• Planète et Univers/Océan, Atmosphère
  
• Sciences de l'environnement/Biodiversité et Ecologie
  

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 the 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 averaged 28.9 ±  9.3 and 27.0 ±  7.2 % of total organic carbon (TOC) in the mesocosms and surrounding lagoon respectively and was strongly and positively coupled with TOC during P2 (i.e., days 15–23). TEP concentrations in the mesocosms declined for the first 9 days after DIP fertilization (P1  =  days 5–14) and then gradually increased during the second phase. 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 turnover time 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 cyanobacterial Group C (UCYN-C) diazotrophs that 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 were 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.

N2 fixation fuels  ∼  50 % of new primary production in the oligotrophic South Pacific Ocean. The VAHINE experiment has been designed to track the fate of diazotroph-derived nitrogen (DDN) and carbon within a coastal lagoon ecosystem in a comprehensive way. For this, large-volume ( ∼  50 m3) mesocosms were deployed in the New Caledonian lagoon and were intentionally fertilized with dissolved inorganic phosphorus (DIP) to stimulate N2 fixation. This study examined the temporal dynamics of the prokaryotic community together with the evolution of biogeochemical parameters for 23 consecutive days in one of these mesocosms (M1) and in the Nouméa lagoon using MiSeq 16S rRNA gene sequencing and flow cytometry. Combining these methods allowed for inference of absolute cell numbers from 16S data. We observed clear successions within M1, some of which were not mirrored in the lagoon. The dominating classes in M1 were Alpha- and Gammaproteobacteria, Cyanobacteria, eukaryotic microalgae, Marine Group II Euryarchaeota, Flavobacteriia, and Acidimicrobia. Enclosure led to significant changes in the M1 microbial community, probably initiated by the early decay of Synechococcus and diatoms. However, we did not detect a pronounced bottle effect with a copiotroph-dominated community. The fertilization with  ∼  0.8 µM DIP on day 4 did not have directly observable effects on the overall community within M1, as the data samples obtained from before and 4 days after fertilization clustered together, but likely influenced the development of individual populations later on, like Defluviicoccus-related bacteria and UCYN-C-type diazotrophic cyanobacteria (Cyanothece). Growth of UCYN-C led to among the highest N2-fixation rates ever measured in this region and enhanced growth of nearly all abundant heterotrophic groups in M1. We further show that different Rhodobacteraceae were the most efficient heterotrophs in the investigated system and we observed niche partitioning within the SAR86 clade. Whereas the location in- or outside the mesocosm had a significant effect on community composition, the temporal effect was significantly stronger and similar in both locations, suggesting that overarching abiotic factors were more influential than the enclosure. While temporal community changes were evident, prokaryotic diversity (Shannon index) only declined slightly from  ∼  6.5 to 5.7 or 6.05 in the lagoon and M1, respectively, throughout the experiment, highlighting the importance of multiple and varying sources of organic matter maintaining competition.

Studies investigating the fate of diazotrophs through the microbial food web are lacking, although N2 fixation can fuel up to 50 % of new production in some oligotrophic oceans. In particular, the role played by heterotrophic prokaryotes in this transfer is largely unknown. In the frame of the VAHINE (VAriability of vertical and tropHIc transfer of diazotroph derived N in the south wEst Pacific) experiment, three replicate large-volume (∼ 50 m3) mesocosms were deployed for 23 days in the new Caledonia lagoon and were intentionally fertilized on day 4 with dissolved inorganic phosphorus (DIP) to stimulate N2 fixation. We specifically examined relationships between heterotrophic bacterial production (BP) and N2 fixation or primary production, determined bacterial growth efficiency and established carbon budgets. BP was statistically higher during the second phase of the experiment (P2: days 15–23), when chlorophyll biomass started to increase compared to the first phase (P1: days 5–14). Phosphatase alkaline activity increased drastically during the second phase of the experiment, showing adaptations of microbial populations after utilization of the added DIP. Notably, among autotrophs, Synechococcus abundances increased during P2, possibly related to its capacity to assimilate leucine and to produce alkaline phosphatase. Bacterial growth efficiency based on the carbon budget (27–43 %), was notably higher than generally cited for oligotrophic environments and discussed in links with the presence of abundant species of bacteria expressing proteorhodopsin. The main fates of gross primary production (particulate + dissolved) were respiration (67 %) and export through sedimentation (17 %). BP was highly correlated with particulate primary production and chlorophyll biomass during both phases of the experiment but was slightly correlated, and only during P2 phase, with N2 fixation rates. Heterotrophic bacterial production was strongly stimulated after mineral N enrichment experiments, suggesting N-limitation of heterotrophic bacteria across the experiment. N2 fixation rates corresponded to 17–37 % of the nitrogen demand of heterotrophic bacteria. Our results suggest that most of the diazotroph-derived nitrogen fuelled the heterotrophic bacterial community through indirect processes generating dissolved organic matter and detritus, like mortality, lysis and grazing of both diazotrophs and non-diazotrophs.

In the framework of the VAHINE project, we investigated the spectral characteristics and the variability of dissolved and particulate chromophoric materials throughout a 23-day mesocosm experiment conducted in the southwest Pacific at the mouth of the New Caledonian coral lagoon (22 • 29.073 S–166 • 26.905 E) from 13 January to 4 Febru-ary 2013. Samples were collected in a mesocosm fertilized with phosphate at depths of 1, 6 and 12 m and in the surrounding waters. Light absorption coefficients of chro-mophoric dissolved organic matter (CDOM) [a g (λ)] and particulate matter [a p (λ)] were determined using a point-source integrating-cavity absorption meter (PSICAM), while fluorescent DOM (FDOM) components were determined from excitation-emission matrices (EEMs) combined with parallel factor analysis (PARAFAC). The evolutions of a g (λ) and a p (λ) in the mesocosm were similar to those of total chlorophyll a concentration, Synechococcus spp. and picoeukary-ote abundances, bacterial production, particulate organic nitrogen and total organic carbon concentrations, with roughly a decrease from the beginning of the experiment to days 9– 10, and an increase from days 9–10 to the end of the experiment. In the surrounding waters, the same trend was observed but the increase was much less pronounced, emphasizing the effect of the phosphate fertilization on the mesocosm's plankton community. Correlations suggested that both Syne-chococcus cyanobacteria and heterotrophic bacteria were strongly involved in the production of CDOM and absorption of particulate matter. The increase in phytoplankton biomass during the second part of the experiment led to a higher contribution of particulate material in the absorption budget at 442 nm. The three FDOM components identified (tryptophan-, tyrosine-and ultraviolet C (UVC) humic-like fluorophores) did not follow the evolution of CDOM and particulate matter, suggesting they were driven by different production/degradation processes. Finally, the results of this work support the idea there is indirect coupling between the dynamics of N 2 fixation and that of chromophoric material via the stimulation of Synechococcus bloom.