Notre connaissance de la diversité virale associée aux microorganismes procaryotiques issus des sources hydrothermales océaniques profondes demeure encore limitée. Peu d’études se sont intéressées à l’abondance virale ou à l’impact des virus sur la mortalité microbienne au sein de ces écosystèmes. Le nombre de virus caractérisés, issus de ces environnements, reste faible. Deux virus, PAV1 et TPV1, associés à des archées hyperthermophiles anaérobies appartenant à l’ordre des Thermococcales ont été décrits dans notre laboratoire. Afin de poursuivre nos recherches sur la diversité virale infectant les microorganismes hydrothermaux marins, l’ordre bactérien des Thermotogales a été ciblé. Cet ordre est composé de bactéries chimioorganotrophes anaérobies pour la plupart hyper/thermophiles. Elles partagent la même niche écologique que les Thermococcales et sont métaboliquement proches. De nombreux transferts latéraux de gènes ont, par ailleurs, contribué à l’histoire évolutive des Thermotogales, subodorant l’implication potentielle de virus. La présence de séquences CRISPR a également été rapportée au sein de plusieurs génomes de Thermotogales, suggérant que les Thermotogales sont ou ont certainement déjà été exposées à des infections virales. Pour autant, à ce jour, les seuls éléments génétiques mobiles à avoir réellement été décrits chez les Thermotogales sont 3 miniplasmides et aucun virus. Une cinquantaine de souches de Thermotogales provenant majoritairement de la collection de notre laboratoire (Souchothèque de Bretagne et Collection Ifremer) a été passée au crible quant à la présence d’éventuels bactériovirus associés. A l’issue de ce criblage, des éléments à ADN extra-chromosomiques, incluant 2 plasmides et 7 bactériovirus (du type Siphoviridae) ont été découverts au sein de souches appartenant aux genres Thermosipho et Marnitoga. Des analyses préliminaires ont été réalisées sur ces différents éléments et l’un des nouveaux systèmes hôte/virus a été caractérisé en détail. MPV1 (Marinitoga piezophila virus 1) est un siphovirus-like tempéré isolé d’une bactérie piezophile, il constitue le premier bactériovirus associé à l’ordre des Thermotogales. La souche hôte est piezophile mais aisément cultivable à pression atmosphérique au terme de plusieurs repiquages. Si l’essentiel des analyses a été mené à pression atmosphérique, la production virale s’est avérée tout à fait effective à pression hydrostatique. Nous avons réalisé les analyses de la séquence complète du génome MPV1 (43,7 kb, extrait des capsides virales purifiées) et sa comparaison avec le provirus présent au sein du génome séquencé de Marinitoga piezophila KA3. Les analyses de ce génome viral ont suggéré une proximité évolutive de MPV1 avec les bactériovirus de Firmicutes. Nous avons également mis en évidence que le bactériovirus partage son hôte avec un élément génétique extra chromosomique circulaire de 13,3 kb (pMP1). Ce « ménage à 3 » est surprenant dans le sens où l’élément de 13,3 kb, contenant 13 ORF de fonctions majoritairement inconnues, utilise les capsides virales afin de se propager. Ceci pourrait, ainsi, illustrer un nouvel exemple de piratage moléculaire. / Our knowledge of the viral diversity associated to procaryotic microorganisms inhabiting the deep sea hydrothermal vents is still limited. Only few studies have focused on viral abundance and impact on microbial mortality within these ecosystems. A limited number of viruses from these environments were isolated and characterized. Two viruses, PAV1 and TPV1, associated to hyperthermophilic anaerobic Archaea, Thermococcales order, have ever been described in our laboratory. The topic of this phD thesis was to extend our investigation to other deep sea vent microorganisms in order to deepen our knowledge on the marine hydrothermal virosphere. We decided to focus more precisely on the bacterial order of Thermotogales. This order is composed of anaerobic chemoorganotrophic bacteria that are, for almost, hyper/thermophilic. They share the same ecological niche as the Thermococcales and are metabolically close. Numerous lateral gene transfers have contributed to the evolutionary history of the Thermotogales, implying the potential involvement of viruses. The presence of CRISPRs has also been reported in many genomes, suggesting that Thermotogales certainly are or have been exposed to viral infections. However, up till now, only 3 miniplasmids have been