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Les coccolithophoridés sont des microalgues unicellulaires calcifiantes qui jouent un rôle important dans le cycle du carbone océanique via leurs processus cellulaires de photosynthèse (puits de CO2) et de calcification (source de CO2). L'espèce la plus abondante dans l'océan moderne est l'espèce cosmopolite Emiliania huxleyi, qui est caractérisée par une forte tolérance à de larges gammes de température, d'irradiance et de concentrations en nutriments. La distribution d'E. huxleyi qui s'étend des régions subarctiques aux régions subéquatoriales et des eaux eutrophes aux eaux oligotrophes en fait l'espèce de coccolithophoridés la plus étudiée jusqu'à présent. Cependant, sa réponse physiologique à des conditions environnementales clés comme la co-limitation en lumière et en nutriments reste peu étudiée, que ce soit in situ, où ces conditions sont rencontrées dans les niches profondes de coccolithophoridés des gyres oligotrophes, ou en laboratoire. J'ai ainsi réalisé des expériences de culture et une approche de modélisation numérique afin d'étudier la réponse physiologique d'E. huxleyi en conditions de limitation en nutriments et en lumière, avec pour objectif d'améliorer notre compréhension du contrôle environnemental des niches profondes de coccolithophoridés. / Coccolithophores are unicellular calcifying marine algae that play an important role in the oceanic carbon cycle via their cellular processes of photosynthesis (a CO2 sink) and calcification (a CO2 source). The most abundant coccolithophore species in the modern ocean is the cosmopolitan species Emiliania huxleyi that is characterized by a strong tolerance to a wide range of light, nutrient and temperature conditions. The distribution of Emiliania huxleyi from subarctic to subequatorial regions and from eutrophic to oligotrophic waters makes it the most widely studied coccolithophore species. However, its physiologic response under key environmental conditions such as the co-limitation of light and nutrients remains poorly investigated, both in the laboratory and in the field, such as in deep niches of coccolithophores in oceanic gyres. I conducted laboratory culture and numerical modeling experiments to understand the controls on the physiology of Emiliania huxleyi in low-nutrient and low-light conditions, with the aim of better understanding environmental controls on deep ecological niches of coccolithophores.

Human activities have contributed to the increase in atmospheric greenhouse gases such as carbon dioxide (CO2). This anthropogenic gas emission has led to a rise in the average Earth temperature. Moreover, the ocean constitutes the major sink for anthropogenic CO2 and its dissolution in surface waters has already resulted in an increase of seawater acidity since the beginning of the industrial revolution. This is commonly called ocean acidification. The increase in water temperature could induce modifications of the physical and chemical characteristics of the ocean. Also, the structure and the functioning of marine ecosystems may be altered as a result of ocean acidification. Phytoplankton productivity is one of the primary controls in regulating our climate, for instance via impact on atmospheric CO2 levels. Coccolithophores, of which Emiliania huxleyi is the most abundant species, are considered to be the most important pelagic calcifying organisms on Earth. Coccolithophores are characterized by calcium carbonate platelets (coccoliths) covering the exterior of the cells. They form massive blooms in temperate and sub-polar oceans and in particular along continental margin and in shelf seas. The intrinsic coupling of organic matter production and calcification in coccolithophores underlines their biogeochemical importance in the marine carbon cycle. Both processes are susceptible to change with ocean acidification and warming. Coccolithophores are further known to produce transparent exopolymer particles (TEP) that promote particle aggregation and related processes such as marine snow formation and sinking. Thus, the impact of ocean warming and acidification on coccolithophores needs to be studied and this can be carried out through a transdisciplinary approach. The first part of this thesis consisted of laboratory experiments on E. huxleyi under controlled conditions. The aim was to estimate the effect of increasing water temperature and acidity on E. huxleyi and especially on the calcification. Cultures were conducted at different partial pressures of CO2 (pCO2); the values considered were 180, 380 and 750 ppm corresponding to past, present and future (year 2100) atmospheric pCO2. These experiments were conducted at 13°C and 18°C. The cellular calcite concentration decreases with increasing pCO2. In addition, it decreases by 34 % at 380 ppm and by 7 % at 750 ppm with an increase in temperature of 5°C. Changes in calcite production at future pCO2 values are reflected in deteriorated coccolith morphology, while temperature does not affect coccolith morphology. Our findings suggest that the sole future increase of pCO2 may have a larger negative impact on calcification than its interacting effect with temperature or the increase in temperature alone. The