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Large scale spatio-temporal variation of carbon fluxes along the land-ocean continuum in three hotspot regionsHastie, Adam 03 June 2019 (has links) (PDF)
Previous research has shown a close relationship between the terrestrial and aquatic carbon (C) cycles, namely that part of the C fixed via terrestrial net primary production (NPP) is exported to inland waters. In turn, it has been demonstrated that once in the freshwater system C can not only be transported laterally as dissolved organic carbon (DOC), particulate organic carbon (POC) and dissolved inorganic carbon (DIC) but is also mineralized and evaded back to the atmosphere as CO2, or buried in sediments. A number of hotspot areas of aquatic CO2 evasion have been identified but there are considerable gaps in our knowledge, particularly associated with understanding and accounting for the temporal and spatial variation of aquatic C fluxes at regional to global scales, which we know from local scale studies, to be substantial. In this thesis, three important regional hotspots of LOAC activity were identified, where significant gaps in our understanding remain.For the boreal region, an empirical model is developed to produce the first high resolution maps of boreal lake pCO2 and CO2 evasion, providing a new estimate for total evasion from boreal lakes of 189 (74–347) Tg C yr-1, which is more than double the previous best estimate. The model is also used along with future projections of terrestrial NPP and precipitation, to predict future lake CO2 evasion under future climate change and land-use scenarios, and it is found that even under the most conservative scenario CO2 evasion from boreal lakes may increase 38% by 2100. For the Amazon Basin, the ORCHILEAK land surface model driven by a newly developed wetland forcing file, is used to show that the export of C to and CO2 evasion from inland waters is highly interannually variable; greatest during wet years and lowest during droughts. However, at the same time overall net ecosystem productivity (NEP) and C sequestration is highest during wet years, partly due to reduced decomposition rates in water-logged floodplain soils. Furthermore, it is shown that aquatic C fluxes display greater variation than terrestrial C fluxes, and that this variation significantly dampens the interannual variability in NEP of the Amazon basin by moderating terrestrial variation. Finally, ORCHILEAK is applied to the Congo Basin to investigate the evolution of the integrated aquatic and terrestrial C fluxes from 1861 to the present day, and in turn to 2099 under a future climate and land-use scenario. It is shown that terrestrial and aquatic fluxes increase substantially over time, both over the historical period and into the future, and that these increases are largely driven by atmospheric CO2. The proportion of terrestrial NPP lost to the LOAC also rises from 3% in 1861 to 5% in 2099 and this trend is driven not only by atmospheric CO2 but also by climate change. This is in contrast to the boreal region where the proportion of NPP exported to inland waters is predicted to remain relatively constant, and to the Amazon, where a decrease has been predicted, due to differences in projected climate change. / L’état de l’art dans le domaine a montré qu’il y avait un lien étroit entre les cycles du carbone terrestre et aquatique :en effet, une partie du carbone fixé par photosynthèse (productivité primaire brute) est transférée vers les milieux aquatiques continentaux pour être ensuite transporté latéralement sous forme de carbone organique dissous (COD), de carbone organique particulaire (COP), de carbone inorganique dissous (CID). Durant ce transfert latéral, le carbone peut être minéralisé puis réémis vers l’atmosphère sous forme de CO2 ou enfoui dans les sédiments. Cependant, nous sommes encore loin de bien comprendre et surtout de quantifier les variations temporelles et spatiales des flux de carbones à l’échelle régionale et globale, même si les études faites à l’échelle locale nous montrent qu’elles sont importantes. Au cours de cette thèse, nous nous sommes focalisés sur 3 grandes régions pour lesquelles la connaissance des flux de carbone le long du continuum aquatique reliant les écosystèmes terrestres aux océans étaient encore très parcellaire.Pour la région boréale, un modèle empirique a été développé afin de produire les premières cartes à haute résolution de pCO2 et d’émission de CO2 pour les lacs boréaux. Les résultats du modèle nous ont permis de contraindre les émissions totales de CO2 pour les lacs boréaux à 189 (74-347) Tg C an-1, soit plus du double des estimations précédentes. Ce modèle a ensuite été couplé aux projections de production primaire brute terrestre et de précipitations afin de prédire les émissions de CO2 pour ces lacs pour différents scénarios de changement climatique et d’occupation des sols. Les résultats montrent que même en prenant le scénario le plus conservatif, les émissions de CO2 des lacs boréaux augmenteraient de 38% d’ici 2100.Pour le bassin de l’Amazone, le modèle d’écosystème terrestre ORCHILEAK, paramétré par de nouvelles donnés de forçage des zones humides, a été utilisé pour démontrer que l’export de carbone terrestre vers les réseaux fluviaux ainsi que les émissions de CO2 ont une très grande variabilité interannuelle :émissions élevées lors des années à forte précipitation et basses lors des années sèches. Cependant, la productivité nette de l’écosystème (PNE) Amazone et la fixation nette de carbone à l’échelle du bassin sont plus élevées lors des années humides, en partie dû au taux de décomposition de carbone organique réduit lorsque les sols sont saturés en eau. De plus, les résultats montrent que les flux de carbone des systèmes aquatiques ont une plus grande variabilité que les flux terrestres, ce qui atténue considérablement la variabilité interannuelle de la PNE du bassin de l'Amazone.Pour finir, nous avons appliqué ORCHILEAK au bassin du Congo afin d’étudier l’évolution intégrée des flux de carbone terrestres et aquatiques de 1861 à nos jours, ainsi que de projeter leur devenir au cours du 21eme siècle selon les scénarios de changement climatiques et de changement d’occupation des sols. Nous avons montré que les flux terrestres et aquatiques augmentent de façon significative durant la période historique et dans le futur, cette augmentation étant largement induite par l’augmentation du CO2 atmosphérique et, dans une moindre mesure, par le changement climatique. En particulier, la proportion de la productivité primaire brute terrestre exportée vers le continuum aquatique passe de 3% en 1861 à 5% en 2099. Ce résultat contraste avec ceux obtenu pour la région boréale où cette proportion reste relativement constante et pour l’Amazone où c’est une baisse qui est en fait prédite. Ces différences s’expliquent par des trajectoires de changement climatique distinctes pour ces 3 régions. / Doctorat en Sciences / info:eu-repo/semantics/nonPublished