described within Thermotogales and no viruses. Fifty strains of Thermotogales, mostly from the LM2E culture collection (Ifremer and “UBOCC”), were screened for the presence of potential bacteriovirus. Extrachromosomal DNA elements, including 2 plasmids and 7 bacterioviruses (siphovirus-like), were discovered amongst strains belonging to both Thermosipho and Marinitoga genera. Preliminary studies were performed on these elements and one of the new virus-host systems was characterized in details. MPV1 (Marinitoga piezophila virus 1) is a temperate siphovirus-like isolated from a piezophilic bacterium, it is the first bacteriovirus associated to the Thermotogales order. This host strain is piezophilic but easily cultivable at atmospheric pressure after several subcultures. Whether most experiments were performed at atmospheric pressure, the viral production appeared to be effective at hydrostatic pressure. We reported the analyses of the complete sequence of the MPV1 genome (43.7 kb, extracted from purified virions) and its comparison to the provirus present in the sequenced genome of Marinitoga piezophila KA3. Analyses of the viral genome suggested a close evolutionary relationship of MPV1 to Firmicutes bacterioviruses .We also reported that this bacteriovirus shares its host with a circular extrachromosomal genetic element of 13.3 kb (pMP1). This ‘ménage à trois’ is surprising in the sense where the 13.3kb element, that contains 13 ORFs of mostly unknown function, uses the viral capsid to propagate. Therefore, it would likely correspond to a new example of molecular piracy.

Thesis: Ph. D., Joint Program in Oceanography/Applied Ocean Science and Engineering (Massachusetts Institute of Technology, Department of Biology; and the Woods Hole Oceanographic Institution), 2015. / Cataloged from PDF version of thesis. / Includes bibliographical references. / Hydrothermal vents and coral ecosystems are conspicuous biological hot spots in the deep-sea. These ecosystems face increasing threats from human activities. Having thorough taxonomic inventories as well as understanding species' relatedness, genetic diversity, connectivity patterns, and adaptive potential is fundamental for the implementation of conservation strategies that help mitigate these threats. This thesis provides fundamental high-priority knowledge in taxonomic, evolutionary, and ecological aspects of deep-sea coral and vent species, by harnessing the power of genomic tools and overcoming long-standing methodological barriers. First, I develop bioinformatic tools that help guide the design of studies aiming to characterize eukaryotic genome diversity using restriction-site associated DNA sequencing. Using these tools I find that the predictability of restriction site frequencies in eukaryotic genomes is chiefly determined by the phylogenetic position of the target species and the recognition sequence of the selected restriction enzyme. These tools are then applied to test global-scale historical biogeographic hypotheses of vent fauna using barnacles as model. Phylogeographic inferences suggest that the western Pacific was the place of origin of the major vent barnacle lineage, followed by circumglobal colonization eastward along the southern hemisphere during the Neogene. I suggest that the geological processes and dispersal mechanisms discussed here can explain distribution patterns of many other marine taxa in addition to barnacles. Regional-scale analyses indicate that vent barnacle populations are well connected within basins and ridge systems, and that their diversity patterns do not conform to the predictions from the hypothesis that seamounts are centers of endemism. I then move on to resolve long-standing questions regarding species definitions in recalcitrant deep-sea coral taxa, by unambiguously resolving evolutionary relationships and objectively inferring species boundaries. Finally, I explore the adaptive potential of deep-sea coral species to environmental changes by examining a case of adaptation to shallow water from the deep sea. Candidate positive-selection markers shared between pairs of shallow and deep populations are identified as likely makers for genomic regions involved in adaptation. Overall, the results from this thesis constitute critical baseline data with which to assess potential effects of anthropogenic disturbances on deep-sea ecosystems. / by Santiago Herrera Monroy. / Ph. D.

