Hydrogen is an energy source for hydrothermal vent symbioses

The discovery of deep-sea hydrothermal vents in 1977 revolutionized our understanding of the energy sources that fuel primary productivity on Earth. Hydrothermal vent ecosystems are dominated by animals that live in symbiosis with chemosynthetic bacteria. So far, only two energy sources have been shown to power chemosynthetic symbioses: reduced sulphur compounds and methane. Using metagenome sequencing, single-gene fluorescence in situ hybridization, immunohistochemistry, shipboard incubations and in situ mass spectrometry, we show here that the symbionts of the hydrothermal vent mussel Bathymodiolus from the Mid-Atlantic Ridge use hydrogen to power primary production. In addition, we show that the symbionts of Bathymodiolus mussels from Pacific vents have hupL, the key gene for hydrogen oxidation. Furthermore, the symbionts of other vent animals such as the tubeworm Riftia pachyptila and the shrimp Rimicaris exoculata also have hupL. We propose that the ability to use hydrogen as an energy source is widespread in hydrothermal vent symbioses, particularly at sites where hydrogen is abundant.     

Adaptive visual metamorphosis in a deep-sea hydrothermal vent crab

Hydrothermal vents along the mid-ocean ridges host ephemeral ecosystems of diverse endemic fauna including several crustacean species, some of which undergo planktonic development as larvae up to 1,000 m above and 100 km away from the vents. Little is known about the role of vision in the life history of vent fauna. Here we report that planktonic zoea larvae of the vent crab Bythograea thermydron possess image-forming compound eyes with a visual pigment sensitive to the blue light of mesopelagic waters. As they metamorphose and begin to descend to and settle at the vents, they lose their image-forming optics and develop high-sensitivity naked-retina eyes. The spectral absorbance of the visual pigment in these eyes shifts towards longer wavelengths from larva to postlarva to adult. This progressive visual metamorphosis trades imaging for increased sensitivity, and changes spectral sensitivity from the blue wavelengths of the larval environment towards the dim, longer wavelengths produced in the deeper bathypelagic vent environment of the adults. As hydrothermal vents produce light, vision may supplement thermal and chemical senses to orient postlarval settlement at vent sites.     

Earthquake-induced changes in a hydrothermal system on the Juan de Fuca mid-ocean ridge

Hydrothermal vents on mid-ocean ridges of the northeast Pacific Ocean are known to respond to seismic disturbances, with observed changes in vent temperature. But these disturbances resulted from submarine volcanic activity; until now, there have been no observations of the response of a vent system to non-magmatic, tectonic events. Here we report measurements of hydrothermal vent temperature from several vents on the Juan de Fuca ridge in June 1999, before, during and after an earthquake swarm of apparent tectonic origin. Vent fluid temperatures began to rise 4-11 days after the first earthquake. Following this initial increase, the vent temperatures oscillated for about a month before settling down to higher values. We also observed a tenfold increase in fluid output from the hydrothermal system over a period of at least 80 days, extending along the entire ridge segment. Such a large, segment-wide thermal response to relatively modest tectonic activity is surprising, and raises questions about the sources of excess heat and fluid, and the possible effect on vent biological communities.     

Larval dispersal potential of the tubeworm Riftia pachyptila at deep-sea hydrothermal vents

Hydrothermal vents are ephemeral because of frequent volcanic and tectonic activities associated with crust formation. Although the larvae of hydrothermal vent fauna can rapidly colonize new vent sites separated by tens to hundreds of kilometres, the mechanisms by which these larvae disperse and recruit are not understood. Here we integrate physiological, developmental and hydrodynamic data to estimate the dispersal potential of larvae of the giant tubeworm Riftia pachyptila. At in situ temperatures and pressures (2 degrees C and 250 atm), we estimate that the metabolic lifespan for a larva of R. pachyptila averages 38 days. In the measured flow regime at a fast-spreading ridge axis (9 degrees 50' N; East Pacific Rise), this lifespan results in potential along-ridge dispersal distances that rarely exceed 100 km. This limited dispersal results not from the physiological performance of the embryos and larvae, but instead from transport limitations imposed by periodic reversals in along-ridge flows and sustained episodes of across-ridge flow. The lifespan presented for these larvae can now be used to predict dispersal under current regimes at other hydrothermal vent sites.     