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Linking dissolved organic matter quality and quantity to CO2 and CH4 concentrations in ombrotrophic bog poolsHassan, Mahmud 06 1900 (has links)
Les petits plans d'eau, en particulier ceux riches en matière organique, sont encore négligés en tant que sources naturelles majeures d'émission de carbone (C) dans l'atmosphère et contributeurs importants au bilan mondial de C. Les mares de tourbières riches en matières organiques sont une source nette de C atmosphérique dans les écosystèmes de tourbières, qui sont généralement un puits net de C. Ces mares émettent des gaz à effet de serre (GES) à des taux plus élevés, en particulier de méthane (CH4), par rapport à d'autres petits plans d'eau lentiques (petits lacs et étangs), ce qui peut être attribué à la connectivité hydraulique des bassins donc aux apports en C de la tourbe environnante et aux caractéristiques morphologiques des mares. Cependant, il existe très peu d'informations sur les schémas et les mécanismes de la dynamique du C dans les bassins de tourbières par rapport à leur couvert végétal ainsi qu'à d'autres petits environnements aquatiques. En particulier, la matière organique dissoute (MOD), un important intermédiaire et substrat de C, peut influencer la dynamique des émissions de GES, mais son rôle demeure méconnu à l'échelle intra- et inter-régionale. Dans cette étude, nous avons caractérisé et identifié les patrons intra- et interrégionaux et les mécanismes potentiels contrôlant la quantité et la qualité de la MOD et des concentrations de GES, ainsi que leurs liens en analysant une gamme de variables optiques et chimiques et en compilant les données géographiques (c'est-à-dire le climat, le couvert végétal et morphométrie des bassins) à partir de bassins de tourbières ombrotrophes dans cinq régions de l'est du Canada (Grande plée Bleue, sud du Québec et région de la Minganie, est du Québec) et du sud de la Patagonie chilienne (Punta Arenas, parc Karukinka et île Navarino). Nous avons également effectué un échantillonnage interannuel dans la Grande plée Bleue pour identifier les tendances temporelles des concentrations et de la composition des GES et de MOD. Nous avons trouvé une variabilité interrégionale élevée dans les patrons de MOD et de GES par rapport à la variabilité intrarégionale qui était cohérente avec l'hétérogénéité des propriétés géographiques, en particulier, le climat. Les patrons interrégionaux des concentrations de GES étaient de plus déterminés par la couverture végétale environnante, la morphométrie du bassin et la composition de la MOD de type protéique. D'autre part, bien que nous n'ayons pas observé de patrons temporels significatifs dans les concentrations de GES, de MOD et de la composition de type humique terrestre au cours de l’été dans la Grande plée Bleue, les patrons temporels de GES ont été influencés par la concentration de MOD, la composition de type humique terrestre, et la chimie interne de l'eau. Dans l'ensemble, nos résultats suggèrent que les patrons interrégionaux de la MOD
et des GES, et les liens entre eux, sont principalement contrôlés par le climat (température et précipitations), la couverture végétale et la morphométrie des bassins, tandis que les patrons temporels de la MOD et des GES sont principalement régis par des facteurs à l'échelle locale tels que la morphométrie des bassins et la connectivité hydrologique. / Small waterbodies, especially organic-rich, are still overlooked as a major natural source of carbon (C) emission to the atmosphere and an important contributor in the global C budget. Organic-rich peatland pools are generally net atmospheric C sources embedded in peatland ecosystems, which are generally net C sinks. They emit high areal rates of greenhouse gases (GHG), particularly methane (CH4), compared to other small lentic waterbodies (small lakes and ponds) which may be attributed to peat-pool hydraulic connectivity leading to C loading from the surrounding peat and morphological characteristics. But there is very little information on the patterns and drivers of C dynamics within peatland pools compared to their vegetated areas as well as other small aquatic environments. In particular, the role that dissolved organic matter (DOM), an important intermediate and C substrate, may play in GHG dynamics is poorly known at the intra- and inter-regional scales. In this study, we characterized and identified the intra- and inter regional patterns and drivers of DOM quantity and quality and GHG concentrations as well as their links. We did so by analyzing a range of optical and chemical variables and compiling geographic data (i.e., climate, vegetation cover and pool morphometry) from ombrotrophic peatland pools across five regions in eastern Canada (Grande plée Bleue, southern Québec and Minganie region, eastern Québec) and southern Chilean Patagonia (Punta Arenas, Karukinka Park and Navarino Island). We also conducted inter-annual sampling in Grande plée Bleue to identify the temporal patterns in GHG and DOM concentrations and composition. We found high inter-regional variability in DOM and GHG patterns compared to intra-regional variability which was coherent with the heterogeneity of geographical properties. Inter-regional patterns in GHG concentrations were driven by surrounding vegetation cover, pool morphometry and protein-like DOM composition. On the other hand, although we did not observe significant temporal patterns in GHG and DOM concentrations and terrestrial humic-like composition throughout the growing seasons in Grande plée Bleue, temporal patterns of GHG were influenced by the DOM concentration, terrestrial humic-like composition, and internal water chemistry. Overall, our results suggest that inter-regional patterns in DOM and GHG, and the links among them are predominantly controlled by the broad-scale patterns in climate (temperature and precipitation), vegetation cover, and pool morphometry, while temporal patterns in DOM and GHG are predominantly governed by local-scale drivers such as pool morphometry.
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Understanding Land-Atmosphere Interactions Across Multiple ScalesHuang, Yu January 2024 (has links)
The terrestrial water, energy and carbon cycles are tightly coupled through land-atmosphere (L-A) interactions, not only regulating local plant physiological activities and also modulating regional and global climate. With ongoing anthropogenic greenhouse gas emissions, many of these interactions can be modified and complicated. To better anticipate and adapt to future climate, it is of great importance and necessity to deepen and refine our understanding of the complex L-A interactions. In this dissertation, three topics are investigated across the ecosystem, regional and global scales respectively, throughout which, the critical role of dryness or drying in the context of global warming is highlighted.