The bioavailability of iron is limited by the solubility of Fe(III) at circumneutral pH. In the High Nutrient-Low Chlorophyll (HNLC) zones of the ocean, the natural or anthropogenic addition of iron stimulates primary productivity and consumes carbon dioxide. As a result, iron fertilization has been proposed to mitigate anthropogenic carbon emissions and lower global temperatures. The natural sources of iron to the ocean are not fully constrained and include eolian depositions as well as inputs from continental shelf sediments, rivers, hydrothermal vents, and icebergs. Regardless of their source, the effectiveness of iron additions in promoting carbon fixation depends on the presence of organic ligands either natural or produced by microorganisms that stabilize or solubilize Fe(III) at neutral pH. For example, siderophores are well known to be expressed extracellularly by prokaryotes in the photic zones of the oceans to increase the bioavailability of iron. In this dissertation, the production of iron nanoparticles is demonstrated in vent fluids from the 90 North hydrothermal system. These iron nanoparticles may either catalyze the oxidation of sulfide to thiosulfate and produce a potential electron acceptor for microbial respiration or provide a source of iron that stimulates primary production at great distances from the hydrothermal vents. In addition, dissolved iron under the form of soluble organic-Fe(III) complexes is demonstrated to constitute a significant source of iron in estuarine sediments that receive large amounts of particulate iron from flocculation and precipitation at the salinity transition of this estuary. A novel competitive ligand equilibration absorptive cathodic stripping voltammetry (CLE-ACSV) technique reveals that the speciation of iron changes from largely colloidal or particulate in the upper estuary to truly dissolved organic-Fe(III) in the lower estuary. It is also demonstrated that organic-Fe(III) complexes are produced far below the sediment-water interface, suggesting that dissimilatory iron-reducing bacteria may play an important role in their production. These complexes then diffuse across the sediment-water interface and provide a significant source of iron to the continental shelf. The mechanism of reduction of iron oxides by iron-reducing bacteria is not fully understood and presents a unique physiological problem for the organism, as the terminal reductase has to transfer electrons to a solid electron acceptor. In this dissertation, it is demonstrated for the first time using random mutagenesis that the respiration of solid Fe(III) oxides by Shewanella oneidensis, a model iron-reducing prokaryote, first proceeds through a non-reductive dissolution step involving organic ligands that are released extracellularly by the cells. These soluble complexes are then reduced by the organism to produce Fe(II) and recycle the ligand for additional solubilization. Incubations with deletion mutants of the proteins involved in the respiration of Fe(III) revealed that the type-II secretion system, which translocates proteins on the outer membrane of gram-negative bacteria, is involved in the production of organic-Fe(III) complexes by secreting an endogenous iron-solubilizing ligand or a protein involved in the biosynthesis of this ligand on the outer membrane. In addition, periplasmic decaheme cytochromes produced by Shewanella appear to be involved in the mechanism of production of the endogenous organic ligand either directly or through a sensing mechanism that controls its production. In turn, two decaheme cytochromes positioned on the outer-membrane and hypothesized to be involved in the electron transfer to the mineral surface do not appear to be involved in the solubilization mechanism, suggesting either that the cells regulate the ligand production via periplasmic sensing systems or that these cytochromes are not involved in the solubilization mechanism. Altogether this research shows the production of organic-Fe(III) complexes in sediments generates a significant flux of dissolved iron to support primary production in continental shelf waters and that these complexes may be partly produced by iron-reducing bacteria. Indeed, experiments with a model organism demonstrate dissimilatory iron reducing bacteria produce endogenous organic ligands with high iron-binding constants to non-reductively solubilize iron oxides during the anaerobic respiration of iron oxides. The organic ligand is apparently recycled several times to minimize the energy cost associated with its biosynthesis. These findings demonstrate that the solubilization of iron oxides by organic ligands may be an important, yet underappreciated process in aquatic systems.

