Thesis (S.M.)--Joint Program in Applied Ocean Science and Engineering (Massachusetts Institute of Technology, Dept. of Ocean Engineering; and the Woods Hole Oceanographic Institution), 2005. / Includes bibliographical references (p. 63-65). / Hydrothermal vents discharge superheated, mineral rich water into our oceans, thereby providing a habitat for exotic chemosynthetic biological communities. Hydrothermal fluids are convected upwards until they cool and reach density equilibrium, at which point they advect laterally with the current. The neutrally buoyant plume layer can have length scales on the order of several kilometers, and it therefore provides the best means to detect the presence of vent fields on the seafloor, which typically have length scales on the order of a few meters. This thesis uses field measurements of the velocity, temperature and particulate anomalies associated with the TAG hydrothermal plume to demonstrate that tidal currents exert a strong impact on the plume shape, and to provide new constraints on the thermal power of the TAG hydrothermal system. The results show that the power output of the TAG system is on the order of 6000 MW, which is up to two orders of magnitude greater than previous estimates, and that there is considerably more entrainment than had previously been assumed. / by Sacha Wichers. / S.M.

Thesis (Ph. D.)--Joint Program in Oceanography/Applied Ocean Science and Engineering (Massachusetts Institute of Technology, Dept. of Mechanical Engineering; and the Woods Hole Oceanographic Institution), 2007. / Includes bibliographical references. / Present-day expeditionary oceanography is beginning to shift from a focus on short-term ship and submersible deployments to an ocean observatory mode where long-term temporally-focused studies are feasible. As a result, a critical need for in situ chemical sensors is evolving. New sensors take a significant amount of time to develop; thus, the evaluation of techniques in the laboratory for use in the ocean environment is becoming increasingly important. Laser-induced breakdown spectroscopy (LIBS) possesses many of the characteristics required for such in situ chemical sensing, and is a promising technique for field measurements in extreme environments. Although many LIBS researchers have focused their work on liquid jets or surfaces, little attention has been paid to bulk liquid analysis, and especially to the effect of oceanic pressures on LIBS signals. In this work, laboratory experiments validate the LIBS technique in a simulated deep ocean environment to pressures up to 2.76 x 10⁷ Pa. A key focus of this work is the validation that select elements important for understanding hydrothermal vent fluid chemistry (Na, Ca, Mn, Mg, K, and Li) are detectable using LIBS. A data processing scheme that accurately deals with the extreme nature of laser-induced plasma formation was developed that allows for statistically accurate comparisons of spectra. The use of both single and double pulse LIBS for high pressure bulk aqueous solutions is explored and the system parameters needed for the detection of the key analytes are optimized. Using both single and double pulse LIBS, the limits of detection were found to be higher than expected as a result of the spectrometer used in this experimentation. However, the results of this validation show that LIBS possesses the characteristics to be a viable chemical sensing method for in situ analyte detection in high pressure environments like the deep ocean. / by Anna Pauline Miranda Michel. / Ph.D.