The habitat and nature of early life

Earth is over 4,500 million years old. Massive bombardment of the planet took place for the first 500-700 million years, and the largest impacts would have been capable of sterilizing the planet. Probably until 4,000 million years ago or later, occasional impacts might have heated the ocean over 100 degrees C. Life on Earth dates from before about 3,800 million years ago, and is likely to have gone through one or more hot-ocean 'bottlenecks'. Only hyperthermophiles (organisms optimally living in water at 80-110 degrees C) would have survived. It is possible that early life diversified near hydrothermal vents, but hypotheses that life first occupied other pre-bottleneck habitats are tenable (including transfer from Mars on ejecta from impacts there). Early hyperthermophile life, probably near hydrothermal systems, may have been non-photosynthetic, and many housekeeping proteins and biochemical processes may have an original hydrothermal heritage. The development of anoxygenic and then oxygenic photosynthesis would have allowed life to escape the hydrothermal setting. By about 3,500 million years ago, most of the principal biochemical pathways that sustain the modern biosphere had evolved, and were global in scope.     

Pre-biotic organic matter from comets and asteroids

Several authors have suggested that comets or carbonaceous asteroids contributed large amounts of organic matter to the primitive Earth, and thus possibly played a vital role in the origin of life. But organic matter cannot survive the extremely high temperatures (>10(4) K) reached on impact, which atomize the projectile and break all chemical bonds. Only fragments small enough to be gently decelerated by the atmosphere--principally meteors of 10(-12)-10(-6) g--can deliver their organic matter intact. The amount of such 'soft-landed' organic carbon can be estimated from data for the infall rate of meteoritic matter. At present rates, only approximately 0.006 g cm-2 intact organic carbon would accumulate in 10(8) yr, but at the higher rates of approximately 4 x 10(9) yr ago, about 20 g cm-2 may have accumulated in the few hundred million years between the last cataclysmic impact and the beginning of life. It may have included some biologically important compounds that did not form by abiotic synthesis on Earth.     

Feeding behaviour: hydrothermal vent crabs feast on sea 'snow'

The crab Xenograpsus testudinatus lives at enormously high densities around the sulphur-rich hydrothermal vents found in shallow waters off Taiwan, even though this acidic environment is low in nutrients. Here we show that these crabs swarm out of their crevices at slack water and feed on the vast numbers of zooplankton that are killed by the vents' sulphurous plumes, and that rain down like marine 'snow'. This opportunistic feeding behaviour explains how the crabs are able to survive in the adverse toxic environment of these shallow hydrothermal vents.     

Discovery of abundant hydrothermal venting on the ultraslow-spreading Gakkel ridge in the Arctic Ocean

Submarine hydrothermal venting along mid-ocean ridges is an important contributor to ridge thermal structure, and the global distribution of such vents has implications for heat and mass fluxes from the Earth's crust and mantle and for the biogeography of vent-endemic organisms. Previous studies have predicted that the incidence of hydrothermal venting would be extremely low on ultraslow-spreading ridges (ridges with full spreading rates <2 cm x yr(-1)-which make up 25 per cent of the global ridge length), and that such vent systems would be hosted in ultramafic in addition to volcanic rocks. Here we present evidence for active hydrothermal venting on the Gakkel ridge, which is the slowest spreading (0.6-1.3 cm x yr(-1)) and least explored mid-ocean ridge. On the basis of water column profiles of light scattering, temperature and manganese concentration along 1,100 km of the rift valley, we identify hydrothermal plumes dispersing from at least nine to twelve discrete vent sites. Our discovery of such abundant venting, and its apparent localization near volcanic centres, requires a reassessment of the geologic conditions that control hydrothermal circulation on ultraslow-spreading ridges.     

Volcanic production of polyphosphates and its relevance to prebiotic evolution

Phosphates would probably have been essential compounds for prebiotic evolution on the primitive Earth. In this context, there have been several studies of condensation of water-soluble phosphates to polyphosphates and phosphorylation and condensation or polymerization of biomolecules with polyphosphates. But most of the phosphorus on the early Earth would have been in the form of water-insoluble apatite, and the origin of the water-soluble polyphosphates required for prebiotic evolution has therefore been a mystery. Here we show, both from experiments that simulate magmatic conditions and from analysis of volatile condensates in volcanic gas, that volcanic activity can produce water-soluble polyphosphates through partial hydrolysis of P4O10. This mechanism seems to be the only viable route identified so far for the production of these species on the primitive Earth.     