Nos connaissances sur la diversité virale associée aux micro-organismes présents dans les sources hydrothermales océaniques profondes restent encore limitées. Seules quelques études concernant l’abondance virale et l’impact de ceux-ci sur la mortalité microbienne dans ces écosystèmes sont disponibles. En effet, seuls 6 bactériovirus et 2 archéovirus provenant de ces écosystèmes ont été caractérisés à l’heure actuelle. Les deux archéovirus infectent des archées anaérobies hyperthermophiles appartenant à l’ordre des Thermococcales et ont été décrits au laboratoire.Afin d’étendre nos connaissances sur la diversité virale associée aux micro-organismes colonisant ces environnements, il a été décidé d’élargir les recherches à l’ordre bactérien des Thermotogales. Cet ordre bactérien est composé de bactéries chimio-organotrophes anaérobies en majorité thermophiles ou hyperthermophiles. De nombreux transferts latéraux de gènes ont contribué à l’histoire évolutive des Thermotogales supposant une forte implication des virus dans celle-ci. Ces travaux de thèse ont permis la caractérisation fonctionnelle et génomique de deux nouveaux siphovirus, MCV1 et MCV2, infectant deux souches de Marinitoga camini. Ces souches ont été isolées de deux sites hydrothermaux profonds (Menez Gwen et Lucky Strike) au niveau de la dorsale médio-atlantique. Ces virus mettent en oeuvre un cycle lysogénique avec une production basale sans induction relativement haute (>107 virions/ml). Une comparaison de ces deux génomes viraux à celui de MPV1, virus précédemment isolé de Marinitoga piezophila, a été réalisée, révélant la présence de nombreuses similarités. Un core genome de 35 ORFs partagé par ces trois génomes a été identifié, incluant des protéines impliquées dans le métabolisme de l’ADN, l’assemblage des virions et le cycle lysogénique. Des protéines hypothétiques ont aussi été identifiées parmi ces gènes communs, elles portent donc probablement des fonctions importantes pour ces bactériovirus. Par ailleurs, 60% des gènes de ces virus ayant une correspondance dans les bases de données, après exclusion des Thermotogales, partagent des similarités avec lesFirmicutes et les bactériovirus qui leurs sont associés. Le génome d’une autre Thermotogales, Thermosipho sp. AT1244-VC14 a été étudié ainsi que son système CRISPR-cas. Ces résultats indiquent que cette souche, qui porte un système CRISPR-cas qui semble complet et fonctionnel, a probablement déjà été infectée par MCV1, MCV2 ou un virus similaire. Ces travaux permettent d’étendre nos connaissances sur les virus portés par les bactéries du phylum Thermotogae, encore peu décrits à ce jour. Les éléments génétiques mobiles associés à ce phylum sont particulièrement intéressants car ils ont probablement eu un impact important dans l’évolution de ces communautés microbiennes ainsi que dans leur adaptation aux conditions physico-chimiques extrêmes et fluctuantes présentes dans les écosystèmes qu’elles colonisent. / Our knowledge of the viral diversity associated to microorganisms inhabiting the deep-sea hydrothermal vents is still limited. Only a few studies have focused on viral abundance and impact on microbial mortality within these ecosystems. A limited number of viruses (6 bacterioviruses and 2 archaeoviruses) were isolated from these environments and characterized. Two viruses associated to hyperthermophilic anaerobic Archaea, from the Thermococcales order, have been described in our laboratory. In order to deepen our knowledge on the viral diversity of these extreme environments, we have extended our investigation to the bacterial order of Thermotogales. This order is composed of anaerobic chemoorganotrophic bacteria that are, for the most part, hyper/thermophilic. Numerous lateral gene transfers have contributed to the evolutionary history of the Thermotogales, implying the potential involvement of viruses. Here, we will report the characterization of two new siphoviruses MCV1 and MCV2 that infect two strains of Marinitoga camini. Those bacterial strains were isolated from two deep-sea hydrothermal vents sites (Menez Gwen and Lucky strike) in the Mid Atlantic Ridge. These viruses are temperate with a high basal production of virions (>107 virions/mL). Comparative genomics with MPV1, a virus isolated from M. piezophila, was performed and show that those bacterioviruses share numerous similarities. A set of “core genes” shared by all these three viruses was identified and includes proteins involved in DNA metabolism, head and tail assembly and lysogenic cycle. Shared hypothetical proteins were also identified, suggesting that these unknown proteins probably provide important functions for these viruses. Interestingly, for genes with blastp matches in Genbank, over 60% have their top matches, outside the Thermotoga, to genes from Firmicutes and bacterioviruses associated to Firmicutes. We also analyzed the genome of Thermosipho sp. 1244 and studied his CRISPR-cas system. Our results indicated that thisThermosipho strain, with a complete and functional CRISPR-cas system, had already been infected by MCV1, MCV2 or a similar virus. The analyses presented here extend our knowledges about these newly discovered viruses in the deeply branching bacterial phylum Thermotogae. This bacterial order and associated mobile genetic elements are significant for addressing long-term evolutionary adaptation to fluctuant and extreme physicochemical conditions.

Thesis (Ph. D.)--Joint Program in Oceanography (Massachusetts Institute of Technology, Dept. of Earth, Atmospheric, and Planetary Sciences; and the Woods Hole Oceanographic Institution), 1988. / Includes bibliographical references (p. 145-152). / by Sarah Alden Little. / Ph.D.