Au niveau des fonds océaniques, souvent considérés comme des déserts aux conditions de vie extrêmes, des oasis luxuriants de vie existent pourtant. Les sources hydrothermales et les suintements froids, principalement localisés au niveau des dorsales océaniques et des marges continentales, vont permettre le développement de communautés microbiennes et animales très particulières. Le Bassin de Guaymas, situé dans le Golfe de Californie (Mexique) présente la particularité de regrouper à la fois une zone de sources hydrothermales et une zone de suintements froids, situées à une soixantaine de kilomètres l’une de l’autre, et toutes deux recouvertes par une épaisse couche sédimentaire. Ces deux zones sont également colonisées par des étendues de bivalves, des buissons de vers tubicoles ainsi que de tapis microbiens blancs ou colorés comparables. Afin de mieux comprendre le fonctionnement global de ces deux types d’écosystèmes et notamment le rôle structurant des communautés microbiennes sédimentaires sur la répartition des différents assemblages de surface, les travaux entrepris dans cette thèse se proposaient d’étudier les communautés microbiennes sédimentaires associés à la zone de suintements froids (Marge de Sonora) et la zone de sources hydrothermales (Southern Trough) du Bassin de Guaymas. Pour cela, la diversité des communautés microbiennes (Bacteria et Archaea) de différents habitats caractérisés par une faune et des profils géochimiques particuliers, a été étudiée grâce à l’utilisation d’une technique de séquençage haut-débit, le pyroséquençage par la technique du 454, combinée à d’autres techniques comme le FISH ou la PCR quantitative. Cette étude a permis de déterminer que la structure et la diversité des communautés microbiennes dans ces sédiments étaient très spécifiques de ces environnements. Par ailleurs, les colonisateurs présents en surface des sédiments reflétaient des profils géochimiques et des communautés microbiennes très différentes au sein des sédiments. Dans les sédiments colonisés par les tapis microbiens, riches en méthane, les communautés microbiennes dominantes (ANME, Deltaproteobacteria…) utiliseraient notamment les émissions de méthane des couches sédimentaires profondes et produiraient d'importantes concentrations de sulfures, nécessaires à l’installation des communautés microbiennes thiotrophes de surface formant les tapis. Ces fortes concentrations en sulfure excluraient en revanche les communautés animales de ce périmètre. A contrario, dans les sédiments présentant de faibles concentrations en méthane et en sulfure, permettant l'installation d’assemblages faunistiques variés, les communautés microbiennes méthanotrophes anaérobies et productrices de sulfure étaient minoritaires. L'activité et le métabolisme de ces colonisateurs de surface pourraient par ailleurs permettre le développement des lignées microbiennes détectées dans ces habitats (MBG-D, Chloroflexi…). L'analyse des larges jeux de données obtenus au cours de cette étude nous a donc permis de mettre en évidence un système dynamique complexe fonctionnant en équilibre entre les communautés microbiennes sédimentaires, les organismes colonisant la surface du sédiment et la composition géochimique des eaux interstitielles. / Whereas the deep-sea environment is often considered to be a desert, hydrothermal vents andcold seeps provide “oases” of biological activity on the ocean floor. Vent and seep ecosystems support complex food webs based on microbial chemoautotrophic primary production. These hydrothermal vent and cold seeps ecosystems both release hydrocarbon- and sulfide-rich fluids,fueling various surface assemblages such as mat-forming giant bacteria or symbiont-bearinginvertebrates (e.g. bivalves, tubeworms). In the Guaymas Basin, the nearby presence at a few tens of kilometers of cold seeps and hydrothermal vents coupled with comparable sedimentary settings and depths offer a unique opportunity to assess and compare the microbial community composition of these ecosystems. Tobetter understand their overall functioning, we studied sedimentary microbial communities associated with cold seep and hydrothermal vent areas in the Guaymas Basin. The diversity of microbial communities inhabiting sediments was studied using high throughput sequencing (454pyrosequencing), combined with complementary approaches, such as FISH and quantitative PCR. This study reveals that sediments found in the Guaymas Basin were colonized by microbial communities typically found in these types of ecosystems. Our results revealed a high similarity between microbial communities composition associated with the cold seep and hydrothermal vent areas as a probable consequence of the sedimentary context. Nonetheless, thermophilic and hyperthermophilic lineages (e.g.: Thermodesulfobacteria, Desulfurococcales, etc) were exclusively identified in hydrothermally influenced sediments highlighting the strong influence of temperature gradients and other hydrothermally-related factors on microbial community composition. Furthermore, sediments populated by different surface assemblages show distinct porewater geochemistry features and are associated with distinct microbial communities. Indeed, in the sediments underlying microbial mats characterized by high methane porewater concentrations,microbial communities were dominated by anaerobic methane oxidizers (ANME), known to produce sulfide which provides high fluxes of sulfide to the seafloor. In contrast, sediment associated microbial communities underlying faunal assemblages were characterized by a lower biomass and lower methane porewater concentrations in sediments, limiting porewater sulfide concentrations. Without elevated and toxic sulfide concentrations, faunal assemblages can colonize the surface. Together, geochemical and microbial surveys indicate that porewater methane concentrations play an important role in the microbial community structure and subsequently in the establishment of the surface colonizers. Furthermore, presence and activity of the surface colonizers influence the underlying microbial communities probably because of modification of energy source availabilities. Finally, the existence of similar microbial populations between the two ecosystems also raises the question of their dispersal mechanisms. Our results support the hypothesis of a potential continuity among deep-sea ecosystems. In absence of physical borders, environmental conditions (temperature, specific compounds associated withhydrothermal fluids) might select specific and highly adapted microorganisms from the pool of microorganisms dispersed globally across the seafloor.