深海热液生物群落研究综述

深海热液口吸其生物群落自1977年发现以来．受到海洋学界和生物界的广泛关注，并在热液生物生理、生态和生物地理学等方面取得了显著的进展．本文就该领域的研究进展做一简要的综述，主要内容包括热液生物群落的基本特征、种类组成和时空变化．     

Possible methane-induced polar warming in the early Eocene

Reconstructions of early Eocene climate depict a world in which the polar environments support mammals and reptiles, deciduous forests, warm oceans and rare frost conditions. At the same time, tropical sea surface temperatures are interpreted to have been the same as or slightly cooler than present values. The question of how to warm polar regions of Earth without noticeably warming the tropics remains unresolved; increased amounts of greenhouse gases would be expected to warm all latitudes equally. Oceanic heat transport has been postulated as a mechanism for heating high latitudes, but it is difficult to explain the dynamics that would achieve this. Here we consider estimates of Eocene wetland areas and suggest that the flux of methane, an important greenhouse gas, may have been substantially greater during the Eocene than at present. Elevated methane concentrations would have enhanced early Eocene global warming, and also might specifically have prevented severe winter cooling of polar regions because of the potential of atmospheric methane to promote the formation of optically thick, polar stratospheric ice clouds.     

Chemical speciation drives hydrothermal vent ecology

The physiology and biochemistry of many taxa inhabiting deep-sea hydrothermal vents have been elucidated; however, the physicochemical factors controlling the distribution of these organisms at a given vent site remain an enigma after 20 years of research. The chemical speciation of particular elements has been suggested as key to controlling biological community structure in these extreme aquatic environments. Implementation of electrochemical technology has allowed us to make in situ measurements of chemical speciation at vents located at the East Pacific Rise (9 degrees 50' N) and on a scale relevant to the biology. Here we report that significant differences in oxygen, iron and sulphur speciation strongly correlate with the distribution of specific taxa in different microhabitats. In higher temperature (> 30 degrees C) microhabitats, the appreciable formation of soluble iron-sulphide molecular clusters markedly reduces the availability of free H2S/HS- to vent (micro)organisms, thus controlling the available habitat.     

A ubiquitous thermoacidophilic archaeon from deep-sea hydrothermal vents

Deep-sea hydrothermal vents are important in global biogeochemical cycles, providing biological oases at the sea floor that are supported by the thermal and chemical flux from the Earth's interior. As hot, acidic and reduced hydrothermal fluids mix with cold, alkaline and oxygenated sea water, minerals precipitate to form porous sulphide-sulphate deposits. These structures provide microhabitats for a diversity of prokaryotes that exploit the geochemical and physical gradients in this dynamic ecosystem. It has been proposed that fluid pH in the actively venting sulphide structures is generally low (pH < 4.5), yet no extreme thermoacidophile has been isolated from vent deposits. Culture-independent surveys based on ribosomal RNA genes from deep-sea hydrothermal deposits have identified a widespread euryarchaeotal lineage, DHVE2 (deep-sea hydrothermal vent euryarchaeotic 2). Despite the ubiquity and apparent deep-sea endemism of DHVE2, cultivation of this group has been unsuccessful and thus its metabolism remains a mystery. Here we report the isolation and cultivation of a member of the DHVE2 group, which is an obligate thermoacidophilic sulphur- or iron-reducing heterotroph capable of growing from pH 3.3 to 5.8 and between 55 and 75 degrees C. In addition, we demonstrate that this isolate constitutes up to 15% of the archaeal population, providing evidence that thermoacidophiles may be key players in the sulphur and iron cycling at deep-sea vents.     

