ALBERT

Description: Dinitrogen fixation is now recognized as one of the major sources of bio-available nitrogen in the ocean. Thus, N2 fixation sustains a significant part of the global primary production by supplying the most common limiting nutrient for phytoplankton growth. The “Oligotrophy to UlTra-oligotrophy PACific Experiment” (OUTPACE) improved the data coverage of the western tropical South Pacific, an area recently recognized as a hotspot of N2 fixation. This new development leads us to develop and test an explicit N2 fixation formulation based on the Trichodesmium physiology (the most studied nitrogen fixer) within a 3-D coupled dynamical–biogeochemical model (ROMS-PISCES). We performed a climatological numerical simulation that is able to reproduce the main physical (e.g. sea surface temperature) and biogeochemical patterns (nutrient and chlorophyll concentrations, as well as N2 fixation) in the tropical Pacific. This simulation displayed a Trichodesmium regional distribution that extends from 150∘ E to 120∘ W in the south tropical Pacific, and from 120∘ E to 140∘ W in the north tropical Pacific. The local simulated maximuma were found around islands (Hawaii, Fiji, Samoa, New Caledonia, Vanuatu). We assessed that 15 % of the total primary production may be due to Trichodesmium in the low-nutrient low-chlorophyll regions (LNLC) of the tropical Pacific. Comparison between our explicit and the often used (in biogeochemical models) implicit parameterization of N2 fixation showed that the latter leads to an underestimation of N2 fixation rates by about 25 % in LNLC regions. Finally, we established that iron fluxes from island sediments control the spatial distribution of Trichodesmium biomasses in the western tropical South Pacific. Note, this last result does not take into account the iron supply from rivers and hydrothermal sources, which may well be of importance in a region known for its strong precipitation rates and volcanic activity.

Description: We assessed the influence of the marine diazotrophic cyanobacterium Trichodesmium on the bio-optical properties of western tropical South Pacific (WTSP) waters (18–22∘ S, 160∘ E–160∘ W) during the February–March 2015 OUTPACE cruise. We performed measurements of backscattering and absorption coefficients, irradiance, and radiance in the euphotic zone with a Satlantic MicroPro free-fall profiler and took Underwater Vision Profiler 5 (UPV5) pictures for counting the largest Trichodesmium spp. colonies. Pigment concentrations were determined by fluorimetry and high-performance liquid chromatography and picoplankton abundance by flow cytometry. Trichome concentration was estimated from pigment algorithms and validated by surface visual counts. The abundance of large colonies counted by the UVP5 (maximum 7093 colonies m−3) was well correlated to the trichome concentrations (maximum 2093 trichomes L−1) with an aggregation factor of 600. In the Melanesian archipelago, a maximum of 4715 trichomes L−1 was enumerated in pump samples (3.2 m) at 20∘ S, 167 30∘ E. High Trichodesmium abundance was always associated with absorption peaks of mycosporine-like amino acids (330, 360 nm) and high particulate backscattering, but not with high Chl a fluorescence or blue particulate absorption (440 nm). Along the west-to-east transect, Trichodesmium together with Prochlorococcus represented the major part of total chlorophyll concentration; the contribution of other groups were relatively small or negligible. The Trichodesmium contribution to total chlorophyll concentration was the highest in the Melanesian archipelago around New Caledonia and Vanuatu (60 %), progressively decreased to the vicinity of the islands of Fiji (30 %), and reached a minimum in the South Pacific Gyre where Prochlorococcus dominated chlorophyll concentration. The contribution of Trichodesmium to zeaxanthin was respectively 50, 40 and 20 % for these regions. During the OUTPACE cruise, the relationship between normalized water-leaving radiance (nLw) in the ultraviolet and visible and chlorophyll concentration was similar to that found during the BIOSOPE cruise in the eastern tropical Pacific. Principal component analysis (PCA) of OUTPACE data showed that nLw at 305, 325, 340, 380, 412 and 440 nm was strongly correlated to chlorophyll and zeaxanthin, while nLw at 490 and 565 nm exhibited lower correlations. These results, as well as differences in the PCA of BIOSOPE data, indicated that nLw variability in the greenish blue and yellowish green during OUTPACE was influenced by other variables associated with Trichodesmium presence, such as backscattering coefficient, phycoerythrin fluorescence and/or zeaxanthin absorption, suggesting that Trichodesmium detection should involve examination of nLw in this spectral domain.