The current research project investigated the anoxygenic phototrophic and metal(loid) transforming bacteria of hypersaline and deep ocean hydrothermal environments. The East German Creek brine springs, an unusual flowing hypersaline system, was enumerated using classical techniques. Subterranean sulfide supported purple sulfur and nonsulfur bacteria, but at the highly oxygenated surface, aerobic anoxygenic phototrophs (AAP) were numerically dominant (up to 16-36% of cultivable bacteria). Strains (EG8, EG13, EG17, EG19) with unusual phylogenetic affiliation and novel photosynthetic and metal(loid) reducing traits were described taxonomically. Chromocurvus halotolerans gen. nov., sp. nov. was proposed as a second example of a gammaproteobacterial AAP. It exhibited bent rod-shaped cells, unusual among AAP. Facultatively anaerobic Charonomicrobium ambiphototrophicum gen. nov., sp. nov. was capable of both aerobic and anaerobic anoxygenic photosynthesis, and incapable of photoautotrophy, distinguishing it from both AAP and purple nonsulfur bacteria. Roseovarius vanadiphilum sp. nov. surprisingly produced 4.5 times more biomass and 2 times more bacteriochlorophyll (BChl) at extremely high NaVO3 concentration (7.5 g/l) than in metal-free medium. A second novel metabolic mode, anaerobic respiration on the toxic metalloid tellurate, was described for a relative of non-phototrophic Shewanella frigidimarina (ER-Te-48), from deep ocean hydrothermal vent Paralvinella worms at Explorer Ridge in the Pacific Ocean. Other strains respired on SeO32- (ER-Se-17L), VO3- (ER-V-6), and VO43- (AV-V-25). These organisms provided the first examples of anaerobic respiration on Te, Se and V at hydrothermal vents. High level resistance of AAP to metal(loid)s prompted investigation of the influence of TeO32- on photosynthetic pigment production in species including Erythromicrobium ramosum (from a terrestrial hydrothermal system) and Erythrobacter litoralis (from a hypersaline supralittoral system). Tellurite enhanced photosynthetic pigment production up to 3.4 times, consistent with an antioxidant carotenoid-based defense mechanism. However, in E. litoralis BChl precursors such as Mg protoporphyrin or its monomethyl ester also accumulated, indicating biosynthetic pathway interruption. In hydrothermal and hypersaline ecosystems, largely devoid of eukaryotic phototrophs but often enriched in metal(loid)s, AAP and metal(loid) reducers are key modulators of nutrient and toxin availability. The presented results on their ecology, physiology and biochemistry have important implications for theoretical understanding of extreme environments and hold potential for biotechnological applications.

