Discovery of active off-axis hydrothermal vents at 9° 54'N East Pacific Rise.

Comprehensive knowledge of the distribution of active hydrothermal vent fields along midocean ridges is essential to understanding global chemical and heat fluxes and endemic faunal distributions. However, current knowledge is biased by a historical preference for on-axis surveys. A scarcity of high-resolution bathymetric surveys in off-axis regions limits vent identification, which implies that the number of vents may be underestimated. Here, we present the discovery of an active, high-temperature, off-axis hydrothermal field on a fast-spreading ridge. The vent field is located 750 m east of the East Pacific Rise axis and ∼7 km north of on-axis vents at 9° 50'N, which are situated in a 50- to 100-m-wide trough. This site is currently the largest vent field known on the East Pacific Rise between 9 and 10° N. Its proximity to a normal fault suggests that hydrothermal fluid pathways are tectonically controlled. Geochemical evidence reveals deep fluid circulation to depths only 160 m above the axial magma lens. Relative to on-axis vents at 9° 50'N, these off-axis fluids attain higher temperatures and pressures. This tectonically controlled vent field may therefore exhibit greater stability in fluid composition, in contrast to more dynamic, dike-controlled, on-axis vents. The location of this site indicates that high-temperature convective circulation cells extend to greater distances off axis than previously realized. Thorough high-resolution mapping is necessary to understand the distribution, frequency, and physical controls on active off-axis vent fields so that their contribution to global heat and chemical fluxes and role in metacommunity dynamics can be determined.

Holocene carbonate record of Lake Kivu reflects the history of hydrothermal activity.

The sediment record of Lake Kivu reveals a complex volcanogenic and climatic Holocene history. Investigation of the inorganic carbonate record dates the onset of carbonate deposition in the mid-Holocene in Kivu's deep northern and eastern basins and identifies conditions enabling deposition. The magnitude and timing of carbonate-rich sedimentation is not so much controlled by climate but, instead, linked strongly to hydrothermal activity in the basin. Sublacustrine springs supply the vast majority of the calcium and carbonate ions required for supersaturation with respect to aragonite. This major hydrothermal activity that permanently stratifies Lake Kivu today was initiated ∼3,100 y before present (3.1 ka), when carbonate-rich sediments first appeared in the Holocene record. Aragonite is the dominant CaCO3 mineral present in the lake deposits. Both δ13C and δ18O of the aragonite are enriched above the expected kinetic fractionation of meteoric waters, suggesting a volcanogenic influence on the formation waters. Repeated major fluctuations in the carbonate record after 3.1 ka therefore most likely reflect the historical variation in hydrothermal inputs.

Hydrothermal impacts on trace element and isotope ocean biogeochemistry.

Hydrothermal activity occurs in all ocean basins, releasing high concentrations of key trace elements and isotopes (TEIs) into the oceans. Importantly, the calculated rate of entrainment of the entire ocean volume through turbulently mixing buoyant hydrothermal plumes is so vigorous as to be comparable to that of deep-ocean thermohaline circulation. Consequently, biogeochemical processes active within deep-ocean hydrothermal plumes have long been known to have the potential to impact global-scale biogeochemical cycles. More recently, new results from GEOTRACES have revealed that plumes rich in dissolved Fe, an important micronutrient that is limiting to productivity in some areas, are widespread above mid-ocean ridges and extend out into the deep-ocean interior. While Fe is only one element among the full suite of TEIs of interest to GEOTRACES, these preliminary results are important because they illustrate how inputs from seafloor venting might impact the global biogeochemical budgets of many other TEIs. To determine the global impact of seafloor venting, however, requires two key questions to be addressed: (i) What processes are active close to vent sites that regulate the initial high-temperature hydrothermal fluxes for the full suite of TEIs that are dispersed through non-buoyant hydrothermal plumes? (ii) How do those processes vary, globally, in response to changing geologic settings at the seafloor and/or the geochemistry of the overlying ocean water? In this paper, we review key findings from recent work in this realm, highlight a series of key hypotheses arising from that research and propose a series of new GEOTRACES modelling, section and process studies that could be implemented, nationally and internationally, to address these issues.This article is part of the themed issue 'Biological and climatic impacts of ocean trace element chemistry'.

Ecology of Bathymodiolus puteoserpentis mussels from the Snake Pit vent field (Mid-Atlantic Ridge).