𝐂𝐡𝐚𝐩𝐭𝐞𝐫 𝟏: Evapotranspiration (ET) is a key component that connects the continental water, carbon and energy cycles and a proxy that measures the coupling strength between the biosphere and atmosphere. A wide range of biophysical factors, which usually exhibit nonlinearity and strong covariation, collectively modulate ET and complicate the overall understanding of ET dynamics. In the first study, the causal discovery frameworks PCMCI+ and Latent PCMCI are utilized with integrated priori physical knowledge to identify the dominant drivers and constraints of ET in the growing seasons across sites, with a particular focus on the role of site dryness degree. The Dryness Index (DI), defined as the ratio of annual mean net radiation to precipitation, has been introduced to assess the water availability relative to energy supply at different locations. By analyzing the daily observations from 115 flux tower sites and satellite remote sensing, it has been discovered that the feedbacks around ET are mediated by the degree of dryness: at sites with adequate water supply (using PCMCI+, the DI value averaged from such sites is 1.33), the atmospheric conditions, including incoming solar radiation and atmospheric demand for water (indicated by vapor pressure deficit, VPD), prevail in driving ET; in contrast, in semi-arid and arid areas where the water stress is high (using PCMCI+, the DI value averaged from such sites is 3.32), soil water content is the primary factor to constrain ET due to the plant regulation of stomatal conductance as part of the water conservation strategy. Additionally, as DI increases across sites, the sign of the contemporaneous causal relationship between VPD and ET can reverse from positive—indicating that atmospheric demand for water drives ET—to negative—reflecting that plant stomatal closure limits ET in response to the dryer atmosphere.
𝐂𝐡𝐚𝐩𝐭𝐞𝐫 𝟐: As summer heatwaves and droughts are becoming more frequent and intense, such as in Western Europe, there is a growing interest in unraveling the physical mechanisms behind their occurrences and their changes. Soil desiccation is critical for the intensification and propagation of heatwaves, but its relative importance compared to other well-known large-scale atmospheric mechanisms, such as persistent atmospheric blocking systems and horizontal warm advection, remains elusive, especially in the context of a changing climate. In the second study, we utilize machine learning along with intervention experiments to estimate the respective contributions of soil water content 𝐶_𝑠𝑤𝑐 and atmospheric circulation 𝐶_𝑎𝑡𝑚 to daily maximum temperature in Western Europe, with a particular focus on the 2022 summer events. Our results reveal that during the two unparalleled heatwave events that occurred in June and July of 2022, the impact 𝐶_𝑠𝑤𝑐 on the heatwave intensity was on average approximately 40% of 𝐶_𝑎𝑡𝑚, and was comparable to 𝐶_𝑎𝑡𝑚 in continental dry-to-wet transition regions. Reviewing heatwaves in recent three decades, the percentage of heatwave areas that are significantly influenced by soil moisture-air temperature coupling has increased by 11.4% per decade. Additionally, for regions that have experienced heatwaves in at least 5 out of the past 33 years, about 21.7% areas, mostly in the transition zones, witness a significant increase in 𝐶_𝑠𝑤𝑐; while only 2.5% exhibit a substantial increase in 𝐶_𝑎𝑡𝑚. Furthermore, we find within the transitional climates, the intensification of heat extremes is mainly resulted from soil moisture depletion rather than atmospheric anomalies; while in (dry) Spain and the (wet) northern areas of central Europe, it is the variations in atmospheric circulation and soil desiccation that jointly fuel the persistent heatwaves. Our study emphasizes the observation-based large and increasing importance of soil moisture coupling in intensifying summer heatwaves and provides insights into future climates in extra-tropical regions like Western Europe, where a warmer and drier future is projected.
𝐂𝐡𝐚𝐩𝐭𝐞𝐫 𝟑: Earth system models (ESMs) and climate simulations are extensively employed to study the dynamics of climate and project long-term changes in the climate system. Despite their widespread use, large uncertainties persist among these models regarding the estimation of the continental gross primary productivity (GPP) and land carbon sink, which compromise the reliability of projections concerning future atmospheric carbon dioxide (𝐶𝑂₂) concentrations and the assessment of how terrestrial ecosystems respond to and might mitigate some of global warming. In ESMs, convection and clouds are one major source of such uncertainties—they are not only the most uncertain factors in the modeling of ``physical'' climate and also significantly affect the land carbon cycle through complex interactions involving radiation, moisture, and thermal pathways. In the third study, to isolate the role of clouds on the terrestrial carbon cycle, two models—the Community Earth System Model (CESM) and its super-parameterized counterpart (SPCESM, abbreviated as SP), which only differ in their representation of convection and clouds, are analyzed under present-day climatology to assess the impact of cloud representations on GPP. Compared with CESM, SP shows a 12.8% decrease in total cloud fraction within the 60°𝑆 ∼ 60°𝑁 range, which results in a notable GPP decline of 5.6 𝑃𝑔𝐶 𝑦𝑟⁻¹. This divergence, equivalent to 4.4% of terrestrial GPP in CESM, is comparable to the inter-annual variability in GPP and the uncertainty of GPP observed across climate models with diverse representations, extending beyond just cloud-related processes.
Further analysis decomposes the GPP divergence between CESM and SP into two additive components and demonstrates that three-quarters of the difference is attributed to the negative impact from reduced cloud cover on light use efficiency (LUE) from CESM to SP, while the remaining one quarter is due to the positive impact from enhanced photosynthetically active radiation (PAR). An explainable machine learning model equipped with SHAP values further identifies two primary mechanisms underlying the lower LUE estimation in SP. Firstly, diminished clouds lead to higher air temperatures and reduced precipitation, creating a drier environment that prompts plants to regulate stomatal conductance to minimize water loss through transpiration, thereby suppressing the exchange rate of 𝐶𝑂₂ between biosphere and atmosphere. Secondly, the reduction in diffused radiation restricts the photosynthesis of shaded leaves. Combined, these two mechanisms reduce plant LUE, outweigh the beneficial impacts of increased PAR on photosynthesis, and ultimately lead to the declined terrestrial biosphere productivity in SP. Overall, we identify the representation of clouds as a key process for the terrestrial carbon cycle.