The current research project investigated the anoxygenic phototrophic and metal(loid) transforming bacteria of hypersaline and deep ocean hydrothermal environments. The East German Creek brine springs, an unusual flowing hypersaline system, was enumerated using classical techniques. Subterranean sulfide supported purple sulfur and nonsulfur bacteria, but at the highly oxygenated surface, aerobic anoxygenic phototrophs (AAP) were numerically dominant (up to 16-36% of cultivable bacteria). Strains (EG8, EG13, EG17, EG19) with unusual phylogenetic affiliation and novel photosynthetic and metal(loid) reducing traits were described taxonomically. Chromocurvus halotolerans gen. nov., sp. nov. was proposed as a second example of a gammaproteobacterial AAP. It exhibited bent rod-shaped cells, unusual among AAP. Facultatively anaerobic Charonomicrobium ambiphototrophicum gen. nov., sp. nov. was capable of both aerobic and anaerobic anoxygenic photosynthesis, and incapable of photoautotrophy, distinguishing it from both AAP and purple nonsulfur bacteria. Roseovarius vanadiphilum sp. nov. surprisingly produced 4.5 times more biomass and 2 times more bacteriochlorophyll (BChl) at extremely high NaVO3 concentration (7.5 g/l) than in metal-free medium. A second novel metabolic mode, anaerobic respiration on the toxic metalloid tellurate, was described for a relative of non-phototrophic Shewanella frigidimarina (ER-Te-48), from deep ocean hydrothermal vent Paralvinella worms at Explorer Ridge in the Pacific Ocean. Other strains respired on SeO32- (ER-Se-17L), VO3- (ER-V-6), and VO43- (AV-V-25). These organisms provided the first examples of anaerobic respiration on Te, Se and V at hydrothermal vents. High level resistance of AAP to metal(loid)s prompted investigation of the influence of TeO32- on photosynthetic pigment production in species including Erythromicrobium ramosum (from a terrestrial hydrothermal system) and Erythrobacter litoralis (from a hypersaline supralittoral system). Tellurite enhanced photosynthetic pigment production up to 3.4 times, consistent with an antioxidant carotenoid-based defense mechanism. However, in E. litoralis BChl precursors such as Mg protoporphyrin or its monomethyl ester also accumulated, indicating biosynthetic pathway interruption. In hydrothermal and hypersaline ecosystems, largely devoid of eukaryotic phototrophs but often enriched in metal(loid)s, AAP and metal(loid) reducers are key modulators of nutrient and toxin availability. The presented results on their ecology, physiology and biochemistry have important implications for theoretical understanding of extreme environments and hold potential for biotechnological applications.

Les écosystèmes hydrothermaux profonds hébergent des communautés présentant de fortes biomasses, issues de l’activité chimiotrophique des microorganismes, avec de nombreux exemples d’associations symbiotiques entre ces derniers et les organismes de la mégafaune dominante. La connaissance du cycle de vie de ces espèces, y compris de leurs symbiontes, et de la façon dont elles sont capables de disperser et de coloniser de nouveaux sites est incontournable pour la compréhension du fonctionnement des communautés hydrothermales.Dans cette étude, sont présentées de nombreuses avancées portant sur la distribution, la reproduction, la dispersion et le cycle de vie d’une espèce dominante des écosystèmes hydrothermaux de la dorsale Médio-Atlantique, la crevette alvinocarididé Rimicaris exoculata, et des espèces proches. Les outils méthodologiques utilisés incluent la description morphologique de larves, l’étude de la structure de populations et de leur état de reproduction, des approches moléculaires appliquées à l’identification des espèces via la reconstruction phylogénétique, la génétique populationnelle et l’étude de la diversité bactérienne. La plupart des observations et analyses ont été réalisées grâce aux prélèvements de la mission BICOSE qui s’est déroulée de janvier à février 2014 sur la dorsale Médio-Atlantique.L’analyse morphologique détaillée des premiers stades larvaires (zoé I) de quatre espèces d’Alvinocarididae (R.exoculata, Mirocaris fortunata, Nautilocaris saintlaurentae et Alvinocaris muricola), indique une combinaison de traits caractéristiques de cette famille et unique parmi les crevettes Caridés. Le premier stade larvaire lécithotrophe présente vraisemblablement une durée de développement prolongée, avec une transition vers la planctotrophie au cours des stades ultérieurs. La capture de ces larves près du fond suggère par ailleurs une dispersion bathypélagique. L’étude réalisée sur les populations de R. exoculata des champs hydrothermaux de TAG et Snake Pit met en évidence une ségrégation spatiale des sexes et des stades de vie. Les femelles, les sub-adultes et les juvéniles occupent la paroi des fumeurs actifs, tandis que les mâles se retrouvent