evolution of culture experiments allows a better comprehension of the development of a bloom in natural environments. Indeed, in order to predict the future evolution of calcifying organisms, it is required to better understand the present-day biogeochemistry and ecology of pelagic calcifying communities under field conditions. The second part of this dissertation was dedicated to results obtained during field investigations in the northern Bay of Biscay, where frequent and recurrent coccolithophorid blooms were observed. Cruises, assisted by remote sensing, were carried out along the continental margin in 2006 (29 May – 10 June), 2007 (7 May – 24 May) and 2008 (5 May – 23 May). Relevant biogeochemical parameters were measured in the water column (temperature, salinity, dissolved oxygen, Chlorophyll-a and nutrient concentrations) in order to determine the status of the bloom at the time of the different campaigns. Calcification has been shown to be extremely important in the study area. In addition, TEP production was significant at some stations, suggesting that the northern Bay of Biscay could constitute an area of important carbon export. Mortality factors for coccolithophores were studied and the first results of lysis rates measured in this region were presented. Results obtained during culture experiments and comparison with data reported in the literature help to better understand and to predict the future of coccolithophores in a context of climate change. Data obtained during either culture experiments or field investigations allowed a better understanding of the TEP dynamics. Finally, the high lysis rates obtained demonstrate the importance of this process in bloom decline. Nevertheless, it is clear that we only begin to understand the effects of global change on marine biogeochemistry, carbon cycling and potential feedbacks on increasing atmospheric CO2. Thus, further research with a combination of laboratory experiments, field measurements and modelling are encouraged.

Coccolithophores are single-celled marine phytoplankton, which produce intricate calcium carbonate platelets or ‘coccoliths’. Emiliania huxleyi is the most abundant and widespread coccolithophore, and is one of the most productive calcifying species on earth, playing a key role in global carbon, carbonate and sulphur cycles. Despite much research into coccolithophore biology, the underlying function of their coccoliths is still unknown. The main aim of the research reported in this thesis was to examine the impact of calcification on metabolism in coccolithophores. Calcification is a significant global process, so it is important to discover what effect it has on the metabolism of cells. The major metabolites each have different costs and benefits to the cell, which will vary depending on the habitat and environmental conditions the cell is in. By comparing the metabolite profiles of different strains, including calcifying, non-calcifying, haploid and diploid cells, differences in metabolite composition and potential patterns related to cell type were investigated. Low molecular weight (LMW) metabolites were characterised using a combination of metabolomic techniques. In agreement with previous research, dimethylsulphoniopropionate (DMSP) was the most abundant compound, followed by mannitol and glycine betaine (GBT). Less abundant sugars, polyols and amino acids were also identified. Environmental factors were manipulated to investigate how the principal metabolites were affected by salinity, different light intensities and nutrient (phosphate and nitrate) limitation. The data revealed a striking difference between haploid and diploid cells of the same strain, with the haploid containing lower concentrations of most of the major metabolites. Thus it is proposed that haploid cells have a different osmoregulatory strategy from the diploid cells. A negative correlation was found between DMSP and mannitol, suggesting that mannitol has a dual function, not only as a major storage compound but also as a principal compatible solute. Untargeted metabolite profiling is becoming a popular tool to investigate phenotypes and varying environmental conditions. LC-ESI-QTOF-MS/MS analyses of a wide range of metabolites showed that it is an effective method to identify differences and similarities between E. huxleyi strains grown in different conditions. Strain and growth phase appear to be the more important factors in differentiating metabolite profiles. Surprisingly there were no obvious metabolite profiling differences between calcifying and non-calcifying cells. Untargeted analysis can, however, be used to identify the compounds that did display differences, and which may be important biomarkers, so warrant further investigation. A range of metabolite profiling techniques highlighted important differences between strains, which will hopefully lead onto further research into the metabolome of E. huxleyi, and the unravelling of important metabolic pathways. There has been little research into the LMW metabolites of E. huxleyi, and especially comparisons between strains. Thus the use of metabolomics is a novel way to investigate the difference between cell types and the possible functions of calcification.