Thesis: Ph. D., Joint Program in Oceanography/Applied Ocean Science and Engineering (Massachusetts Institute of Technology, Department of Earth, Atmospheric, and Planetary Sciences; and the Woods Hole Oceanographic Institution), 2018. / Cataloged from PDF version of thesis. / Includes bibliographical references. / This thesis examines abiotic processes controlling the transformation and distribution of carbon compounds in seafloor hydrothermal systems hosted in ultramafic rock. These processes have a direct impact on carbon budgets in the oceanic lithosphere and on the sustenance of microorganisms inhabiting hydrothermal vent ecosystems. Where mantle peridotite interacts with carbon-bearing aqueous fluids in the subseafloor, dissolved inorganic carbon can precipitate as carbonate minerals or undergo reduction by H2(aq) to form reduced carbon species. In Chapters 2 and 3, I conduct laboratory experiments to assess the relative extents of carbonate formation and CO2 reduction during alteration of peridotite by C02(aq)-rich fluids. Results from these experiments reveal that formation of carbonate minerals is favorable on laboratory timescales, even at high H2(aq) concentrations generated by serpentinization reactions. Although CO2(aq) attains rapid metastable equilibrium with formate, formation of thermodynamically stable CH4(aq) is kinetically limited on timescales relevant for active fluid circulation in the subseafloor. It has been proposed that CH4 and potentially longer-chain hydrocarbons may be sourced, instead, from fluid inclusions hosted in plutonic and mantle rocks. Chapter 4 analyzes CH4-rich fluid inclusions in olivine-rich basement rocks from the Von Damm hydrothermal field and the Zambales ophiolite to better understand the origin of abiotic hydrocarbons in ultramaficinfluenced hydrothermal systems. Comparisons of hydrocarbon abundances and stable isotopic compositions in fluid inclusions and associated vent fluids suggest that fluid inclusions may provide a significant contribution of abiotic hydrocarbons to both submarine and continental serpentinization systems. / by Niya Gueorguieva Grozeva. / Ph. D.

Les crevettes Alvinocarididae sont emblématiques des sources hydrothermales de la Dorsale Médio-Atlantique, mais les mécanismes qui leur permettent de détecter leur habitat sont énigmatiques. Il est supposé que les signatures chimique et thermique du fluide hydrothermal leur servent de repères pour s’orienter. Les facultés chimio- et thermosensorielles de l’espèce hydrothermale Mirocaris fortunata et de l’espèce côtière Palaemon elegans ont été étudiées avec plusieurs approches. Des traits structuraux du système sensoriel périphérique et central ont été comparés pour inférer sur les facultés olfactives de chaque espèce. L’expression du récepteur ionotropique IR25a dans les organes chimiosensoriels a été mesurée. Une technique d’électroantennographie a été mise au point afin de tester la détection de composés chimiques de fluide hydrothermal par les crevettes. Plusieurs expériences à pression atmosphérique et in situ ont été réalisées afin d’étudier le comportement des espèces hydrothermale et côtière exposées à une odeur de nourriture, du sulfure ou des températures relativement chaudes. L’ensemble des résultats montre que M. fortunata présente un système chimio- et thermosensoriel fonctionnel et apporte des bases substantielles pour approfondir les connaissances sur les adaptations sensorielles en milieu hydrothermal profond. / Alvinocaridid shrimp are an emblematic taxon at hydrothermal vents on the Mid-Atlantic Ridge, but how they locate active vents and detect their habitat is still enigmatic. They might use the chemical and thermal signatures of the hydrothermal fluid as orientation cues. The chemo- and thermosensory abilities of the hydrothermal species Mirocaris fortunata and the coastal species Palaemon elegans were investigated using various approaches. Structural features of the peripheral and central nervous system were used as rough estimates of their olfactory abilities. The co-ionotropic receptor IR25a, involved in chemodetection, was identified and located in the antennal appendages. A new electroantennography method was developed to test the detection of hydrothermal fluid chemicals. Several attraction tests were conducted at atmospheric and in situ pressure to investigate the behavior of the vent and shallow-water species when exposed to a food odor, sulfide and warm temperatures. Altogether, these results provide advances in the knowledge of the chemosensory biology of vent shrimp, and a substantial background for future researches on sensory adaptations to the hydrothermal environment.