Description: Trichodesmium is the major nitrogen-fixing species in the western tropical South Pacific (WTSP) region, a hot spot of diazotrophy. Due to the paucity of in situ observations, remote-sensing methods for detecting Trichodesmium presence on a large scale have been investigated to assess the regional-to-global impact of this organism on primary production and carbon cycling. A number of algorithms have been developed to identify Trichodesmium surface blooms from space, but determining with confidence their accuracy has been difficult, chiefly because of the scarcity of sea-truth information at the time of satellite overpass. Here, we use a series of new cruises as well as airborne surveys over the WTSP to evaluate their ability to detect Trichodesmium surface blooms in the satellite imagery. The evaluation, performed on MODIS data at 250 m and 1 km resolution acquired over the region, shows limitations due to spatial resolution, clouds, and atmospheric correction. A new satellite-based algorithm is designed to alleviate some of these limitations, by exploiting optimally spectral features in the atmospherically corrected reflectance at 531, 645, 678, 748, and 869 nm. This algorithm outperforms former ones near clouds, limiting false positive detection and allowing regional-scale automation. Compared with observations, 80 % of the detected mats are within a 2 km range, demonstrating the good statistical skill of the new algorithm. Application to MODIS imagery acquired during the February-March 2015 OUTPACE campaign reveals the presence of surface blooms northwest and east of New Caledonia and near 20∘ S–172∘ W in qualitative agreement with measured nitrogen fixation rates. Improving Trichodesmium detection requires measuring ocean color at higher spectral and spatial (

Description: 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.

Description: On the global scale, N2 fixation provides the major external source of reactive nitrogen to the surface ocean, surpassing atmospheric and riverine inputs, and sustains  ∼  50 % of new primary production in oligotrophic environments. The main goal of the VAriability of vertical and tropHIc transfer of diazotroph derived N in the south wEst Pacific (VAHINE) project was to study the fate of nitrogen newly fixed by diazotrophs (or diazotroph-derived nitrogen) in oceanic food webs, and how it impacts heterotrophic bacteria, phytoplankton and zooplankton dynamics, stocks and fluxes of biogenic elements and particle export. Three large-volume ( ∼  50 m3) mesocosms were deployed in a tropical oligotrophic ecosystem (the New Caledonia lagoon, south-eastern Pacific) and intentionally fertilized with  ∼  0.8 µM of dissolved inorganic phosphorus (DIP) to stimulate diazotrophy and follow subsequent ecosystem changes. VAHINE was a multidisciplinary project involving close collaborations between biogeochemists, molecular ecologist, chemists, marine opticians and modellers. This introductory paper describes in detail the scientific objectives of the project as well as the implementation plan: the mesocosm description and deployment, the selection of the study site (New Caledonian lagoon), and the logistical and sampling strategy. The main hydrological and biogeochemical conditions of the study site before the mesocosm deployment and during the experiment itself are described, and a general overview of the papers published in this special issue is presented.

Description: The VAHINE mesocosm experiment was designed to trigger a diazotroph bloom and to follow the subsequent transfer of diazotroph-derived nitrogen (DDN) in the rest of the food web. Three mesocosms (50 m3) located inside the Nouméa lagoon (New Caledonia, southwestern Pacific) were enriched with dissolved inorganic phosphorus (DIP) in order to promote N2 fixation in these low-nutrient, low-chlorophyll (LNLC) waters. Initially, the diazotrophic community was dominated by diatom diazotroph associations (DDAs), mainly by Rhizosolenia/Richelia intracellularis, and by Trichodesmium, which fueled enough DDN to sustain the growth of other diverse diatom species and Synechococcus populations that were well adapted to limiting DIP levels. After DIP fertilization (1 µM) on day 4, an initial lag time of 10 days was necessary for the mesocosm ecosystems to start building up biomass. However, changes in community structure were already observed during this first period, with a significant drop of both Synechococcus and diatom populations, while Prochlorococcus benefited from DIP addition. At the end of this first period, corresponding to when most added DIP was consumed, the diazotroph community changed drastically and became dominated by Cyanothece-like (UCYN-C) populations, which were accompanied by a monospecific bloom of the diatom Cylindrotheca closterium. During the second period, biomass increased sharply together with primary production and N2-fixation fluxes near tripled. Diatom populations, as well as Synechococcus and nanophytoeukaryotes, showed a re-increase towards the end of the experiment, showing efficient transfer of DDN to non-diazotrophic phytoplankton.

Description: 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 south-west Pacific at the mouth of the New Caledonian coral lagoon (22°29.073 S–166°26.905 E) from 13 January to 4 February 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 chromophoric dissolved organic matter (CDOM) [ag(λ)] and particulate matter [ap(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 ag(λ) and ap(λ) in the mesocosm were similar to those of total chlorophyll a concentration, Synechococcus spp. and picoeukaryote 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 Synechococcus 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 N2 fixation and that of chromophoric material via the stimulation of Synechococcus bloom.

