Constitutive Extracellular Polysaccharide (EPS) Production by Specific Isolates of Crocosphaera watsonii

Unicellular dinitrogen (N2) fixing cyanobacteria have only recently been identified in the ocean and recognized as important contributors to global N2 fixation. The only cultivated representatives of the open ocean unicellular diazotrophs are multiple isolates of Crocosphaera watsonii. Although constituents of the genus are nearly genetically identical, isolates have been described in two size classes, large ∼5 μm and small ∼3 μm cell diameters. We show here that the large size class constitutively produces substantial amounts of extracellular polysaccharides (EPS) during exponential growth, up to 10 times more than is seen in the small size class, and does so under both N2 fixing and non-N2 fixing conditions. The EPS production exceeds the amount produced by larger phytoplankton such as diatoms and coccolithophores by one to two orders of magnitude, is ∼22% of the total particulate organic C in the culture, and is depleted in N compared to cellular material. The large difference in observed EPS production may be accounted for by consistently higher photochemical efficiency of photosystem II in the large (0.5) vs. small (∼0.35) strains. While it is known that Crocosphaera plays an important role in driving the biological carbon (C) pump through the input of new nitrogen (N) to the open ocean, we hypothesize that this species may also contribute directly to the C cycle through the constitutive production of EPS. Indeed, at two stations in the North Pacific Subtropical Gyre, ∼70% of large Crocosphaera cells observed were embedded in EPS. The evolutionary advantage of releasing such large amounts of fixed C is still unknown, but in regions where Crocosphaera can be abundant (i.e., the warm oligotrophic ocean) this material will likely have important biogeochemical consequences

Western Pacific atmospheric nutrient deposition fluxes, their impact on surface ocean productivity

The atmospheric deposition of both macronutrients and micronutrients plays an important role in driving primary productivity, particularly in the low-latitude ocean. We report aerosol major ion measurements for five ship-based sampling campaigns in the western Pacific from similar to 25 degrees N to 20 degrees S and compare the results with those from Atlantic meridional transects (similar to 50 degrees N to 50 degrees S) with aerosols collected and analyzed in the same laboratory, allowing full incomparability. We discuss sources of the main nutrient species (nitrogen (N), phosphorus (P), and iron (Fe)) in the aerosols and their stoichiometry. Striking north-south gradients are evident over both basins with the Northern Hemisphere more impacted by terrestrial dust sources and anthropogenic emissions and the North Atlantic apparently more impacted than the North Pacific. We estimate the atmospheric supply rates of these nutrients and the potential impact of the atmospheric deposition on the tropical western Pacific. Our results suggest that the atmospheric deposition is P deficient relative to the needs of the resident phytoplankton. These findings suggest that atmospheric supply of N, Fe, and P increases primary productivity utilizing some of the residual excess phosphorus (P*) in the surface waters to compensate for aerosol P deficiency. Regional primary productivity is further enhanced via the stimulation of nitrogen fixation fuelled by the residual atmospheric iron and P*. Our stoichiometric calculations reveal that a P* of 0.1 mu mol L-1 can offset the P deficiency in atmospheric supply for many months. This study suggests that atmospheric deposition may sustain similar to 10% of primary production in both the western tropical Pacific

Effects of African dust deposition on phytoplankton in the western tropical Atlantic Ocean off Barbados

Bioassay incubation experiments conducted with nutrients and local atmospheric aerosol amendments indicate that phosphorus (P) availability limited phytoplankton growth in the low-nutrient low-chlorophyll (LNLC) ocean off Barbados. Atmospheric deposition provides a relatively large influx of new nutrients and trace metals to the surface ocean in this region in comparison to other nutrient sources. However, the impact on native phytoplankton is muted due to the high ratio of nitrogen (N) to P (NO3:SRP > 40) and the low P solubility of these aerosols. Atmospheric deposition induces P limitation in this LNLC region by adding more N and iron (Fe) relative to P. This favors the growth of Prochlorococcus, a genus characterized by low P requirements and highly efficient P acquisition mechanisms. A global three-dimensional marine ecosystem model that includes species-specific phytoplankton elemental quotas/stoichiometry and the atmospheric deposition of N, P, and Fe supports this conclusion. Future increases in aerosol N loading may therefore influence phytoplankton community structure in other LNLC areas, thereby affecting the biological pump and associated carbon sequestration

Abundances of Iron-Binding Photosynthetic and Nitrogen-Fixing Proteins of Trichodesmium Both in Culture and In Situ from the North Atlantic

Marine cyanobacteria of the genus Trichodesmium occur throughout the oligotrophic tropical and subtropical oceans, where they can dominate the diazotrophic community in regions with high inputs of the trace metal iron (Fe). Iron is necessary for the functionality of enzymes involved in the processes of both photosynthesis and nitrogen fixation. We combined laboratory and field-based quantifications of the absolute concentrations of key enzymes involved in both photosynthesis and nitrogen fixation to determine how Trichodesmium allocates resources to these processes. We determined that protein level responses of Trichodesmium to iron-starvation involve down-regulation of the nitrogen fixation apparatus. In contrast, the photosynthetic apparatus is largely maintained, although re-arrangements do occur, including accumulation of the iron-stress-induced chlorophyll-binding protein IsiA. Data from natural populations of Trichodesmium spp. collected in the North Atlantic demonstrated a protein profile similar to iron-starved Trichodesmium in culture, suggestive of acclimation towards a minimal iron requirement even within an oceanic region receiving a high iron-flux. Estimates of cellular metabolic iron requirements are consistent with the availability of this trace metal playing a major role in restricting the biomass and activity of Trichodesmium throughout much of the subtropical ocean