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Earth system dynamics in the Anthropocen / modelling key processes of climate-human interactions in the terrestrial biosphereBeringer, Tim 12 January 2012 (has links)
In nie dagewesener Größenordnung greift der Mensch durch die Verbrennung fossiler Energieträger und der weiträumigen Umgestaltung der Landoberfläche in die globale Umwelt ein. Klimawandel und Übernutzung natürlicher Ressourcen könnten schon in diesem Jahrhundert die Anpassungsfähigkeiten vieler ökologischer und sozialer Systeme übersteigen und somit zu Konflikten und politischer Destabilisierung führen. Vor diesem Hintergrund soll diese Studie zu einem besseren Verständnis der wichtigsten globalen Triebkräfte beitragen, die die Entwicklung der terrestrischen Biosphäre in diesem Jahrhundert prägen werden: Klimawandel und menschliche Landnutzung. Auf der Basis eines Dynamischen Globalen Vegetationsmodells werden im ersten Teil der vorliegenden Arbeit zwei große klimatische Störungen des globalen Kohlenstoffkreislaufs untersucht, die innerhalb der letzten drei Jahrzehnte beobachtet wurden. Im Fordergrund steht die Frage, wie sich die Veränderungen von Temperatur-, Niederschlags- und Strahlungsbedingungen auf pflanzliche Produktivität und Zersetzungsprozesse im Boden auswirkten. Es zeigt sich, dass vermehrte Kohlenstoffspeicherung in der Landbiosphäre den überwiegenden Teil der atmosphärischen CO2 Anomalien erklärt. Der zweite Teil dieser Arbeit beschäftigt sich mit der weltweit steigenden Nachfrage nach Bioenergie, die aufgrund des flächenintensiven Anbaus von Biomasse zur wichtigsten Triebkraft für zukünftige Landnutzungsänderungen werden könnte. Aus der Kombination von Vegetationsmodellierung und räumlichen Datenanalysen werden globale Bioenergiepotentiale unter Berücksichtigung verschiedener Nachhaltigkeitsanforderungen bestimmt und mögliche ökologische Auswirkungen des großräumigen Anbaus von Energiepflanzen abgeschätzt. Im Jahr 2050 könnten demnach 15-25% des weltweiten Energiebedarfs durch Bioenergie abgedeckt werden. Dafür müssten allerdings natürliche Ökosysteme in großem Umfang in Agrarland umgewandelt werden. / Human activities, primarily the combustion of fossil fuels and the global modification of the land surface, are transforming the Earth System at unprecedented scale. Climate change and the overexploitation of natural resources may soon overwhelm the adaptive capacities of many ecosystems and societies, which could lead to substantial losses in human well-being and political destabilization. In this context, it is the goal of this thesis to contribute to a better understanding of the most important global drivers that will determine the future of the land biosphere during this century: climate change and human land use. Based on a Dynamic Global Vegetation Model (DGVM), the first part of this thesis examines two large climatic disturbances of the terrestrial carbon cycle that were observed during the last three decades. These analyses focus on the effects of changes in temperature, precipitation and radiation on plant productivity and soil decomposition. Results indicate that increased carbon storage in the land biosphere explains the most part of the atmospheric CO2 anomaly. The second part of this thesis addresses the worldwide increasing demand for bioenergy that may become the most important driver of future land use change due to the large area requirements of biomass cultivation. A combination of vegetation modeling and spatial data analyses is used to assess global bioenergy potentials that consider various sustainability requirements for food security, biodiversity protection and the reduction of greenhouse gas emissions and to evaluate the environmental impacts of large-scale energy crop cultivation. The results indicate that bioenergy may provide between 15 and 25% of the global energy demand in 2050. Exploiting these potentials, however, requires the conversion of large amounts of natural vegetation into agricultural land affecting a large number of ecosystems already fragmented and degraded by land use change.
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Elevation Effects on Key Processes of Carbon Cycling in South Ecuadorian Mountain Forests / Der Einfluss der Meereshöhe auf Schlüsselprozesse des Kohlenstoffkreislaufs in Südecuadorianischen BergregenwäldernMoser, Gerald 24 January 2008 (has links)
No description available.
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A Reconnaissance Study of Water and Carbon Fluxes in Tropical Watersheds of Peninsular Malaysia: Stable Isotope ConstraintsIshak, Muhammad Izzuddin Syakir 04 February 2014 (has links)
Evapotranspiration is a nexus for planetary energy and carbon cycles, as yet poorly constrained. Here I use stable isotopes of oxygen and hydrogen to partition flux of water due to plant transpiration from the direct evaporative flux from soils, water bodies and plant. The study areas, Langat and Kelantan watersheds represent examples of domains dominated by the respective Southwest and Northeast monsoons on the two sides of the main orographic barrier (Titiwangsa mountain range). Mean annual rainfall for the Langat watershed, obtained from 30 years of hydrological data, is 2145 ± 237 mm. Tentatively, 48% of this precipitation returns to the atmosphere via transpiration (T), with 33% partitioned into discharge (Q), 8% into interception (In), and 11% into evaporation (Ed). In the Kelantan watershed, the mean annual rainfall, also based on the 30 year hydrological data, is 2383 ± 120 mm. Similar to Langat, the T accounts for 43% of precipitation (P), 45% is discharged into South China Sea (Q), 12% partitioned into interception (In) and tentatively 0% for evaporation (Ed). Ed for the Langat watershed represents only a small proportion in terms of volumetric significance, up to almost ~11% with strong effect on the isotopic fingerprints of waters associated with the summer Southwest Monsoon (SWM). Note, however, that insignificant Ed for the Kelantan watershed may be an artefact of rain and river water sampling at only coastal downstream portion of the watershed. High humidity (80%) also was recorded for the Malaysian Peninsula watershed.
T appropriates about half of all solar energy absorbed by the continents, here ~1000*103 g H2O m-2 yr-1 similar to other tropical regions at 900-1200*103 g H2O m-2 yr-1. The associated carbon fluxes are ~ 1300 g C m-2yr-1, independent of P. Vegetation responses to solar irradiance, via T and photosynthesis reflects the importance of stomatal regulation of the water and carbon fluxes. In order to maintain high transpiration in the tropical region, “constant” water supply is required for continuous pumping of water that delivers nutrients to the plant, suggesting that water and carbon cycle are co-driven by the energy of the sun. The existence of the water conveyor belt may be precondition for nutrient delivery, hence operation of the carbon cycle. Potentially, this may change our perspective on the role that biology plays in the water cycle. In such perspective, the global water cycle is the medium that redistributes the incoming solar energy across the planet, and the anatomical structures of plants then help to optimize the loop of energy transfer via evaporation and precipitation in the hydrologic cycle.
The main features of aquatic geochemistry of the Langat and Kelantan rivers inferred from the Principal Component Analysis are controlled by three components that explain 80% and 82% of total variances. These components are reflecting of the geogenic factor with superimposed pollution, the latter particularly pronounced in urbanized sections of the Langat river and dominant in downstream of the Kelantan river. There is no correlation between seasonal variations in major ion chemistry and environmental variables such as precipitation, discharge, temperature or solar activity.