Thesis: Ph. D., Joint Program in Oceanography/Applied Ocean Science and Engineering (Massachusetts Institute of Technology, Department of Earth, Atmospheric, and Planetary Sciences; and the Woods Hole Oceanographic Institution), 2019 / Cataloged from PDF version of thesis. / Includes bibliographical references. / Nitrogen (N) species in hydrothermal vent fluids serve as both a nutrient and energy source for the chemosynthetic ecosystems surrounding deep-sea vents. While numerous pathways have been identified in which N-species can be produced and consumed in the context of submarine hydrothermal vent systems, their exact nature has been largely limited to interpretation of variations in concentrations. This thesis applies stable isotope approaches to further constrain the sources and fate of N-species in deep-sea vents across a variety of geological settings. First, I discuss isotope fractionation and reaction kinetics during abiotic reduction of nitrate (NO₃⁻) to ammonium ([sigma]NH₄⁺ = NH₃+NH₄⁺) under hydrothermal conditions. Results of lab experiments conducted at high temperatures and pressures revealed a wide degree of N isotope fractionation as affected by temperature, fluid/rock ratio, and pH-all which exert control over reaction rates. / Moreover, a clear pattern in terms of reaction products can be discerned with the reaction producing [sigma]NH₄⁺ only at high pH, but both [sigma]NH₄⁺ and N₂ at low pH. This challenges previous assumptions that NO₃⁻ is always quantitatively converted to NH₄⁺ during submarine hydrothermal circulation. Next, I report measurements of [sigma]NH₄⁺ concentrations and N isotopic composition ([delta]¹⁵N[subscript NH4]) from vent fluid samples, together with the largest compilation to date of these measurements made from other studies of deep-sea vent systems for comparison. The importance of different processes at sediment-influenced and unsedimented systems are discussed with a focus on how they ultimately yield observed vent [sigma]NH₄⁺ values. / Notable findings include the role that phase separation might play under some conditions and a description of how an unsedimented site from Mid-Cayman Rise with unexpectedly high NH4+ may be uniquely influenced by N₂ reduction to [sigma]NH₄⁺. Lastly, I explore [sigma]NH₄⁺ dynamics in the context of low-temperature vent sites at 9°50'N East Pacific Rise to investigate dynamics of microbially-mediated N transformations. Through both measurements of natural samples, as well as isotopic characterization of N species from incubation experiments and model simulations thereof, an exceptionally high variability observed in [delta]¹⁵N[subscript NH4] values emphasizes the complexity of these microbe-rich systems. / In sum, this thesis highlights the role of microbial processes in low temperature systems, demonstrates a more mechanistic understanding of lesser-understood abiotic N reactions and improves the coverage of available data on deep-sea vent [sigma]NH₄⁺ measurements. / by Chawalit "Net" Charoenpong. / Ph. D. / Ph.D. Joint Program in Oceanography/Applied Ocean Science and Engineering (Massachusetts Institute of Technology, Department of Earth, Atmospheric, and Planetary Sciences; and the Woods Hole Oceanographic Institution)

En bientôt quarante ans de recherche, de nombreuses connaissances ont été acquises sur la géologie des champs hydrothermaux, la chimie des fluides qui en réchappent et l’écologie des communautés qui les habitent. Celles-ci s’organisent en assemblages denses, distribués le long de la zone de dilution du fluide hydrothermal dans l’eau de mer, et dominés visuellement par une poignée d’espèces. La forte variabilité spatio-temporelle du fluide hydrothermal a une forte influence sur la distribution des communautés. Cependant, les mécanismes à l’origine de la réponse de la faune à cette variabilité sont peu compris. Pour cela, une approche modélisatrice est présentée. Les données collectées pendant plus de 20 ans sur l’édifice Tour Eiffel, sur le champ hydrothermal Lucky Strike (ride médio-Atlantique) ont été intégrées afin d’en extraire les composantes principales. L’étude intégrative des biomasses sur l’édifice montre que celles-ci sont largement dominées par la modiole Bathymodiolus azoricus. Ce bivalve est susceptible d’avoir une influence importante sur le fonctionnement de l’écosystème, et fait donc l’objet d’un premier modèle. La recherche de données pour le contraindre ont mené à mesurer des taux métaboliques in situ. Une fois le modèle paramétré, le modèle a fourni des estimations de flux encore inconnu. La simulation d’interruption du flux hydrothermal a fourni des indices sur la réponse de la modiole à la variabilité de son environnement. / In nearly forty years of research, significant insights have been gained on vent field geology, on the chemistry of emitted fluid and on the ecology of the communities inhabiting hydrothermal ecosystems. The fauna forms dense assemblages, distributed along the hydrothermal fluid/sea water mixing gradient, and visually dominated only by a few species. The high spatio-temporal variability of the hydrothermal fluid has a strong influence on species distribution. However, the mechanisms determining the species response to this variability is still poorly understood. In order to investigate this issue, a modelling approach is presented. Data collected for more than 20 years on the Eiffel Tower edifice, on the Lucky Strike vent filed (Mid-Atlantic Ridge) were integrated in order to identify meaningful elements for our problem. An integrative study of the faunal biomasses on the edifice showed that these are dominated by the mussel Bathymodiolus azoricus. This bivalve is likely to have a significant influence on the ecosystem functioning and is thus the object of a first model. The search for data to constrain it led to the measurement of in situ metabolic rates. Once parametrized, the model provided quantitative estimates of unknown fluxes. The simulation of hydrothermal flow interruption provided some insights on the mussel biomass response to its environment variability.