Description: Dinitrogen fixation is now recognized as one of the major sources of bio-available nitrogen in the ocean. Thus, nitrogen fixation sustains a significant part of the global primary production by providing an input of the most common limiting nutrient for phytoplankton growth. Evidences of the Western Tropical South Pacific being a hotspot of nitrogen fixation, and a data coverage complemented by OUTPACE, lead us to develop an explicit nitrogen fixation compartment based on the Trichodesmium physiology (the most studied nitrogen fixer) within a 3D coupled dynamical-biogeochemical model (ROMS-PISCES). We performed a first 20-year tropical Pacific simulation that is able to reproduce the main physical (e.g. Sea Surface Temperature) and biogeochemical conditions (nutrients, and chlorophyll concentrations as well as dinitrogen fixation). This simulation showed a possible Trichodesmium regional distribution that extends from 150° E to 120° W in the south tropical Pacific, and from 120° E to 140° W in the north tropical Pacific. The local simulated maximums were around islands (Hawaii, Fiji, Samoa, New Caledonia, Vanuatu). We assessed that 15 % of the total primary production may be due to Trichodesmium in the Low Nutrient, Low Chlorophyll regions (LNLC). We also argue that implicit parameterization of N2 fixation (often used in biogeochemical models) leads to underestimate nitrogen fixation rates by about 25% in LNLC regions compared to our explicit formulation. Finally, we showed that iron fluxes from island sediments control the spatial distribution and the abundance of Trichodesmium in the western tropical south Pacific. Noteworthy, this last result does not take into account the iron supply from rivers and hydrothermal sources, which may well be of importance in a region known for its strong precipitation rates and volcanic activity.

Description: Trichodesmium is the main nitrogen-fixing species in the South Pacific region, a hotspot for diazotrophy. Due to the paucity of in situ observations, methods for detecting Trichodesmium presence on a large scale have been investigated to assess the regional-to-global impact of these species on primary production and carbon cycling. A number of satellite-derived algorithms have been developed to identify Trichodesmium surface blooms, but determining with confidence their accuracy has been difficult, chiefly because of the scarcity of sea-truth information at time of satellite overpass. Here, we use a series of new cruises as well as airborne observational surveys in the South Pacific to quantify statistically the ability of these algorithms to discern correctly Trichodesmium surface blooms in the satellite imagery. The evaluation, performed on MODIS data at 250 m and 1 km resolution acquired over the South West Pacific, shows limitations due to spatial resolution, clouds, and atmospheric correction. A new satellite-based algorithm is designed to alleviate some of these limitations, by exploiting optimally spectral features in the atmospherically corrected reflectance at 531, 645, 678, 748, and 869 nm. This algorithm outperforms former ones near clouds, limiting false positive detection, and allowing regional scale automation. Compared with observations, 80 % of the detected mats are within a 2 km range, demonstrating the good statistical skill of the new algorithm. Application to MODIS imagery acquired during the February–March 2015 OUTPACE campaign reveals the presence of surface blooms Northwest and East of New Caledonia and near 20° S–172° W in qualitative agreement with measured nitrogen fixation rates. The new algorithm, however, fails to detect sub-surface booms evidenced in trichome counts. Improving Trichodesmium detection requires measuring ocean color at higher spectral and spatial (

Description: We assessed the influence of the marine diazotrophic cyanobacterium Trichodesmium on the bio-optical properties of South West tropical Pacific waters (18–22° S, 160° E–160° W) during the February–March 2015 OUTPACE cruise. We performed measurements of backscattering and absorption coefficients, irradiance, and radiance, in the euphotic zone, and took Underwater Vision Profiler 5 (UPV5) pictures for counting the largest Trichodesmium spp. colonies. Pigment concentrations were determined by fluorimetry and by high performance liquid chromatography and picoplankton abundance by flow cytometry. Trichome concentration was estimated from pigment algorithms and validated by surface visual counts. In result, the large colonies were well correlated to the trichome concentration estimates (though with a large factor of 600 to 900, due to aggregation processes). Large Trichodesmium abundance was always associated with particulate absorption at a peak of mycosporine-like amino acid absorption, and high particulate backscattering, but not with high fluorescence, high chlorophyll-a concentration, or blue particulate absorption in the water column. Along the West to East transect, Trichodesmium together with Prochlorococcus represented the major part of total chlorophyll and the other groups were negligible. Trichodesmium contribution to chlorophyll was the highest in the Melanesian Archipelago around New Caledonia and Vanuatu, progressively decreased to the vicinity of the Fiji Islands, and reached a minimum in the South Pacific gyre where the contribution of Prochlorococcus was maximum. At the frontal LDB, Trichodesmium and Prochlorococcus has almost same contributions. The relationship between normalized water-leaving radiance, in the ultraviolet and visible domains, nLw, and chlorophyll was generally similar to that found in the Eastern tropical at BIOSOPE. Principal component analysis (PCA) of OUTPACE data showed that nLw were strongly correlated to chlorophyll except in the green and yellow domains. These results, as well as differences in the PCA of BIOSOPE data, suggested that nLw variability in the green and yellow during OUTPACE was influenced by other variables, associated with Trichodesmium presence as the backscattering coefficient, phycoerythrin fluorescence, and/or zeaxanthin absorption. Trichodesmium detection should then involve examination of nLw at the green and yellow wavelengths.