Along the northern Mid-Atlantic Ridge (nMAR), in habitats under moderate (<10 °C) hydrothermal influence on the Snake Pit vent field (SP), large assemblages dominated by Bathymodiolin mussels remain poorly characterised, contrary to those in warmer habitats dominated by gastropods and alvinocaridid shrimps that were recently described. In this study, we assessed and compared the population structure, biomass, diversity and trophic interactions of two Bathymodiolus puteoserpentis assemblages and their associated fauna at SP. Three sampling units distanced by 30 cm were sampled in 2014 during the BICOSE cruise at the top of the Moose site (''Elan'' site), while few meters further down three others, distanced by ∼1 m were obtained in 2018 during the BICOSE 2 cruise at the edifice's base. We observed a micro-scale heterogeneity between these six sampling units partially explained by temperature variations, proximity to hydrothermal fluids and position on the edifice. Meiofauna dominate or co-dominate most of the sampling units, with higher densities at the base of the edifice. In terms of macrofauna, high abundance of Pseudorimula midatlantica gastropods was observed at the top of the vent edifice, while numerous Ophioctenella acies ophiuroids were found at the base. Contrary to what was expected, the apparent health and abundance of mussels seems to indicate a current climax stage of the community. However, the modification of B. puteoserpentis isotopic signatures, low number of juveniles decreasing over the two years and observations made during several French cruises in the study area raise questions about the fate of the B. puteoserpentis population over time, which remains to be verified in a future sampling campaign.

Resilience of the seagrass Posidonia oceanica following pulse-type disturbance.

Understanding the response of species to disturbance and the ability to recover is crucial for preventing their potential collapse and ecosystem phase shifts. Explosive submarine activity, occurring in shallow volcanic vents, can be considered as a natural pulse disturbance, due to its suddenness and high intensity, potentially affecting nearby species and ecosystems. Here, we present the response of Posidonia oceanica, a long-lived seagrass, to an exceptional submarine volcanic explosion, which occurred in the Aeolian Archipelago (Italy, Mediterranean Sea) in 2002, and evaluate its resilience in terms of time required to recover after such a pulse event. The study was carried out in 2011 in the sea area off Panarea Island, in the vicinity of Bottaro Island by adopting a back-dating methodological approach, which allowed a retrospective analysis of the growth performance and stable carbon isotopes (δ13C) in sheaths and rhizomes of P. oceanica, during a 10-year period (2001-2010). After the 2002 explosion, a trajectory shift towards decreasing values for both growth performance and δ13C in sheaths and rhizomes was observed. The decreasing trend reversed in 2004 when recovery took place progressively for all the analysed variables. Full recovery of P. oceanica occurred 8 years after the explosive event with complete restoration of all the variables (rhizome growth performance and δ13C) by 2010. Given the ecological importance of this seagrass in marine coastal ecosystems and its documented large-scale decline, the understanding of its potential recovery in response to environmental changes is imperative.

Arsenic and high affinity phosphate uptake gene distribution in shallow submarine hydrothermal sediments.

The toxicity of arsenic (As) towards life on Earth is apparent in the dense distribution of genes associated with As detoxification across the tree of life. The ability to defend against As is particularly vital for survival in As-rich shallow submarine hydrothermal ecosystems along the Hellenic Volcanic Arc (HVA), where life is exposed to hydrothermal fluids containing up to 3000 times more As than present in seawater. We propose that the removal of dissolved As and phosphorus (P) by sulfide and Fe(III)(oxyhydr)oxide minerals during sediment-seawater interaction, produces nutrient-deficient porewaters containing < 2.0 ppb P. The porewater arsenite-As(III) to arsenate-As(V) ratios, combined with sulfide concentration in the sediment and/or porewater, suggest a hydrothermally-induced seafloor redox gradient. This gradient overlaps with changing high affinity phosphate uptake gene abundance. High affinity phosphate uptake and As cycling genes are depleted in the sulfide-rich settings, relative to the more oxidizing habitats where mainly Fe(III)(oxyhydr)oxides are precipitated. In addition, a habitat-wide low As-respiring and As-oxidizing gene content relative to As resistance gene richness, suggests that As detoxification is prioritized over metabolic As cycling in the sediments. Collectively, the data point to redox control on Fe and S mineralization as a decisive factor in the regulation of high affinity phosphate uptake and As cycling gene content in shallow submarine hydrothermal ecosystems along the HVA.

Spiculosiphon oceana (Foraminifera) a new bio-indicator of acidic environments related to fluid emissions of the Zannone Hydrothermal Field (central Tyrrhenian Sea).

The new record of a shallow-water submarine hydrothermal field (<150 m w.d.) in the western Mediterranean Sea (Tyrrhenian Sea, Italy) allows us to study CO2 fluid impact on benthic foraminifers. Benthic foraminifers calcification process is sensitive to ocean acidification and to local chemical and physical parameters of seawater and pore water. Thus, foraminifers can record specific environmental conditions related to hydrothermal fluids, but at present their response to such activity is poorly defined. The major outcome of this study is the finding of a very uncommon taxon for the Mediterranean Sea, i.e., the Spiculosiphon oceana, a giant foraminifer agglutinating spicules of sponges. This evidence, along with the strong decrease of calcareous tests in the foraminiferal assemblages associated to hydrothermal activity, provides new insights on the meiofauna living in natural stressed environment. In particular, observations obtained from this study allow us to consider S. oceana a potential tolerant species of high CO2 concentrations (about 2-4 times higher than the normal marine values) and a proxy of acidic environments as well as of recent ocean acidification processes.

