ALBERT

Description: Dinitrogen (N2) fixation rates were measured during early spring across the different provinces of Mediterranean Sea surface waters. N2 fixation rates, measured using 15N2 enriched seawater, were lowest in the eastern basin and increased westward with a maximum at the Strait of Gibraltar (0.10 to 2.35 nmol N L−1 d−1, respectively). These rates were 3–7 fold higher than N2 fixation rates measured previously in the Mediterranean Sea during summertime. Moreover, comparisons between N2 fixation rates measured during dark versus natural light incubations (48 h) show higher rates during dark incubations at the eastern Mediterranean stations but lower rates at the western stations. This suggests that heterotrophic diazotrophy has a significant role in the Eastern Mediterranean while autotrophic diazotrophy has a more dominant role in the Western basin.

Description: Dinitrogen (N2) fixation rates were measured during early spring across the different provinces of Mediterranean Sea surface waters. N2 fixation rates, measured using 15N2 enriched seawater, were lowest in the eastern basin and increased westward with a maximum at the Strait of Gibraltar (0.10 to 2.35 nmol N L−1 d−1, respectively). These rates were 3–7 fold higher than N2 fixation rates measured previously in the Mediterranean Sea during summertime and we estimated that methodological differences alone did not account for the seasonal changes we observed. Higher contribution of N2 fixation to primary production (4–8%) was measured in the western basin compared to the eastern basin (∼2%). Our data indicates that these differences between basins may be attributed to changes in N2-fixing planktonic communities and that heterotrophic diazotrophy may play a significant role in the eastern Mediterranean while autotrophic diazotrophy has a more dominant role in the western basin.

Description: Marine N2 fixing microorganisms, termed diazotrophs, are a key functional group in marine pelagic ecosystems. The biological fixation of dinitrogen (N2) to bioavailable nitrogen provides an important new source of nitrogen for pelagic marine ecosystems and influences primary productivity and organic matter export to the deep ocean. As one of a series of efforts to collect biomass and rates specific to different phytoplankton functional groups, we have constructed a database on diazotrophic organisms in the global pelagic upper ocean by compiling about 12 000 direct field measurements of cyanobacterial diazotroph abundances (based on microscopic cell counts or qPCR assays targeting the nifH genes) and N2 fixation rates. Biomass conversion factors are estimated based on cell sizes to convert abundance data to diazotrophic biomass. The database is limited spatially, lacking large regions of the ocean especially in the Indian Ocean. The data are approximately log-normal distributed, and large variances exist in most sub-databases with non-zero values differing 5 to 8 orders of magnitude. Reporting the geometric mean and the range of one geometric standard error below and above the geometric mean, the pelagic N2 fixation rate in the global ocean is estimated to be 62 (52–73) Tg N yr−1 and the pelagic diazotrophic biomass in the global ocean is estimated to be 2.1 (1.4–3.1) Tg C from cell counts and to 89 (43–150) Tg C from nifH-based abundances. Reporting the arithmetic mean and one standard error instead, these three global estimates are 140 ± 9.2 Tg N yr−1, 18 ± 1.8 Tg C and 590 ± 70 Tg C, respectively. Uncertainties related to biomass conversion factors can change the estimate of geometric mean pelagic diazotrophic biomass in the global ocean by about ±70%. It was recently established that the most commonly applied method used to measure N2 fixation has underestimated the true rates. As a result, one can expect that future rate measurements will shift the mean N2 fixation rate upward and may result in significantly higher estimates for the global N2 fixation. The evolving database can nevertheless be used to study spatial and temporal distributions and variations of marine N2 fixation, to validate geochemical estimates and to parameterize and validate biogeochemical models, keeping in mind that future rate measurements may rise in the future. The database is stored in PANGAEA (doi:10.1594/PANGAEA.774851).