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Interactions between the microbial network and the organic matter in the Southern Ocean: impacts on the biological carbon pump / Interactions entre le réseau microbien et la matière organique dans l'Océan Antarctique: impacts sur la pompe biologique à carboneDumont, Isabelle 03 July 2009 (has links)
<p align="justify">The Southern Ocean (ca. 20% of the world ocean surface) is a key place for the regulation of Earth climate thanks to its capacity to absorb atmospheric carbon dioxide (CO2) by physico-chemical and biological mechanisms. The biological carbon pump is a major pathway of absorption of CO2 through which the CO2 incorporated into autotrophic microorganisms in surface waters is transferred to deep waters. This process is influenced by the extent of the primary production and by the intensity of the remineralization of organic matter along the water column. So, the annual cycle of sea ice, through its in situ production and remineralization processes but also, through the release of microorganisms, organic and inorganic nutrients (in particular iron)into the ocean has an impact on the carbon cycle of the Southern Ocean, notably by promoting the initiation of phytoplanktonic blooms at time of ice melting.</p><p><p align="justify">The present work focussed on the distribution of organic matter (OM) and its interactions with the microbial network (algae, bacteria and protozoa) in sea ice and ocean, with a special attention to the factors which regulate the biological carbon pump of the Southern Ocean. This thesis gathers data collected from a) late winter to summer in the Western Pacific sector, Western Weddell Sea and Bellingshausen Sea during three sea ice cruises ARISE, ISPOL-drifting station and SIMBA-drifting station and b) summer in the Sub-Antarctic and Polar Front Zone during the oceanographic cruise SAZ-Sense.</p><p><p align="justify">The sea ice covers were typical of first-year pack ice with thickness ranging between 0.3 and 1.2 m, and composed of granular and columnar ice. Sea ice temperature ranging between -8.9°C and -0.4°C, brines volume ranging between 2.9 to 28.2% and brines salinity from 10 to >100 were observed. These extreme physicochemical factors experienced by the microorganisms trapped into the semi-solid sea ice matrix therefore constitute an extreme change as compared to the open ocean. Sea ice algae were mainly composed of diatoms but autotrophic flagellates (such as dinoflagellates or Phaeocystis sp.) were also typically found in surface ice layers. Maximal algal biomass was usually observed in the bottom ice layers except during SIMBA where the maxima was localised in the top ice layers likely because of the snow and ice thickness which limit the light available in the ice cover. During early spring, the algal growth was controlled by the space availability (i.e. brine volume) while in spring/summer (ISPOL, SIMBA) the major nutrients availability inside sea ice may have controlled algal growth. At all seasons, high concentrations of dissolved and particulate organic matter were measured in sea ice as compared to the water column. Dissolved monomers (saccharides and amino acids) were accumulated in sea ice, in particular in winter. During spring and summer, polysaccharides constitute the main fraction of the dissolved saccharides pool. High concentrations of transparent exopolymeric particles (TEP), mainly constituted with saccharides, were present and their gel properties greatly influence the internal habitat of sea ice, by retaining the nutrients and by preventing the protozoa grazing pressure, inducing therefore an algal accumulation. The composition as well as the vertical distribution of OM in sea ice was linked to sea ice algae.</p><p><p align="justify">Besides, the distribution of microorganisms and organic compounds in the sea ice was also greatly influenced by the thermodynamics of the sea ice cover, as evidenced during a melting period for ISPOL and during a floodfreeze cycle for SIMBA. The bacteria distribution in the sea ice was not correlated with those of algae and organic matter. Indeed, the utilization of the accumulated organic matter by bacteria seemed to be limited by an external factor such as temperature, salinity or toxins rather than by the nature of the organic substrates, which are partly composed of labile monomeric saccharides. Thus the disconnection of the microbial loop leading to the OM accumulation was highlighted in sea ice.</p><p><p align="justify">In addition the biofilm formed by TEP was also involved in the retention of cells and other compounds(DOM, POM, and inorganic nutrients such as phosphate and iron) to the brine channels walls and thus in the timing of release of ice constituents when ice melts. The sequence of release in marginal ice zone, as studied in a microcosm experiments realized in controlled and trace-metal clean conditions, was likely favourable to the development of blooms in the marginal ice zone. Moreover microorganisms derived from sea ice (mainly <10 µm) seems able to thrive and grow in the water column as also the supply of organic nutrients and Fe seems to benefit to the pelagic microbial community.</p><p><p align="justify">Finally, the influence of the remineralization of organic matter by heterotrophic bacterioplankton on carbon export and biological carbon pump efficiency was investigated in the epipelagic (0-100 m) and mesopelagic(100-700 m) zones during the summer in the sub-Antarctic and Polar Front zones (SAZ and PFZ) of the Australian sector (Southern Ocean). Opposite to sea ice, bacterial biomass and activities followed Chl a and organic matter distributions. Bacterial abundance, biomass and activities drastically decreased below depths of 100-200 m. Nevertheless, depth-integrated rates through the thickness of the different water masses showed that the mesopelagic contribution of bacteria represents a non-negligible fraction, in particular in a diatom-dominated system./</p><p><br><p><p align="justify">L’océan Antarctique (± 20% de la surface totale des océans) est un endroit essentiel pour la régulation du climat de notre planète grâce à sa capacité d’absorber le dioxyde de carbone (CO2) atmosphérique par des mécanismes physico-chimique et biologique. La pompe biologique à carbone est un processus majeur de fixation de CO2 par les organismes autotrophes à la surface de l’océan et de transfert de carbone organique vers le fond de l’océan. Ce processus est influencé par l’importance de la production primaire ainsi que par l’intensité de la reminéralisation de la matière organique dans la colonne d’eau. Ainsi, le cycle annuel de la glace via sa production/reminéralisation in situ mais aussi via l’ensemencement de l’océan avec des microorganismes et des nutriments organiques et inorganiques (en particulier le fer) a un impact sur le cycle du carbone dans l’Océan Antarctique, notamment en favorisant l’initiation d’efflorescences phytoplanctoniques dans la zone marginale de glace.