majoritairement dispersés à la périphérie inactive des sites. L’identification de plusieurs cohortes d’individus, retrouvées au niveau des habitats des deux champs hydrothermaux indique par ailleurs un recrutement discontinu. Enfin, l’observation, pour la première fois, d’un grand nombre de femelles gravides sur les deux champs hydrothermaux, suggère une reproduction saisonnière, avec quelques différences mineures en terme de fécondité entre les populations des deux champs.Les embryons portés par les femelles jusqu’à l’éclosion des larves sont exposés aux fluides hydrothermaux. Nos résultats, encore partiels, d’analyses par clonage d’assemblages bactériens se développant sur les oeufs au cours de cette phase d’incubation indiquent une spécificité qui pourrait être le reflet d’une fonction symbiotique s’établissant à un stade précoce du cycle de vie de la crevette. La similarité de ces assemblages bactériens avec ceux colonisant le céphalothorax des crevettes adultes, suggère un possible rôle de détoxification et/ou de nutrition.Enfin la découverte, sur TAG, d’importantes « nurseries » de post-larves appartenant à R. chacei, espèce cohabitant avec R. exoculata mais relativement peu abondante, pose la question de l’origine de ce recrutement. Cette question s’inscrit également dans le débat taxonomique récurrent des délimitations d’espèces chez les Alvinocaridiés. Ainsi, de récents travaux de génétique suggèrent que R. chacei pourrait être identique à R. hybisae, une espèce des sites hydrothermaux de la Ride des Caïmans, qui paradoxalement, présente une écologie et un développement symbiotique beaucoup plus similaire à celui de R. exoculata que de R. chacei. Nos analyses de génétique populationnelle et une reconstruction phylogénétique réalisée avec plusieurs gènes […] / Deep-water hydrothermal vent host high-biomass communities based on chemoautotrophy supported by the metabolic activity of free-living and symbiotic bacteria associated to invertebrates, especially megafauna. Knowledge on the mechanisms of dispersal and the life cycle of vent species is essential to our understanding of the vent communities in terms of distribution, structure and temporal variation. In this study, I present some advances regarding the dispersal and life cycle of a dominant species of the Mid-Atlantic Ridge (MAR) vent ecosystems, the alvinocaridid shrimp Rimicaris exoculata, and other related species. The methodological approaches applied include morphological descriptions of larvae, analysis of population biology and reproduction, and molecular genetics for species identification, phylogenetic reconstructions, population genetics and bacterial diversity. Most observations and studies presented here were conducted on samples collected in January-February 2014, during the BICOSE cruise on the MAR.Based on the analysis of Zoea I larvae of four species (R. exoculata, Mirocaris fortunata, Nautilocaris saintlaurentae andAlvinocaris muricola), we conclude that the alvinocaridid first larval stage is lecitotrophic with extended development, allowing large dispersal without external food requirement. A bathypelagical larval dispersal and a shift to a planktotrophic stage during the larval period is proposed. In terms of population biology, collections performed at the TAG and Snake Pit vent fields show variations in the population structure among habitats, according to sex and life stage. Large aggregations of shrimps found close of the vent emission comprise mostly females and young individuals, whereas scattered adults found at the vent periphery were mostly males. Multiple cohorts were found in both vents fields, denoting a discontinuous recruitment. Brooding females were observed in significant numbers close to the vent emission, which contrasts with their constant lack in previous field studies and suggests a seasonal reproduction with a brooding period the winter season. In addition, differences in the reproductive effort were detected between vent fields, including egg number, egg size and proportion of aborted females. The egg surface of R. exoculata is colonized by episymbiotic bacteria. Cloning approaches show that the bacterial assemblages on eggs seem to be specific, suggesting their symbiotic role, and evolve according to the egg development. The bacterial assemblages on eggs and their variation during the embryonic development remind the episymbiotic communities found in the branchial chamber of adults, suggesting similar detoxification or nutrition role. In other Rimicaris species, questions about life cycle, vent connectivity and speciation have been raised recently. Genetic studies suggest that two species with contrasting distribution, morphology and ecology, R. hybisae and R. chacei, are the same species. This question is related also with the source of a massive recruitment of R. chacei found at TAG vent field, despite the low density of