mediated sulfate reduction may account for the removal of organic matter in fluids within the upper oceanic crust and underscore the potential quantitative impact of microbial processes in deep subsurface marine crustal fluids on marine and global biogeochemical carbon cycling.

Controls on thallium uptake during hydrothermal alteration of the upper ocean crust

Hydrothermal circulation is a fundamental component of global biogeochemical cycles. However, the magnitude of the high temperature axial hydrothermal fluid flux remains disputed, and the lower temperature ridge flank fluid flux is difficult to quantify. Thallium (Tl) isotopes behave differently in axial compared to ridge flank systems, with Tl near-quantitatively stripped from the intrusive crust by high temperature hydrothermal reactions, but added to the lavas during low temperature reaction with seawater. This contrasting behavior provides a unique approach to determine the fluid fluxes associated with axial and ridge flank environments. Unfortunately, our understanding of the Tl isotopic mass balance is hindered by poor knowledge of the mineralogical, physical and chemical controls on Tl-uptake by the ocean crust.Here we use analyses of basaltic volcanic upper crust from Integrated Ocean Drilling Program Hole U1301B on the Juan de Fuca Ridge flank, combined with published analyses of dredged seafloor basalts and upper crustal basalts from Holes 504B and 896A, to investigate the controls on Tl-uptake by mid-ocean ridge basalts and evaluate when in the evolution of the ridge flank hydrothermal system Tl-uptake occurs.Seafloor basalts indicate an association between basaltic uptake of Tl from cold seawater and uptake of Cs and Rb, which are known to partition into K-rich phases. Although there is no clear relationship between Tl and K contents of seafloor basalts, the data do not rule out the incorporation of at least some Tl into the same minerals as the alkali elements. In contrast, we find no relationship between the Tl content and either the abundance of secondary phyllosilicate minerals, or the K, Cs or Rb contents in upper crustal basalts. We conclude that the uptake of Tl and alkali elements during hydrothermal alteration of the upper crust involves different processes and/or mineral phases compared to those that govern seafloor weathering. Furthermore, a correlation between the Tl and S concentrations of upper crustal basalts from Holes U1301B, 504B and 896A indicates that Tl is primarily incorporated into secondary sulfides. Given that some of these secondary sulfides formed as a result of microbial sulfate reduction, microbial action is at least indirectly responsible for Tl-uptake.Thallium-enrichment of ridge flank basalts requires a Tl-bearing fluid and physical, chemical and microbial conditions that favor secondary sulfide formation. Uptake of Tl occurs in reducing environments in the background rocks away from fluid flow pathways during early ‘open’ circulation of oxidizing seawater but more pervasively throughout the system during later ‘restricted’ circulation of reducing fluids. The Tl-isotope system is therefore a useful tracer of the fluid flux through both the ‘open’ and ‘restricted’ ridge flank hydrothermal regimes.

Dissolved hydrogen and methane in the oceanic basaltic biosphere

The oceanic basaltic crust is the largest aquifer on Earth and has the potential to harbor substantial subsurface microbial ecosystems, which hitherto remains largely uncharacterized and is analogous to extraterrestrial subsurface habitats. Within the sediment-buried 3.5 Myr old basaltic crust of the eastern Juan de Fuca Ridge flank, the circulating basement fluids have moderate temperature (∼65 °C) and low to undetectable dissolved oxygen and nitrate concentrations. Sulfate, present in high concentrations, is therefore expected to serve as the major electron acceptor in this subsurface environment. This study focused on the availability and potential sources of two important electron donors, methane (CH4) and hydrogen (H2), for the subseafloor biosphere. High integrity basement fluids were collected via fluid delivery lines associated with Integrated Ocean Drilling Program (IODP) Circulation Obviation Retrofit Kits (CORKs) that extend from basement depths to outlet ports at the seafloor. Two new CORKs installed during IODP 327 in 2010, 1362A and 1362B, were sampled in 2011 and 2013. The two CORKs are superior than earlier style CORKs in that they are equipped with coated casing and polytetrafluoroethylene fluid delivery lines, reducing the interaction between casing materials with the environment. Additional samples were collected from an earlier style CORK at Borehole 1301A.The basement fluids are enriched in H2 (0.05–1.8 μmol/kg), suggesting that the ocean basaltic aquifer can support H2-driven metabolism. The basement fluids also contain significant amount of CH4 (5–32 μmol/kg), revealing CH4 as an available substrate for subseafloor basaltic habitats. The δ13C values of CH4 from the three boreholes ranged from −22.5 to −58‰, while the δ2H values ranged from −316 to 57‰. The isotopic compositions of CH4 and the molecular compositions of hydrocarbons suggest that CH4 in the basement fluids is of both biogenic and abiotic origins, varying among sites and sampling times. The δ2H values of CH4 in CORK 1301A fluid samples are much more positive than found in all other marine environments investigated to date and are best explained by the partial microbial oxidation of biogenic CH4. In conclusion, our study shows that CH4 and H2 are persistently available to fuel the deep biosphere and that CH4 is both produced and potentially consumed by microorganisms in the oceanic basement.