</p><p><p align="justify">Plus précisément, nous avons étudié les interactions entre le réseau microbien (algues, bactéries et protozoaires) et la matière organique dans le but d’évaluer leurs impacts potentiels sur la pompe biologique de carbone dans l’Océan Austral. Deux écosystèmes différents ont été étudiés :la glace de mer et le milieu océanique grâce à des échantillons prélevés lors des campagnes de glace ARISE, ISPOL et SIMBA et lors de la campagne océanographique SAZ-Sense, couvrant une période allant de la fin de l’hiver à l’été.</p><p><p align="justify">La glace de mer est un environnement très particulier dans lequel les microorganismes planctoniques se trouvent piégés lors de la formation de la banquise et dans lesquels ils subissent des conditions extrêmes de température et de salinité, notamment. Les banquises en océan ouvert étudiées (0,3 à 1,2 m d’épaisseur, températures de -8.9°C à -0.4°C, volumes relatifs de saumure de 2.9 à 28.2% et salinités de saumures entre 10 et jusque >100) étaient composées de glace columnaire et granulaire. Les algues de glace étaient principalement des diatomées mais des flagellés autotrophes (tels que des dinoflagellés ou Phaeocystis sp.) ont été typiquement observés dans les couches de glace de surface. Les biomasses algales maximales se trouvaient généralement dans la couche de glace de fond sauf à SIMBA où les maxima se trouvaient en surface, probablement en raison de l’épaisseur des couches de neige et de glace, limitant la lumière disponible dans la colonne de glace. Au début du printemps, la croissance algale était contrôlée par l’espace disponible (càd le volume des saumures) tandis qu’au printemps/été, la disponibilité en nutriments majeurs a pu la contrôler. A toutes les saisons, des concentrations élevées en matière organique (MO) dissoute et particulaire on été mesurées dans la glace de mer par rapport à l’océan. Des monomères dissous (sucres et acides aminés) étaient accumulés dans la glace, surtout en hiver. Au printemps et été, les polysaccharides dissous dominaient le réservoir de sucres. La MO était présente sous forme de TEP qui par leurs propriétés de gel modifie l’habitat interne de la glace. Ce biofilm retient les nutriments et gêne le mouvement des microorganismes. La composition et la distribution de la MO dans la glace étaient en partie reliées aux algues de glace. De plus, la thermodynamique de la couverture de glace peut contrôler la distribution des microorganismes et de la MO, comme observé lors de la fonte de la glace à ISPOL et lors du refroidissement de la banquise à SIMBA. La distribution des bactéries n’est pas corrélée avec celle des algues et de la MO dans la glace. En effet, la consommation de la MO par les bactéries semble être limitée non pas par la nature chimique des substrats mais par un facteur extérieur affectant le métabolisme bactérien tel que la température, la salinité ou une toxine. Le dysfonctionnement de la boucle microbienne menant à l’accumulation de la MO dans la glace a donc été mis en évidence dans nos échantillons.</p><p><p align="justify">De plus, le biofilm formé par les TEP est aussi impliquée dans l’attachement des cellules et autres composés aux parois des canaux de saumure et donc dans la séquence de largage lors de la fonte. Cette séquence semble propice au développement d’efflorescences phytoplanctoniques dans la zone marginale de glace. Les microorganismes originaires de la glace (surtout ceux de taille < 10 μm) semblent capables de croître dans la colonne d’eau et l’apport en nutriments organiques et inorganiques apparaît favorable à la croissance des microorganismes pélagiques.</p><p><p align="justify">Enfin, l’influence des activités hétérotrophes sur l’export de carbone et l’efficacité de la pompe biologique à carbone a été évaluée dans la couche de surface (0-100 m) et mésopélagique (100-700 m) de l’océan. Au contraire de la glace, les biomasses et activités bactériennes suivaient les distributions de la chlorophyll a et de la MO. Elles diminuent fortement en dessous de 100-200 m, néanmoins les valeurs intégrées sur la hauteur de la colonne d’eau indiquent que la reminéralisation de la MO par les bactéries dans la zone mésopélagique est loin d’être négligeable, spécialement dans une région dominée par les diatomées.</p> / Doctorat en Sciences agronomiques et ingénierie biologique / info:eu-repo/semantics/nonPublished
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A Reconnaissance Study of Water and Carbon Fluxes in Tropical Watersheds of Peninsular Malaysia: Stable Isotope ConstraintsIshak, Muhammad Izzuddin Syakir January 2014 (has links)
Evapotranspiration is a nexus for planetary energy and carbon cycles, as yet poorly constrained. Here I use stable isotopes of oxygen and hydrogen to partition flux of water due to plant transpiration from the direct evaporative flux from soils, water bodies and plant. The study areas, Langat and Kelantan watersheds represent examples of domains dominated by the respective Southwest and Northeast monsoons on the two sides of the main orographic barrier (Titiwangsa mountain range). Mean annual rainfall for the Langat watershed, obtained from 30 years of hydrological data, is 2145 ± 237 mm. Tentatively, 48% of this precipitation returns to the atmosphere via transpiration (T), with 33% partitioned into discharge (Q), 8% into interception (In), and 11% into evaporation (Ed). In the Kelantan watershed, the mean annual rainfall, also based on the 30 year hydrological data, is 2383 ± 120 mm. Similar to Langat, the T accounts for 43% of precipitation (P), 45% is discharged into South China Sea (Q), 12% partitioned into interception (In) and tentatively 0% for evaporation (Ed). Ed for the Langat watershed represents only a small proportion in terms of volumetric significance, up to almost ~11% with strong effect on the isotopic fingerprints of waters associated with the summer Southwest Monsoon (SWM). Note, however, that insignificant Ed for the Kelantan watershed may be an artefact of rain and river water sampling at only coastal downstream portion of the watershed. High humidity (80%) also was recorded for the Malaysian Peninsula watershed.
T appropriates about half of all solar energy absorbed by the continents, here ~1000*103 g H2O m-2 yr-1 similar to other tropical regions at 900-1200*103 g H2O m-2 yr-1. The associated carbon fluxes are ~ 1300 g C m-2yr-1, independent of P. Vegetation responses to solar irradiance, via T and photosynthesis reflects the importance of stomatal regulation of the water and carbon fluxes. In order to maintain high transpiration in the tropical region, “constant” water supply is required for continuous pumping of water that delivers nutrients to the plant, suggesting that water and carbon cycle are co-driven by the energy of the sun. The existence of the water conveyor belt may be precondition for nutrient delivery, hence operation of the carbon cycle. Potentially, this may change our perspective on the role that biology plays in the water cycle. In such perspective, the global water cycle is the medium that redistributes the incoming solar energy across the planet, and the anatomical structures of plants then help to optimize the loop of energy transfer via evaporation and precipitation in the hydrologic cycle.