Nous avons étudié l’écologie microbienne d’un site hydrothermal alcalin sous-marin peu profond (0-50 m) localisé dans la baie de Prony (PHF) dans le sud de Nouvelle-Calédonie (Pacifique Sud-Ouest) comparable par son fonctionnement au site hydrothermal alcalin profond (800m) de Lost City (LCHF) (Dorsale médio-atlantique). PHF au même titre que LCHF est un système hydrothermal ultramafique associé à des réactions de serpentinisation des roches du manteau terrestre, libérant des fluides anoxiques très alcalins (jusqu'à pH 11) riche en calcium, en hydrogène et en méthane dissous. Le site hydrothermal de Prony (PHF) est caractérisé par de grandes cheminées de carbonate émettant des fluides dont la température n’excède pas 40°C. Les approches moléculaires ont démontré une prédominance des bactéries (Firmicutes, Deltaproteobacteria …) sur les archées (Methanosarcinales). En ce qui concerne les mises en culture, elles sont restées vaines pour la plupart des grands groupes trophiques recherchés (bactéries sulfato-réductrices, archées méthanogènes) excepté pour les fermentaires relevant du phylum des Firmicutes. Deux nouvelles bactéries appartenant à ce phylum ont été isolées. Il s’agit (i) de Vallitalea pronyensis vraisemblablement associée aux cheminées hydrothermales et (ii) d’Alkaliphilus hydrothermalis qui serait plutôt indigène aux fluides alcalins émis par les cheminées si l’on en juge par ses conditions physico-chimiques optimales de croissance en adéquation avec celles des fluides. Ces deux microorganismes représentent les premiers anaérobies stricts isolés de systèmes hydrothermaux alcalins serpentinisés à ce jour. / We studied the microbial ecology of an alkaline hydrothermal submarine shallow field (0-50 m) located in Prony Bay (PHF) in the south of New Caledonia (SW Pacific) similar to the deep alkaline hydrothermal site (800m) of Lost City (LCHF) (Mid-Atlantic Ridge). Similarly to LCHF, PHF is an ultramafic hydrothermal system functioning on the basis of serpentinization reactions of the mantle rocks, releasing anoxic, highly alkaline fluids (to pH 11) rich in calcium, and in dissolved hydrogen and methane. The Prony hydrothermal field (PHF) is characterized by large carbonate chimneys emitting fluids with temperatures not exceeding 40 °C. Molecular approaches revealed a prevalence of Bacteria (Firmicutes, Deltaproteobacteria…) over Archaea (Methanosarcinales). Regarding microbial cultures, they were unsuccessful for most major trophic groups (sulfate-reducing bacteria, methanogens) with the exception of fermentative representatives of the phylum Firmicutes. Two novel bacteria belonging to this phylum were isolated. They include (i) Vallitalea pronyensis which is likely associated with hydrothermal vents and (ii) Alkaliphilus hydrothermalis which should be indigenous to alkaline fluids emitted from chimneys since its growth optimal physicochemical conditions match those of fluids. These two bacteria represent the first anaerobic microorganisms isolated from alkaline hydrothermal serpentinized systems so far.