Coccolithophores are unicellular microscopic algae (Haptophyta) surrounded by calcium carbonate plates that are produced during their life cycle. These species, whose contemporary contributor is Emiliania huxleyi, are mainly found in the sub-polar and temperate oceans, where they produce huge blooms visible from space. Coccolithophores are sensitive to ocean acidification that results from the ongoing accumulation of anthropogenic carbon dioxide (CO2) in the atmosphere. The response of these organisms to global change appears to be related to the reduction of their ability to produce calcium carbonate at the cellular level. At the community levels, one anticipates changes in the carbon fluxes associated to their blooms as calcification is reduced. However, the consequences of such environmental changes on this species are speculative and require improvements in the description of the mechanisms controlling the organic and inorganic carbon production and export. The first aspect of this work was to study the response of these organisms to artificially modified CO2 concentrations representative of the conditions occurring in the past (glacial) and those expected by the end of the century (2100). Two different levels were examined: the continuous monospecific cultures (chemostats) allowed us to work at the cellular level while the mesocosms gave light to the mechanisms taking place in an isolated fraction of the natural community. The second aspect of this work consisted of field studies carried out during four cruises (2002, 2003, 2004 and 2006) in the northern Bay of Biscay, where the occurrence of E. huxleyi blooms were observed in late spring and early summer. We describe the vertical profiles of biogeochemical variables (nutrients, chlorophyll-a, dissolved inorganic chemistry, particulate carbon, transparent exopolymer particles (TEP)) and study processes such as primary production, calcification and bacterial production. The properties of these blooms are compared with those reported in the literature and enriched with original measurements such as the abundance and concentration of TEP that could play an important role in carbon export to the deep ocean, modifying the properties of the settling ballasted aggregates.

Cette étude a été motivée par la nécessité de pouvoir décrire la variabilité biogéochimique avec la même résolution que les paramètres physiques. Des profileurs optiques ont permis de mesurer à haute fréquence les propriétés optiques inhérentes dans la colonne d'eau. Par comparaison avec des données discrètes acquises simultanément à nos mesures, les coefficients optiques à des longueurs d'onde précises ont été convertis en grandeurs biogéochimiques quantitatives. Nous avons estimé la concentration en chlorophylle a et en carbone particulaire total ainsi que la répartition verticale de la matière organique dissoute colorée (CDOM). La nature qualitative du matériel particulaire et dissous a été abordée en combinant différents coefficients optiques ainsi qu'en analysant leur dépendance spectrale. En préalable aux campagnes dans l'Atlantique nord (campagnes POMME), une expérience en milieu contrôlé a été menée sur une population d'organismes phytoplanctoniques calcifiants (Emiliania huxleyi) afin de caractériser ses propriétés biooptiques ainsi que leurs modifications consécutives à une augmentation de la pCO2 atmosphérique du niveau actuel (360 ppm) au niveau prévu pour la fin du siècle (700 ppm). Lors des campagnes entre le Portugal continental et les Açores nous avons effectué le suivi biooptique de la variabilité à méso-échelle à trois périodes clés de l'année : fin de l'hiver, printemps et fin de l'été. L'observation à haute résolution de la colonne d'eau a permis de mettre en évidence certaines relations entre les caractéristiques hydrologiques et biogéochimiques. Le déploiement de nos instruments le long de radiales avec un système tracté autorisant une résolution de 2 milles (Tow-Yo) a conduit à préciser l'influence des structures tourbillonnaires de moyenne échelle.