Phylogenetic diversity of microorganisms in subseafloor crustal fluids from Holes 1025C and 1026B along the Juan de Fuca Ridge flank.

To expand investigations into the phylogenetic diversity of microorganisms inhabiting the subseafloor biosphere, basalt-hosted crustal fluids were sampled from Circulation Obviation Retrofit Kits (CORKs) affixed to Holes 1025C and 1026B along the Juan de Fuca Ridge (JdFR) flank using a clean fluid pumping system. These boreholes penetrate the crustal aquifer of young ocean crust (1.24 and 3.51 million years old, respectively), but differ with respect to borehole depth and temperature at the sediment-basement interface (147 m and 39°C vs. 295 m and 64°C, respectively). Cloning and sequencing of PCR-amplified small subunit ribosomal RNA genes revealed that fluids retrieved from Hole 1025C were dominated by relatives of the genus Desulfobulbus of the Deltaproteobacteria (56% of clones) and Candidatus Desulforudis of the Firmicutes (17%). Fluids sampled from Hole 1026B also contained plausible deep subseafloor inhabitants amongst the most abundant clone lineages; however, both geochemical analysis and microbial community structure reveal the borehole to be compromised by bottom seawater intrusion. Regardless, this study provides independent support for previous observations seeking to identify phylogenetic groups of microorganisms common to the deep ocean crustal biosphere, and extends previous observations by identifying additional lineages that may be prevalent in this unique environment.

Energy yields from chemolithotrophic metabolisms in igneous basement of the Juan de Fuca ridge flank system

The permeable rocks of the upper oceanic basement contain seawater-sourced fluids estimated to be ~2% of the global ocean volume. This represents a very large potential subsurface biosphere supported by chemosynthesis. Recent collection of high integrity samples of basement fluid from the sedimented young basaltic basement on the Juan de Fuca Ridge flanks, off the coasts of Vancouver Island (Canada) and Washington (USA), and subsequent chemical analyses permit numerical modeling of metabolic redox reaction energetics. Here, values of Gibbs free energy for potential chemolithotrophic net reactions were calculated in basement fluid and in zones where basement fluid and entrained seawater may mix; the energy yields are reported both on a per mole electrons transferred and on a per kg of basement fluid basis. In pure basement fluid, energy yields from the anaerobic respiration processes investigated are anemic, releasing <0.3J/kg basement fluid for all reactions except methane oxidation by ferric iron, which releases ~0.6J/kg basement fluid. In mixed solutions, aerobic oxidation of hydrogen, methane, and sulfide is the most exergonic on a per mole electron basis. Per kg of basement fluid, the aerobic oxidation of ammonia is by far the most exergonic at low temperature and high seawater:basement fluid ratio, decreasing by more than two orders of magnitude at the highest temperature (63°C) and lowest seawater:basement fluid ratio investigated. Compared with mixing zones in deep-sea hydrothermal systems, oceanic basement aquifers appear to be very low energy systems, but because of their expanse, may support what has been labeled the ‘starving majority’.

Microbial communities at the borehole observatory on the Costa Rica Rift flank (Ocean Drilling Program Hole 896A)