The main features of aquatic geochemistry of the Langat and Kelantan rivers inferred from the Principal Component Analysis are controlled by three components that explain 80% and 82% of total variances. These components are reflecting of the geogenic factor with superimposed pollution, the latter particularly pronounced in urbanized sections of the Langat river and dominant in downstream of the Kelantan river. There is no correlation between seasonal variations in major ion chemistry and environmental variables such as precipitation, discharge, temperature or solar activity.
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Variability of summer CH4 and CO2 flux rates in and between three large Swedish lakes : A spatio-temporal field study / Sommarvariation i CH4- och CO2-flöden i och mellan tre stora svenska sjöar : En rumslig och tidsmässig fältstudieNilsson, Isak, Seifarth, Filip January 2020 (has links)
Understanding of natural greenhouse gas (GHG) cycles is crucial for making GHG budgets, which work as basis in climate change and global warming policy programs. Lakes as a source for GHG activity have only recently been included in global GHG budgets, and most studies of lake GHG flux rates are conducted on lakes <10 km2, which only comprise roughly half of the global lake area—making data of large lake flux rates scarce. CO2 and CH4 are the primary contributors of GHGs, and lakes house production processes and receive these gasses via lateral transport. This study utilized a floating chamber method with CO2 sensors to study CH4 and CO2 flux rates from three large Swedish lakes. To do this, chambers were anchored at shallow depth, as well as passively drifted on open water. Sampling was conducted during two periods in the summer 2019, late June–early July and August. For CH4, spatial difference was found between deep and shallow transects within lakes, no temporal difference was found between study periods. Difference between lakes within the deep and shallow chamber groups was found. One possible instance of deep-water ebullition was recorded, and a correlation between CH4 flux rate and water temperature was observed. For CO2, no difference between deep and shallow chambers or measurement periods was found. One instance within the deeper chamber group was found to be different between two of the lakes, despite all three lakes being of different size, depth and trophic state. The study results indicate CO2 water concentrations near saturation with atmosphere during the sampling periods. No correlation between CO2 flux rate and water temperature was observed. Unexpected small-scale variability patterns in CO2 flux was observed while chambers were passively drifting. While some observed patterns for the two gases could be explained by previous findings, some of our observations could not be explained on the basis of previous literature, highlighting the need for further study of GHG flux rates from large lakes. / Förståelsen av naturliga växthuscykler är avgörande för att göra budgetar för växthusgaser, eftersom dessa uppskattningar och budgetar agerar som grund i policyprogram för arbete med klimatförändringar och global uppvärmning. Sjöar har endast nyligen inkluderats i globala växthusgodsbudgetar som en källa för växthusgasutsläpp, och de flesta studier av flödeshastigheter genomförs på sjöar <10 km2, vilka endast utgör ungefär hälften av den globala sjöarealen –vilket leder till att data om växthusgasflöden från stora sjöar saknas. CO2och CH4 är de mest potenta växthusgaserna, och sjöar hyser produktionsprocesser samt tar emot dessa gaser från kringliggande miljöer. Denna studie nyttjade en kammarmetod med CO2-sensorer för att studera CH4och CO2-flödeshastigheter från tre stora svenska sjöar. Detta gjordes dels genom att ankra kammare på grunt djup, och dels genom att låta kammare driva passivt på öppet vatten. Provtagningen genomfördes under två perioder sommaren 2019, slutet av juni-början av juli och augusti. För CH4 hittades rumslig skillnad mellan djupa och grunda transekter i sjöar, men ingen tidsmässig skillnad hittades mellan studieperioder. Skillnader mellan sjöar i de djupa och grunda kammargrupperna hittades. Ett möjligt fall av metanbubblor från djupt vatten registrerades, liksom en korrelation mellan CH4-flödeshastighet och vattentemperatur. För CO2 hittades ingen skillnad mellan djupa och grunda kammare eller mätperioder. En skillnad i den djupa kammargruppen hittades mellan två av sjöarna, trots att alla tre var av olika storlek, djup och trofiklass. Studiens resultat indikerar att koncentrationer av CO2 i vatten var nära mättnad med atmosfären under studieperioderna. Ingen korrelation mellan CO2-flödeshastighet och vattentemperatur observerades. Oväntade småskaliga variabilitetsmönster i CO2-flöde observerades medan kamrar drev passivt. Medan vissa observerade mönster för de två gaserna kan förklaras av tidigare kunskap, kan andra av våra observationer inte förklaras av tidigare litteratur, och detta tydliggör behovet av fortsatt forskning på växthusgasflöden från stora sjöar. / <p>We also thank the METLAKE project funded by the European Research Council (ERC; grant agreement No 725546).</p> / METLAKE (ERC; grant agreement No 725546)
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Origine, composition et destinée de la matière organique dissoute et ses interactions avec les communautés de procaryotes dans la mer du LabradorLaBrie, Richard 12 1900 (has links)
Dans les océans, les procaryotes sont des acteurs clés dans le cycle du carbone puisqu’ils consomment une fraction importante de la matière organique dissoute (MOD) relâchée par les producteurs primaires. Puisque cette matière organique est très complexe et de biodisponibilité variable, les communautés de procaryotes qui la consomme sont très diversifiées et spécialisées pour certains types de composés organiques. En utilisant cette matière organique, les procaryotes contribuent à réintroduire ce carbone dans le réseau trophique, une source d’énergie essentielle dans les gyres oligotrophes de l’océan. Toutefois, puisque cette consommation n’est pas parfaite, une quantité importante de carbone est relâchée sous forme de CO2 lors de la respiration, mais aussi sous forme de MOD récalcitrante, contribuant à séquestrer du carbone dans les océans.
Le but de cette thèse est d’une part, de dresser un portrait global de la biodisponibilité de la MOD et d’autre part, de déterminer l’influence de la biodisponibilité de cette dernière sur la composition et le métabolisme des procaryotes dans la mer du Labrador, une mer dont le rôle est critique dans la régulation du climat. Plus spécifiquement, nous identifions pour la première fois comment la distribution spatiale des procaryotes influencent leur métabolisme et est influencée par leur préférence alimentaire dans les eaux de surface de la mer du Labrador. Finalement, nous regardons comment la matière organique produite en surface est transformée et séquestrée en profondeur suite à la convection hivernale dans la mer du Labrador.