A re-evaluation of the magnitude and impacts of anthropogenic atmospheric nitrogen inputs on the ocean

We report a new synthesis of best estimates of the inputs of fixed nitrogen to the world ocean via atmospheric deposition, and compare this to fluvial inputs and di-nitrogen fixation. We evaluate the scale of human perturbation of these fluxes. Fluvial inputs dominate inputs to the continental shelf, and we estimate about 75% of this fluvial nitrogen escapes from the shelf to the open ocean. Biological di-nitrogen fixation is the main external source of nitrogen to the open ocean, i.e. beyond the continental shelf. Atmospheric deposition is the primary mechanism by which land based nitrogen inputs, and hence human perturbations of the nitrogen cycle, reach the open ocean. We estimate that anthropogenic inputs are currently leading to an increase in overall ocean carbon sequestration of ~0.4% (equivalent to an uptake of 0.15 Pg C yr-1 and less than the Duce et al., 2008 estimate). The resulting reduction in climate change forcing from this ocean CO2 uptake is offset to a small extent by an increase in ocean N2O emissions. We identify four important feedbacks in the ocean atmosphere nitrogen system that need to be better quantified to improve our understanding of the perturbation of ocean biogeochemistry by atmospheric nitrogen inputs. These feedbacks are recycling of (1) ammonia and (2) organic nitrogen from the ocean to the atmosphere and back, (3) the suppression of nitrogen fixation by increased nitrogen concentrations in surface waters from atmospheric deposition, and (4) increased loss of nitrogen from the ocean by denitrification due to increased productivity stimulated by atmospheric inputs

Combining autocracy and majority voting: the canonical succession rules of the Latin Church

The autocratic turn of the Latin Church in the XI-XIII century, a reaction to the secular power interferences, concentrated the decision-making power in the hands of the top hierarchy, and finally in the hands of the pope. A fundamental step was the change and the constitutionalisation of the procedures for leadership replacement, which were open successions where the contest for power was governed by elections. The autocratic reform limited the active electorate to the clergy only and gradually substituted the episcopal elections by the pope’s direct appointment. Besides, the voting rules changed from unanimity to the dual principle of maioritas et sanioritas (where the majority was identified with the greater part by number and wisdom) and finally to the numerical rule of qualified majority. This evolution aimed at preserving the elections from external interferences and at eliminating the elements of arbitrariness. The most important succession, the papal election, was protected by institutionalising a selectorate and its decision-making rules. The selectorate and the elections did not insert accountability and representation mechanisms but only protected the quality of the autocratic leadership and its autonomy

Global-scale variations of the ratios of carbon to phosphorus in exported marine organic matter

The ratio of carbon (C) to phosphorus (P) in marine phytoplankton is thought to be constant throughout the worlds'oceans. Known as the Redfield ratio, this relationship describes the links between carbon and phosphorus cycling and marine ecosystems. However, variations in the stoichiometry of phytoplankton have recently been identified, in particular strong latitudinal variability. Here we assess the impact of this variability in the C:P ratio of biomass on the C:P ratio of organic matter that is exported to the deep ocean using a biogeochemical inverse-model based on a data-constrained ocean circulation model and a global database of dissolved inorganic carbon and phosphate measurements. We identify global patterns of variability in the C:P ratios of exported organic matter, with higher values in the nutrient-depleted subtropical gyres, where organic matter export is relatively low, and lower ratios in nutrient-rich upwelling zones and high-latitude regions, where organic matter export is high. This suggests that total carbon export is relatively constant throughout the oceans, in agreement with recent estimates of carbon fluxes. We conclude that the latitudinal patterns of C:P in exported organic matter are consistent with the large-scale stoichiometric variations in phytoplankton C:P. We suggest that a future expansion of nutrient-depleted waters could result in a shift to more efficient C export that compensates for the expected decline in productivity

Transferts électrodynamiques et hydrodynamiques en électrodialyse

Après le rappel du principe de l'électrodialyse, les membranes ioniques permsélectives, triant les ions selon le signe de !eur charge, sont décrites. Le nombre de transport d'un ion dans la membrane qui lui est perméable est beaucoup plus élevé que dans la solution, d'où l'apparition d'une polarisation de concentration qui entraîne elle-même une limite de la densité de courant. Cette intensité limite dépend du régime hydrodynamique de la cellule, et la théorie permet de préciser le couplage entre les transferts électrodynamiques et les transferts par diffusion et convection. On en déduit les conséquences sur la technologie et les performances des appareils, ainsi que sur les domaines d'application de l'électrodialyse