The microbiology of subsurface, hydrothermally influenced basaltic crust flanking mid-ocean ridges has remained understudied, due to the difficulty in accessing the subsurface environment. The instrumented boreholes resulting from scientific ocean drilling offer access to samples of the formation fluids circulating through oceanic crust. We analyzed the phylogenetic diversity of bacterial communities of fluid and microbial mat samples collected in situ from the observatory at Ocean Drilling Program Hole 896A, drilled into ~6.5 million-year-old basaltic crust on the flank of the Costa Rica Rift in the equatorial Pacific Ocean. Bacterial 16S rRNA gene sequences recovered from borehole fluid and from a microbial mat coating the outer surface of the fluid port revealed both unique and shared phylotypes. The dominant bacterial clones from both samples were related to the autotrophic, sulfur-oxidizing genus Thiomicrospira. Both samples yielded diverse gamma- and alphaproteobacterial phylotypes, as well as members of the Bacteroidetes, Planctomycetes, and Verrucomicrobia. Analysis of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) genes (cbbL and cbbM) from the sampling port mat and from the borehole fluid demonstrated autotrophic carbon assimilation potential for in situ microbial communities; most cbbL genes were related to those of the sulfur-oxidizing genera Thioalkalivibrio and Thiomicrospira, and cbbM genes were affiliated with uncultured phylotypes from hydrothermal vent plumes and marine sediments. Several 16S rRNA gene phylotypes from the 896A observatory grouped with phylotypes recovered from seawater-exposed basalts and sulfide deposits at inactive hydrothermal vents, but there is little overlap with hydrothermally influenced basaltic boreholes 1026B and U1301A on the Juan de Fuca Ridge flank, suggesting that site-specific characteristics of Hole 896A (i.e., seawater mixing into borehole fluids) affect the microbial community composition.

Acetogenesis in Deep Subseafloor Sediments of The Juan de Fuca Ridge Flank: A Synthesis of Geochemical, Thermodynamic, and Gene-based Evidence

In deep subsurface sediments of the Juan de Fuca Ridge Flank, porewater acetate that is depleted in 13 C relative to sedimentary organic matter indicates an acetogenic component to total acetate production. Thermodynamic calculations indicate common fermentation products or lignin monomers as potential substrates for acetogenesis. The classic autotrophic reaction may contribute as well, provided that dihydrogen (H 2 ) concentrations are not drawn down to the thermodynamic thresholds of the energetically more favorable processes of sulfate reduction and methanogenesis. A high diversity of novel formyl tetrahydrofolate synthetase (fhs) genes throughout the upper half of the sediment column indicates the genetic potential for acetogenesis. Our results suggest that a substantial fraction of the acetate produced in marine sediment porewaters may derive from acetogenesis, in addition to the conventionally invoked sources fermentation and sulfate reduction.

Dehalogenation Activities and Distribution of Reductive Dehalogenase Homologous Genes in Marine Subsurface Sediments

Halogenated organic compounds serve as terminal electron acceptors for anaerobic respiration in a diverse range of microorganisms. Here, we report on the widespread distribution and diversity of reductive dehalogenase homologous (rdhA) genes in marine subsurface sediments. A total of 32 putative rdhA phylotypes were detected in sediments from the southeast Pacific off Peru, the eastern equatorial Pacific, the Juan de Fuca Ridge flank off Oregon, and the northwest Pacific off Japan, collected at a maximum depth of 358 m below the seafloor. In addition, significant dehalogenation activity involving 2,4,6-tribromophenol and trichloroethene was observed in sediment slurry from the Nankai Trough Forearc Basin. These results suggest that dehalorespiration is an important energy-yielding pathway in the subseafloor microbial ecosystem.

Sedimentation patterns, folding, and fluid upflow above a buried basement ridge: Results from 2‐D and 3‐D seismic surveys at the eastern Juan de Fuca Ridge flank

During R/V Sonne Cruise SO 149, new two‐dimensional (2‐D) and 3‐D multifrequency seismic data were collected at the eastern flank of the Juan de Fuca Ridge, off the west coast of North America. Two‐dimensional overview lines indicate that turbidity currents moving southward are partly trapped between a buried basement elevation called First Ridge and another elongated basement obstruction further to the west. As a consequence, sediments form a local topographic high and a distinct wedge‐shaped section in between the two ridge segments. On the basis of these seismic observations a simple conceptual model is developed to explain local sediment accumulation patterns at the flanks and on top of First Ridge. This model is consistent with seismic, coring, and drilling results that provide quantitative constraints for the remaining seafloor relief above a fully buried basement high as well as sediment porosity values on top of First Ridge that are higher than those in adjacent troughs. In general, higher porosity values may explain lower seismic reflection amplitudes, but the general trend of porosity increase above First Ridge due to vertical grain size sorting within a turbidity current has to be distinguished from additional, overprinting effects that confine the major portion of the observed reflection amplitude decrease to narrow, vertical zones. At the location of such zones, seismic data indicate that pronounced basement peaks are associated with distinct folding of the sediment layers above. It is thus suggested that the folding and the local seismic amplitude decrease are related. Folding and the associated extensional strain field forming throughout such folds can explain locally high layer inclination and increased porosity, which both affect seismic reflection amplitudes. Folding and associated fracturing may cause locally increased permeability and have the potential to focus hydrothermally driven fluid upflow and to develop the plumbing system required to explain the observed fluid seepage through a fine‐grained sediment seal.