The evolution of the marine phosphate reservoir

Phosphorus is a biolimiting nutrient that has an important role in regulating the burial of organic matter and the redox state of the ocean-atmosphere system. The ratio of phosphorus to iron in iron-oxide-rich sedimentary rocks can be used to track dissolved phosphate concentrations if the dissolved silica concentration of sea water is estimated. Here we present iron and phosphorus concentration ratios from distal hydrothermal sediments and iron formations through time to study the evolution of the marine phosphate reservoir. The data suggest that phosphate concentrations have been relatively constant over the Phanerozoic eon, the past 542 million years (Myr) of Earth's history. In contrast, phosphate concentrations seem to have been elevated in Precambrian oceans. Specifically, there is a peak in phosphorus-to-iron ratios in Neoproterozoic iron formations dating from ～750 to ～635?Myr ago, indicating unusually high dissolved phosphate concentrations in the aftermath of widespread, low-latitude 'snowball Earth' glaciations. An enhanced postglacial phosphate flux would have caused high rates of primary productivity and organic carbon burial and a transition to more oxidizing conditions in the ocean and atmosphere. The snowball Earth glaciations and Neoproterozoic oxidation are both suggested as triggers for the evolution and radiation of metazoans. We propose that these two factors are intimately linked; a glacially induced nutrient surplus could have led to an increase in atmospheric oxygen, paving the way for the rise of metazoan life.     

Early-stage mineralization of hydrothermal tubeworms： New insights into the role of microorganisms in the process of mineralization

As an important part of marine hydrothermal system, hydrothermal vent faunas live in hydrothermal inorganic environment and closely interact with hydrothermal inorganic environment. Sometimes, they can participate in the mineralization process of modern hydrothermal site. Hydrothermal vent faunas, particularly vesUmentiferan and polychaete tubeworms, are occasionally preserved in the geological record. Study on the early mineralization process of hydrothermal vent fauna is significant for understanding the interaction between mineral and organism, and also the formation and preservation mechanism of geological fossil in hydrothermal environment. In this paper, the early stage of mineralization of Vestimentiferan Ridgeia piscesae tubes collected from Juan de Fuca Ridge is studied. The results showed that a lot of filamentous microorganisms were unevenly distributed on the surface of internal wall and in the interspace of the wall of tubeworm. In some cases, microorganisms aggregated as thin layers in or on the wall of tubeworm. The surfaces of microbial cells and the products of microbial degradation may play an important role in the early mineralization of tubeworm. Semitransparent thin layers of organic matter containing sulfur and sulfur granules were commonly found on the wall of tubeworm with lower degree of mineralization. The degradation production of these semitransparent thin layers may accelerate the mineralization of tube wall during the early stage. EDS results showed that on the tube walls some chemical elements such as Fe, P, Ca and Si are selectively enriched from ambient hydrothermal environment. Interestingly, P, Ca and Si covary with Fe content. Because element S originated from the bio-oxidation of H2S by symbiotic microorganism in the tissue of tubeworm, it can be considered as a biomarker when studying the mineralization process of tube wall. Based on the characteristics of tubeworms with different degrees of mineralization, we suggested that the early mineralization stage of tube wall was mai     

Palaeoclimatology: evidence for hot early oceans?

The oxygen isotopes in sedimentary cherts (siliceous sediments) have been used to argue that the Precambrian oceans were hot--with temperatures of up to 70 degrees C at 3.3 Gyr before present. Robert and Chaussidon measure silicon isotopes in cherts and arrive at a similar conclusion. We suggest here that both isotope trends may be caused by variations in seawater isotope composition, rather than in ocean temperatures. If so, then the climate of the early Earth may have been temperate, as it is today, and therefore more consistent with evidence for Precambrian glaciations and with constraints inferred from biological evolution.     

Volcanic carbon dioxide vents show ecosystem effects of ocean acidification

The atmospheric partial pressure of carbon dioxide (p(CO(2))) will almost certainly be double that of pre-industrial levels by 2100 and will be considerably higher than at any time during the past few million years. The oceans are a principal sink for anthropogenic CO(2) where it is estimated to have caused a 30% increase in the concentration of H(+) in ocean surface waters since the early 1900s and may lead to a drop in seawater pH of up to 0.5 units by 2100 (refs 2, 3). Our understanding of how increased ocean acidity may affect marine ecosystems is at present very limited as almost all studies have been in vitro, short-term, rapid perturbation experiments on isolated elements of the ecosystem. Here we show the effects of acidification on benthic ecosystems at shallow coastal sites where volcanic CO(2) vents lower the pH of the water column. Along gradients of normal pH (8.1-8.2) to lowered pH (mean 7.8-7.9, minimum 7.4-7.5), typical rocky shore communities with abundant calcareous organisms shifted to communities lacking scleractinian corals with significant reductions in sea urchin and coralline algal abundance. To our knowledge, this is the first ecosystem-scale validation of predictions that these important groups of organisms are susceptible to elevated amounts of p(CO(2)). Sea-grass production was highest in an area at mean pH 7.6 (1,827 (mu)atm p(CO(2))) where coralline algal biomass was significantly reduced and gastropod shells were dissolving due to periods of carbonate sub-saturation. The species populating the vent sites comprise a suite of organisms that are resilient to naturally high concentrations of p(CO(2)) and indicate that ocean acidification may benefit highly invasive non-native algal species. Our results provide the first in situ insights into how shallow water marine communities might change when susceptible organisms are removed owing to ocean acidification.     