Description: We evaluated the seasonal contribution of heterotrophic and autotrophic diazotrophy to the total dinitrogen (N2) fixation in a representative pelagic station in the northern Gulf of Aqaba in early spring when the water column was mixed and during summer under full thermal stratification. N2 fixation rates were low during the mixed period (∼ 0.1 nmol N L−1 d−1) and were significantly coupled with both primary and bacterial productivity. During the stratified period N2 fixation rates were four-fold higher (∼ 0.4 nmol N L−1 d−1) and were significantly correlated solely with bacterial productivity. Furthermore, while experimental enrichment of seawater by phosphorus (P) enhanced bacterial productivity and N2 fixation rates during both seasons primary productivity was stimulated by P only in the early spring. Metatranscriptomic analyses from the stratified period identified the major diazotrophic contributors as related to heterotrophic prokaryotes from the Euryarchaeota and Desulfobacterales (Deltaproteobacteria) or Chlorobiales (Chlorobia). Moreover, during this season, experimental amendments to seawater applying a combination of the photosynthetic inhibitor 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) and a mixture of amino acids increased both bacterial productivity and N2 fixation rates. Our findings from the northern Gulf of Aqaba indicate a~shift in the diazotrophic community from phototrophic and heterotrophic populations, including small blooms of the cyanobacterium Trichodesmium, in winter/early spring, to predominantly heterotrophic diazotrophs in summer that may be both P and carbon limited as the additions of P and amino acids illustrated.

Description: N2 fixation rates were measured daily in large (~ 50 m3) mesocosms deployed in the tropical South West Pacific coastal ocean (New Caledonia) to investigate the spatial and temporal dynamics of diazotrophy and the fate of diazotroph-derived nitrogen (DDN) in a low nutrient, low chlorophyll ecosystem. The mesocosms were intentionally fertilized with ~ 0.8 μM dissolved inorganic phosphorus (DIP) to stimulate diazotrophy. Bulk N2 fixation rates were replicable between the three mesocosms, averaged 18.5 ± 1.1 nmol N L−1 d−1 over the 23 days, and increased by a factor of two during the second half of the experiment (days 15 to 23) to reach 27.3 ± 1.0 nmol N L−1 d−1. These rates are higher than the upper range reported for the global ocean, indicating that the waters surrounding New Caledonia are particularly favourable for N2 fixation. During the 23 days of the experiment, N2 fixation rates were positively correlated with seawater temperature, primary production, bacterial production, standing stocks of particulate organic carbon, nitrogen and phosphorus, and alkaline phosphatase activity, and negatively correlated with DIP concentrations, DIP turnover time, nitrate, and dissolved organic nitrogen and phosphorus concentrations. The fate of DDN was investigated during the bloom of the unicellular diazotroph, UCYN-C, that occurred during the second half of the experiment. Quantification of diazotrophs in the sediment traps indicates that ~ 10 % of UCYN-C from the water column were exported daily to the traps, representing as much as 22.4 ± 5.5 % of the total POC exported at the height of the UCYN-C bloom. This export was mainly due to the aggregation of small (5.7 ± 0.8 μm) UCYN-C cells into large (100–500 μm) aggregates. During the same time period, a DDN transfer experiment based on high-resolution nanometer scale secondary ion mass spectrometry (nanoSIMS) coupled with 15N2 isotopic labelling revealed that 16 ± 6 % of the DDN was released to the dissolved pool and 21 ± 4 % was transferred to non-diazotrophic plankton, mainly picoplankton (18 ± 4 %) followed by diatoms (3 ± 2 %) within 24 h of incubation. This is consistent with the observed dramatic increase in picoplankton and diatom abundances, primary production, bacterial production and standing stocks of particulate organic carbon, nitrogen and phosphorus during the second half of the experiment in the mesocosms. These results offer insights into the fate of DDN during a bloom of UCYN-C in low nutrient, low chlorophyll ecosystems.