Le budget de carbone dans les océans n’est toujours pas balancé. Afin de mieux connaître les sources et la biodisponibilité du carbone dans les différents milieux aquatiques, nous avons évalué la biodisponibilité de la MOD à travers le continuum aquatique, des lacs jusqu’à l’océan. En menant une méta-analyse sur le sujet, nos résultats montrent que la proportion de matière organique labile, c’est-à-dire facilement utilisable par les procaryotes, est d’environ 6% dans tous les environnements aquatiques. Toutefois, la proportion de matière organique semi-labile, celle qui nécessite plus de transformation par les procaryotes, est grandement liée à la proximité au milieu terrestre. Les seuls écosystèmes aquatiques déviant de ces deux constats sont ceux en période d’efflorescence algale: ils contiennent beaucoup plus de carbone labile et semi-labile que ceux à l’équilibre. Nous avons estimé que le carbone semi-labile peut soutenir 62% de la biomasse de procaryotes dans les lacs et les milieux côtiers.
Dans un deuxième temps, nous évaluons l’influence de la MOD sur le métabolisme et les communautés de procaryotes. Nous avons fait trois missions océanographiques sur la mer du Labrador à bord du navire Hudson pour déterminer la composition de la MOD et la communauté des procaryotes ainsi que leur métabolisme. En utilisant une approche novatrice, la modélisation de la distribution spatiale de l’abondance des procaryotes, nous avons montré à quel point celle-ci est importante pour déterminer leur préférence alimentaire ainsi que leur métabolisme. Nous avons également proposé un nouveau cadre conceptuel qui vise à faciliter la recherche à l’interface de la biogéochimie, de l’écologie microbienne et du métabolisme microbien.
Dans un dernier temps, nous avons comparé la capacité des procaryotes venant de différentes profondeurs océaniques à séquestrer le carbone. Lors de la consommation de la MOD, les procaryotes en relâche une petite fraction sous forme plus récalcitrante. En répétant ce processus, le carbone résiduel devient très récalcitrant et peut résister à la consommation par les procaryotes durant des centaines d’années. Nous avons montré que les procaryotes de l’océan profond sont plus efficaces pour séquestrer le carbone de cette façon. Nos résultats montrent que ce sont les taxons rares des procaryotes qui sont les éléments clés dans cette suite de transformation qui mène à la séquestration du carbone appelée pompe microbienne.
Cette thèse contribue à la compréhension du cycle du carbone dans la mer du Labrador et dans les écosystèmes aquatiques en général. Nous avons proposé une approche novatrice permettant de lier la qualité de la MOD à la composition des communautés de procaryotes qui la dégrade, un défi qui perdure depuis des dizaines d’années. De plus, nous montrons pour la première fois la que la pompe microbienne de carbone est un processus itératif fortement relié à la succession de la communauté de procaryotes. Nous montrons également que la pompe microbienne est active dans chaque strate océanique, mais que les procaryotes rares issus de l’océan profond sont plus efficaces à séquestrer le carbone. Mieux comprendre comment la composition de la MOD influence les procaryotes est primordial puisqu’ils sont centraux au cycle du carbone océanique. / Oceanic prokaryotes are key players in the carbon cycle by consuming dissolved organic mat-ter (DOM) produced by primary producers. As this organic matter is highly complex with varying degree of bioavailability, prokaryotic communities are highly diverse and different taxa target certain types of organic compounds. By consuming this organic matter, prokary-otes reintroduce this carbon into the food web, a critical energy flow in oligotrophic gyres. However, this consumption is not perfect and they release a lot of carbon as CO2 through respiration, but also as recalcitrant DOM. Thus, they contribute to carbon sequestration in aquatic ecosystems.
The objective of this thesis is to characterize DOM bioavailability and its influence on the composition and metabolism of prokaryotic communities in the Labrador Sea, described as one of the Earth’s climate system tipping elements. More precisely, we quantify for the first time how the spatial abundance distribution of prokaryotes influences ecosystem metabolism and organic matter association in the surface waters of the Labrador Sea. Lastly, we look at how DOM produced at the surface is transformed and sequestered following the Labrador Sea winter convective mixing.
The oceanic carbon budget is still unbalanced. In order to better understand its carbon sources and bioavailability, we characterize DOM bioavailability across the aquatic contin-uum, from lakes to the open ocean. Using a meta-analysis, our results show that the propor-tion of labile organic matter, i.e. readily available for prokaryotes, is similar at around 6% in all aquatic ecosystems. However, the proportion of semi-labile organic matter, i.e requiring transformations to be consumed by prokaryotes, is highly related to terrestrial connectivity. The only ecosystems that did not follow these patterns were in a phytoplankton bloom pe-riod and had a high proportion of labile and semi-labile organic matter as their counterparts at equilibrium. Finally, we estimated that semi-labile organic matter could sustain 62% of prokaryotic biomass in lakes and coastal zones.
Second, we evaluated the influence of DOM on prokaryotic metabolism and community composition. In order to determine organic matter composition, prokaryotic community composition and metabolic rates, we did three oceanic cruises in the Labrador Sea onboard the Hudson ship. By using spatial abundance distribution modelling of prokaryotes, we identified strong associations between groups of this novel approach and organic matter composition. We also proposed a framework to bridge the gap between prokaryotic diversity, microbial ecology, and biogeochemistry among methods and across scales.
Lastly, we compared how prokaryotic communities from different oceanic strata could se-quester carbon. When they consume organic matter, prokaryotes release a small amount in recalcitrant forms. Through this iterative process, called the microbial carbon pump, prokaryotes contribute to carbon sequestration by creating highly recalcitrant compounds that resist further degradation for hundreds of years. We have shown that all prokaryotes enable the microbial carbon pump, but that prokaryotes from deeper strata are more effi-cient. Our results also conclusively show that the rare prokaryotic taxa are key players in the microbial carbon pump.
This thesis contributes to better understand the carbon cycle in the Labrador Sea and in all aquatic ecosystems. We proposed a novel framework to relate biogeochemistry, prokaryotic diversity and microbial ecology which has been a challenge for decades. Moreover, we con-clusively showed for the first time that the iterative process of the microbial carbon pump is related to prokaryotic succession. We also show that it happens in all oceanic strata, but that rare prokaryotes from the deep ocean are more efficient to sequester carbon. Better understanding how DOM composition influences prokaryotes is of prime importance as they are the main drivers of the oceanic carbon cycle.
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