adults. Analysis of population genetics and phylogenetic reconstructions with multiple genes show that R. chacei and R. hybisae are separate lineages with recent or undergoing speciation. These species, as R. exoculata and other alvinocaridids, show a genetic population model associated with a migration pool. An extended larval period could contribute to the wide dispersal and high genetic flow between populations. Implications of these findings and perspectives of future research are discussed in terms of additional experiments and field sampling required to characterize the larval period of alvinocaridids, the variations of symbiosis of the different life stages and sexes inhabiting different habitats, the quantitative and functional characterization of the episymbiosis on eggs, and the evolutionary processes associated with the speciation in Rimicaris. / Las emisiones hidrotermales profundas albergan comunidades de elevada biomasa basadas en quimioautotrofía, soportadas por la actividad metabólica de bacterias de vida libre y bacterias simbiontes asociadas a invertebrados marinos, especialmente megafauna. El conocimiento de los mecanismos de dispersión y el ciclo de vida de las especies de ambientes hidrotermales es escencial para comprender los procesos ecológicos de ambientes hidrotermales asociados a la distribución, la estructura comunitaria y la variación temporal. En este estudio, presento algunos avances relacionados a la dispersión y el ciclo de vida de una especie dominante de los sistemas de emision hidrotermal de la dorsal medioatlántica. La aproximaciones metodológicas aplicadas en este estudio incluyen el estudio de la morfología larvaria, el análisis de la biologia poblacional y de la reproducciôn, así como genética molecular con fines de identificación, reconstructión filogenética, genética de poblaciones y análisis de diversidad de bacterias. La mayoría de las observaciones y análisis presentados en el presente estudio fueron realizados con muestras colectadas en enero y febrero de 2014 durante el crucero oceanográfico BICOSE en la dorsal medioatlántica (campos TAG y Snake Pit). El análisis morfológico de la larva Zoea I de cuatro especies (R. exoculata, Mirocaris fortunata, Nautilocaris saintlaurentae y Alvinocaris muricola) permite concluir que el primer estadio larvario de la familia Alvinocarididae es lecitotrôfico con una duración del desarrollo extendida, permitiendo la dispersión a grandes distancias sin requerimiento de una fuente externa de nutrición. Se propone para estas especies una dispersión batipelágica y un cambio a un estadio planctotrófico durante el periodo larvario. En relación a la biología poblacional, fue observada una variación en la estructura poblacional entre hábitats en relación al sexo y el estado de desarrollo.Las agregaciones densas de camarones encontradas cerca de las emisiones hidrotermales están compuestas principales de hembras y juveniles, mientras la mayoría de adultos dispersos encontrados en la periferia de las chimeneas fueron machos. Varias cohortes de tallas fueron identificadas en ambas poblaciones, lo cual denota un reclutamiento discontinuo. Una gran cantidad de hembras ovígeras fueron observadas cerca de la emisión hidrotermal, lo cual contrasta con la casi completa ausencia de hembras ovígeras en muestreos previos y sugiere una reproducción estacional con incubación y desove durante el invierno. La superficie de los huevos de R.exoculata está colonizada por bacterias episimbiontes. Los análisis de clonación muestran que los ensambles bacterianos parecen ser específicos, lo cual sugiere una relación simbiótica. Además estos ensambles cambian en relación al desarrollo embrionario. Los ensambles de bacterias observados en los huevos son similares a las comunidades episimbiontes encontradas en la cámara branquial de los adultos, sugiriendo la ocurrencia de procesos de detoxificación o nutrición similares. En otras especies del género Rimicaris, interrogantes en relación al ciclo de vida, la conectividad entre sistemas hidrotermales y la especiación han surgido recientemente. Estudios genéticos sugieren que dos especies alopátricas y con diferencias en morfología y ecología, R. hybisae y R. chacei representan una especie única. Esta hipótesis se encuentra relacionada además con el origen de un reclutamiento masivo de R. chacei encontrado en el campo TAG, a pesar de la baja densidad de adultos. Análisis de genética poblacional y reconstrucciones filogenéticas utilizando varios genes muestran que R. chacei y R. hybisae son linajes separados producto de una especiación reciente o en proceso. Estas especies, al igual que R. exoculata y otros alvinocarídidos, muestran patrones de conectividad asociados al modelo de migración colectiva (migration pool). Implicaciones de estos hallazgos y perspectivas de futuras […]

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