Description: A combined dataset of near real time multi-satellite observations and in situ measurements from a high-resolution survey, is used for characterizing physical-biogeochemical properties of a patch stretching from the coast to the open sea in the Levantine basin of the Eastern Mediterranean. Spatial analysis of the combined dataset indicates that the patch is a semi-enclosed system, bounded within the mixed layer and separated from ambient waters by transport barriers induced by horizontal stirring. As such, the patch is characterized by physical-biogeochemical properties that significantly differ from those of the waters surrounding it, with lower salinity, higher temperatures, higher concentrations of silicic acid and chlorophyll a, and higher abundance of Synechococcus and Picoeukaryotes cells. Based on estimates of patch dimensions (~ 40 km width and ~ 25 m depth) and propagation speed (~ 0.09 m s−1), the volume flux associated with the patch is found to be in the order of 0.1 Sv. Our observations suggest that horizontal stirring by surface currents is likely to have an important impact on the ultra-oligotrophic Levantine basin ecosystem, through (1) transport of nutrients and coastal derived material, and (2) formation of local, dynamically isolated, niches. In addition, this work provides a satellite-based framework for planning and executing high resolution sampling strategies in the interface between coast and the open sea.

Description: In the marine environment, transparent exopolymeric particles (TEP) produced from abiotic and biotic sources link the particulate and dissolved carbon pools and are essential vectors enhancing vertical carbon flux. We characterized spatial and temporal dynamics of TEP during the VAHINE experiment that investigated the fate of diazotroph derived nitrogen and carbon in three, replicate, dissolved inorganic phosphorus (DIP)-fertilized 50 m3 enclosures in an oligotrophic New Caledonian lagoon. During the 23 days of the experiment, we did not observe any depth dependent changes in TEP concentrations in the three sampled-depths (1, 6, 12 m). TEP carbon (TEP-C) content per mesocosm averaged 28.9 ± 9.3% and 27.0 ± 7.2% of TOC in the mesocosms and surrounding lagoon respectively and was strongly and positively coupled with TOC during P2. TEP concentrations declined for the first 9 days after DIP fertilization (P1 = days 5-14) and then gradually increased during the second phase (P2 = days 15-23). Temporal changes in TEP concentrations paralleled the growth and mortality rates of the diatom-diazotroph association of Rhizosolenia and Richelia that predominated the diazotroph community during P1. By P2, increasing total primary and heterotrophic bacterial production consumed the supplemented P and reduced availability of DIP. For this period, TEP concentrations were negatively correlated with DIP availability and turnovertime of DIP (TDIP) while positively associated with enhanced alkaline phosphatase activity (APA) that occurs when the microbial populations are P-stressed. During P2, increasing bacterial production (BP) was positively correlated with higher TEP concentrations which were also coupled with the increased growth rates and aggregation of the unicellular UCYN-C diazotrophs which bloomed during this period. We conclude that the composite processes responsible for the formation and breakdown of TEP yielded a relatively stable TEP pool available as both a carbon source and facilitating aggregation and flux throughout the experiment. TEP was probably mostly influenced by abiotic physical processes during P1 while biological activity (BP, diazotrophic growth and aggregation, export production) mainly impacted TEP concentrations during P2 when DIP-availability was limited.

Description: A combined dataset of near-real-time multi-satellite observations and in situ measurements from a high-resolution survey is used for characterizing physical-biogeochemical properties of a patch stretching from the coast to the open sea in the Levantine Basin (LB) of the eastern Mediterranean (EM). Spatial analysis of the combined dataset indicates that the patch is a semi-enclosed system, bounded within the mixed layer and separated from ambient waters by transport barriers induced by horizontal stirring. As such, the patch is characterized by physical-biogeochemical properties that significantly differ from those of the waters surrounding it, with lower salinity and higher temperatures, concentrations of silicic acid and chlorophyll a, and abundance of Synechococcus and picoeukaryote cells. Based on estimates of patch dimensions (∼40 km width and ∼25 m depth) and propagation speed (∼0.09 m s−1), the volume flux associated with the patch is found to be on the order of 0.1 Sv. Our observations suggest that horizontal stirring by surface currents is likely to have an important impact on the ultra-oligotrophic Levantine Basin ecosystem, through (1) transport of nutrients and coastally derived material, and (2) formation of local, dynamically isolated niches. In addition, this work provides a satellite-based framework for planning and executing high-resolution sampling strategies in the interface between the coast and the open sea.