Microbial Biomarker Transition in High-Altitude Sinter Mounds From El Tatio (Chile) Through Different Stages of Hydrothermal Activity.

Geothermal springs support microbial communities at elevated temperatures in an ecosystem with high preservation potential that makes them interesting analogs for early evolution of the biogeosphere. The El Tatio geysers field in the Atacama Desert has astrobiological relevance due to the unique occurrence of geothermal features with steep hydrothermal gradients in an otherwise high altitude, hyper-arid environment. We present here results of our multidisciplinary field and molecular study of biogeochemical evidence for habitability and preservation in silica sinter at El Tatio. We sampled three morphologically similar geyser mounds characterized by differences in water activity (i.e., episodic liquid water, steam, and inactive geyser lacking hydrothermal activity). Multiple approaches were employed to determine (past and present) biological signatures and dominant metabolism. Lipid biomarkers indicated relative abundance of thermophiles (dicarboxylic acids) and sulfate reducing bacteria (branched carboxylic acids) in the sinter collected from the liquid water mound; photosynthetic microorganisms such as cyanobacteria (alkanes and isoprenoids) in the steam sinter mound; and archaea (squalane and crocetane) as well as purple sulfur bacteria (cyclopropyl acids) in the dry sinter from the inactive geyser. The three sinter structures preserved biosignatures representative of primary (thermophilic) and secondary (including endoliths and environmental contaminants) microbial communities. Sequencing of environmental 16S rRNA genes and immuno-assays generally corroborated the lipid-based microbial identification. The multiplex immunoassays and the compound-specific isotopic analysis of carboxylic acids, alkanols, and alkanes indicated that the principal microbial pathway for carbon fixation in the three sinter mounds was through the Calvin cycle, with a relative larger contribution of the reductive acetyl-CoA pathway in the dry system. Other inferred metabolic traits varied from the liquid mound (iron and sulfur chemistry), to the steam mound (nitrogen cycle), to the dry mound (perchlorate reduction). The combined results revealed different stages of colonization that reflect differences in the lifetime of the mounds, where primary communities dominated the biosignatures preserved in sinters from the still active geysers (liquid and steam mounds), in contrast to the surviving metabolisms and microbial communities at the end of lifetime of the inactive geothermal mound.

Characteristics of hydrocarbons in sediment core samples from the northern Okinawa Trough.

Sediment core samples from the northern Okinawa Trough (OT) were analyzed to determine abundances and distributions of hydrocarbons by gas chromatography-mass spectrometer (GC-MS). The results show that the n-alkanes in this sediment core conform to a bimodal distribution, and exhibit an odd-to-even predominance of high molecular weights compared to an even-to-odd predominance in low molecular weight n-alkanes with maxima at C16 and C18. The concentrations of bitumen, alkanes and polyaromatic hydrocarbons (PAHs) were higher in samples S10-07 than all others. Three maturity parameters as well as the ratios between parent phenanthrenes (Ps) and methylphenanthrenes (MPs) in samples S10-07 and S10-17 were higher. The distribution and composition of hydrocarbons in sample S10-07 suggest that one, or several, undetected hydrothermal fields may be present in the region of this sediment core. Results also suggest that volcanism may be the main reason for the observed distribution and composition of hydrocarbons in S10-17 sample.

Proteomic responses to metal-induced oxidative stress in hydrothermal vent-living mussels, Bathymodiolus sp., on the Southwest Indian Ridge.

Bathymodiolin mussels are amongst the dominant fauna occupying hydrothermal vent ecosystems throughout the World's oceans. This subfamily inhabits a highly ephemeral and variable environment, where exceptionally high concentrations of reduced sulphur species and heavy metals necessitate adaptation of specialised detoxification mechanisms. Whilst cellular responses to common anthropogenic pollutants are well-studied in shallow-water species, they remain limited in deep-sea vent fauna. Bathymodiolus sp. were sampled from two newly-discovered vent sites on the Southwest Indian Ridge (Tiamat and Knuckers Gaff) by the remotely operated vehicle (ROV) Kiel 6000 during the RRS James Cook cruise, JC 067 in November 2011. Here, we use redox proteomics to investigate the effects of tissue metal accumulation on protein expression and thiol oxidation in gill. Following 2D PAGE, we demonstrate a significant difference in intensity in 30 protein spots in this organ between the two vent sites out of 205 matched spots. We also see significant variations in thiol oxidation in 15 spots, out of 143 matched. At Tiamat, 23 protein spots are up-regulated compared to Knuckers Gaff and we identify 5 of these with important roles in metabolism, cell structure, stress response, and redox homeostasis. We suggest that increased metal exposure triggers changes in the proteome, regulating tissue uptake. This is evident both between vent sites and across a chemical gradient within the Knuckers Gaff vent site. Our findings highlight the importance of proteomic plasticity in successful adaptation to the spatially and temporally fluctuating chemical environments that are characteristic of hydrothermal vent habitats.