The visibility of 350 degrees C black-body radiation by the shrimp Rimicaris exoculata and man

The eye of the 'eyeless' shrimp Rimicaris exoculata is unusual in having no image-forming optics and a high concentration of rhodopsin. The shrimps swarm around 350 degrees C hydrothermal 'black smoker' vents in the Mid-Atlantic Ridge. There is no other known source of visible light in the shrimp's environment. The spectral sensitivity of rhodopsin is well matched to typical spectra of bioluminescence of organisms found at lesser depths, but other animals detect such emissions without the unusual features of the R. exoculata eye. These two features are most easily understood as an adaptation for the detection of extremely faint sources of light. Physical calculations presented here indicate that the shrimp could see the black-body radiation of the 350 degrees C vents, even though these sources are practically invisible to the human eye. This would be useful to the shrimp as it feeds on sulphide-loving bacteria very near to the vents but must avoid the lethal 350 degrees C vents themselves.     

Evidence from massive siderite beds for a CO2-rich atmosphere before approximately 1.8 billion years ago

It is generally thought that, in order to compensate for lower solar flux and maintain liquid oceans on the early Earth, methane must have been an important greenhouse gas before approximately 2.2 billion years (Gyr) ago. This is based upon a simple thermodynamic calculation that relates the absence of siderite (FeCO3) in some pre-2.2-Gyr palaeosols to atmospheric CO2 concentrations that would have been too low to have provided the necessary greenhouse effect. Using multi-dimensional thermodynamic analyses and geological evidence, we show here that the absence of siderite in palaeosols does not constrain atmospheric CO2 concentrations. Siderite is absent in many palaeosols (both pre- and post-2.2-Gyr in age) because the O2 concentrations and pH conditions in well-aerated soils have favoured the formation of ferric (Fe3+)-rich minerals, such as goethite, rather than siderite. Siderite, however, has formed throughout geological history in subsurface environments, such as euxinic seas, where anaerobic organisms created H2-rich conditions. The abundance of large, massive siderite-rich beds in pre-1.8-Gyr sedimentary sequences and their carbon isotope ratios indicate that the atmospheric CO2 concentration was more than 100 times greater than today, causing the rain and ocean waters to be more acidic than today. We therefore conclude that CO2 alone (without a significant contribution from methane) could have provided the necessary greenhouse effect to maintain liquid oceans on the early Earth.     

Seismic identification of along-axis hydrothermal flow on the East Pacific Rise

Hydrothermal circulation at the axis of mid-ocean ridges affects the chemistry of the lithosphere and overlying ocean, supports chemosynthetic biological communities and is responsible for significant heat transfer from the lithosphere to the ocean. It is commonly thought that flow in these systems is oriented across the ridge axis, with recharge occurring along off-axis faults, but the structure and scale of hydrothermal systems are usually inferred from thermal and geochemical models constrained by the geophysical setting, rather than direct observations. The presence of microearthquakes may shed light on hydrothermal pathways by revealing zones of thermal cracking where cold sea water extracts heat from hot crustal rocks, as well as regions where magmatic and tectonic stresses create fractures that increase porosity and permeability. Here we show that hypocentres beneath a well-studied hydrothermal vent field on the East Pacific Rise cluster in a vertical pipe-like zone near a small axial discontinuity, and in a band that lies directly above the axial magma chamber. The location of the shallow pipe-like cluster relative to the distribution and temperature of hydrothermal vents along this section of the ridge suggests that hydrothermal recharge may be concentrated there as a consequence of the permeability generated by tectonic fracturing. Furthermore, we interpret the band of seismicity above the magma chamber as a zone of hydrothermal cracking, which suggests that hydrothermal circulation may be strongly aligned along the ridge axis. We conclude that models that suggest that hydrothermal cells are oriented across-axis, with diffuse off-axis recharge zones, may not apply to the fast-spreading East Pacific Rise.