Human activities have contributed to the increase in atmospheric greenhouse gases such as carbon dioxide (CO2). This anthropogenic gas emission has led to a rise in the average Earth temperature. Moreover, the ocean constitutes the major sink for anthropogenic CO2 and its dissolution in surface waters has already resulted in an increase of seawater acidity since the beginning of the industrial revolution. This is commonly called ocean acidification. The increase in water temperature could induce modifications of the physical and chemical characteristics of the ocean. Also, the structure and the functioning of marine ecosystems may be altered as a result of ocean acidification. <p>Phytoplankton productivity is one of the primary controls in regulating our climate, for instance via impact on atmospheric CO2 levels. Coccolithophores, of which Emiliania huxleyi is the most abundant species, are considered to be the most important pelagic calcifying organisms on Earth. Coccolithophores are characterized by calcium carbonate platelets (coccoliths) covering the exterior of the cells. They form massive blooms in temperate and sub-polar oceans and in particular along continental margin and in shelf seas. The intrinsic coupling of organic matter production and calcification in coccolithophores underlines their biogeochemical importance in the marine carbon cycle. Both processes are susceptible to change with ocean acidification and warming. Coccolithophores are further known to produce transparent exopolymer particles (TEP) that promote particle aggregation and related processes such as marine snow formation and sinking. Thus, the impact of ocean warming and acidification on coccolithophores needs to be studied and this can be carried out through a transdisciplinary approach.<p>The first part of this thesis consisted of laboratory experiments on E. huxleyi under controlled conditions. The aim was to estimate the effect of increasing water temperature and acidity on E. huxleyi and especially on the calcification. Cultures were conducted at different partial pressures of CO2 (pCO2); the values considered were 180, 380 and 750 ppm corresponding to past, present and future (year 2100) atmospheric pCO2. These experiments were conducted at 13°C and 18°C. The cellular calcite concentration decreases with increasing pCO2. In addition, it decreases by 34 % at 380 ppm and by 7 % at 750 ppm with an increase in temperature of 5°C. Changes in calcite production at future pCO2 values are reflected in deteriorated coccolith morphology, while temperature does not affect coccolith morphology. Our findings suggest that the sole future increase of pCO2 may have a larger negative impact on calcification than its interacting effect with temperature or the increase in temperature alone. The evolution of culture experiments allows a better comprehension of the development of a bloom in natural environments. Indeed, in order to predict the future evolution of calcifying organisms, it is required to better understand the present-day biogeochemistry and ecology of pelagic calcifying communities under field conditions.<p>The second part of this dissertation was dedicated to results obtained during field investigations in the northern Bay of Biscay, where frequent and recurrent coccolithophorid blooms were observed. Cruises, assisted by remote sensing, were carried out along the continental margin in 2006 (29 May – 10 June), 2007 (7 May – 24 May) and 2008 (5 May – 23 May). Relevant biogeochemical parameters were measured in the water column (temperature, salinity, dissolved oxygen, Chlorophyll-a and nutrient concentrations) in order to determine the status of the bloom at the time of the different campaigns. Calcification has been shown to be extremely important in the study area. In addition, TEP production was significant at some stations, suggesting that the northern Bay of Biscay could constitute an area of important carbon export. Mortality factors for coccolithophores were studied and the first results of lysis rates measured in this region were presented. <p>Results obtained during culture experiments and comparison with data reported in the literature help to better understand and to predict the future of coccolithophores in a context of climate change. Data obtained during either culture experiments or field investigations allowed a better understanding of the TEP dynamics. Finally, the high lysis rates obtained demonstrate the importance of this process in bloom decline. Nevertheless, it is clear that we only begin to understand the effects of global change on marine biogeochemistry, carbon cycling and potential feedbacks on increasing atmospheric CO2. Thus, further research with a combination of laboratory experiments, field measurements and modelling are encouraged.<p><p> / Doctorat en Sciences / info:eu-repo/semantics/nonPublished

Coccolithophores are unicellular microscopic algae (Haptophyta) surrounded by calcium carbonate plates that are produced during their life cycle. These species, whose contemporary contributor is Emiliania huxleyi, are mainly found in the sub-polar and temperate oceans, where they produce huge blooms visible from space. Coccolithophores are sensitive to ocean acidification that results from the ongoing accumulation of anthropogenic carbon dioxide (CO2) in the atmosphere. The response of these organisms to global change appears to be related to the reduction of their ability to produce calcium carbonate at the cellular level. At the community levels, one anticipates changes in the carbon fluxes associated to their blooms as calcification is reduced. However, the consequences of such environmental changes on this species are speculative and require improvements in the description of the mechanisms controlling the organic and inorganic carbon production and export.<p><p>The first aspect of this work was to study the response of these organisms to artificially modified CO2 concentrations representative of the conditions occurring in the past (glacial) and those expected by the end of the century (2100). Two different levels were examined: the continuous monospecific cultures (chemostats) allowed us to work at the cellular level while the mesocosms gave light to the mechanisms taking place in an isolated fraction of the natural community. The second aspect of this work consisted of field studies carried out during four cruises (2002, 2003, 2004 and 2006) in the northern Bay of Biscay, where the occurrence of E. huxleyi blooms were observed in late spring and early summer. We describe the vertical profiles of biogeochemical variables (nutrients, chlorophyll-a, dissolved inorganic chemistry, particulate carbon, transparent exopolymer particles (TEP)) and study processes such as primary production, calcification and bacterial production. The properties of these blooms are compared with those reported in the literature and enriched with original measurements such as the abundance and concentration of TEP that could play an important role in carbon export to the deep ocean, modifying the properties of the settling ballasted aggregates.<p> / Doctorat en Sciences / info:eu-repo/semantics/nonPublished