Search

International audience; This study assesses marine community production based on the diel variability of bio-optical properties monitored by two BioGeoChemical-Argo (BGC-Argo) floats. Experiments were conducted in two distinct Mediterranean systems, the northwestern Ligurian Sea and the central Ionian Sea, during summer months. We derived particulate organic carbon (POC) stock and gross community production integrated within the surface, euphotic and subsurface chlorophyll maximum (SCM) layers, using an existing approach applied to diel cycle measurements of the particulate beam attenuation (cp) and backscattering (bbp) coefficients. The diel cycle of cp provided a robust proxy for quantifying biological production in both systems; that of bbp was comparatively less robust. Derived primary production estimates vary by a factor of 2 depending upon the choice of the bio-optical relationship that converts the measured optical coefficient to POC, which is thus a critical step to constrain. Our results indicate a substantial contribution to the water column production of the SCM layer (16 %-42 %), which varies largely with the considered system. In the Ligurian Sea, the SCM is a seasonal feature that behaves as a subsurface biomass maximum (SBM) with the ability to respond to episodic abiotic forcing by increasing production. In contrast, in the Ionian Sea, the SCM is permanent, primarily induced by phytoplankton photoacclimation, and contributes moderately to water column production. These results clearly demonstrate the strong potential for transmissometers deployed on BGC-Argo profiling floats to quantify non-intrusively in situ biological production of organic carbon in the water column of stratified oligotrophic systems with recurring or permanent SCMs, which are widespread features in the global ocean.

The shallower oxygen-poor water masses of the ocean confine a majority of the microbial communities that can produce up to 90 % of oceanic N2. This effective N2-yielding section encloses a suspended small-particle layer, inferred from particle backscattering (bbp) measurements. It is thus hypothesized that this layer (hereafter, the bbp-layer) is linked to microbial communities involved in N2 yielding such as nitrate-reducing SAR11 as well as sulfur-oxidizing, anammox, and denitrifying bacteria – a hypothesis yet to be evaluated. Here, data collected by three BGC-Argo floats deployed in the Black Sea are used to investigate the origin of this bbp-layer. To this end, we evaluate how the key drivers of N2-yielding bacteria dynamics impact the vertical distribution of bbp and the thickness of the bbp-layer. In conjunction with published data on N2 excess, our results suggest that the bbp-layer is at least partially composed of the bacteria driving N2 yielding for three main reasons: (1) strong correlations are recorded between bbp and nitrate; (2) the top location of the bbp-layer is driven by the ventilation of oxygen-rich subsurface waters, while its thickness is modulated by the amount of nitrate available to produce N2; and (3) the maxima of both bbp and N2 excess coincide at the same isopycnals where bacteria involved in N2 yielding coexist. We thus advance that bbp and O2 can be exploited as a combined proxy to delineate the N2-yielding section of the Black Sea. This proxy can potentially contribute to refining delineation of the effective N2-yielding section of oxygen-deficient zones via data from the growing BGC-Argo float network.

As commonly observed in oligotrophic stratified waters, a subsurface (or deep) chlorophyll maximum (SCM) frequently characterizes the vertical distribution of phytoplankton chlorophyll in the Mediterranean Sea. Occurring far from the surface layer “seen” by ocean colour satellites, SCMs are difficult to observe with adequate spatio-temporal resolution and their biogeochemical impact remains unknown. Biogeochemical-Argo (BGC-Argo) profiling floats represent appropriate tools for studying the dynamics of SCMs. Based on data collected from 36 BGC-Argo floats deployed in the Mediterranean Sea, our study aims to address two main questions. (1) What are the different types of SCMs in the Mediterranean Sea? (2) Which environmental factors control their occurrence and dynamics? First, we analysed the seasonal and regional variations in the chlorophyll concentration (Chl a), particulate backscattering coefficient (bbp), a proxy of the particulate organic carbon (POC) and environmental parameters (photosynthetically active radiation and nitrates) within the SCM layer over the Mediterranean Basin. The vertical profiles of Chl a and bbp were then statistically classified and the seasonal occurrence of each of the different types of SCMs quantified. Finally, a case study was performed on two contrasted regions and the environmental conditions at depth were further investigated to understand the main controls on the SCMs. In the eastern basin, SCMs result, at a first order, from a photoacclimation process. Conversely, SCMs in the western basin reflect a biomass increase at depth benefiting from both light and nitrate resources. Our results also suggest that a variety of intermediate types of SCMs are encountered between these two endmember situations.

The distribution of the chlorophyll a concentration ([Chl a]) in the Mediterranean Sea, mainly obtained from satellite surface observations or from scattered in situ experiments, is updated by analyzing a database of fluorescence profiles converted into [Chl a]. The database, which includes 6790 fluorescence profiles from various origins, was processed with a specific quality control procedure. To ensure homogeneity between the different data sources, 65 % of fluorescence profiles have been intercalibrated on the basis of their concomitant satellite [Chl a] estimation. The climatological pattern of [Chl a] vertical profiles in four key sites of the Mediterranean Sea has been analyzed. Climatological results confirm previous findings over the range of existing [Chl a] values and throughout the principal Mediterranean trophic regimes. They also provide new insights into the seasonal variability in the shape of the vertical [Chl a] profile, inaccessible through remote-sensing observations. An analysis based on the recognition of the general shape of the fluorescence profile was also performed. Although the shape of [Chl a] vertical distribution characterized by a deep chlorophyll maximum (DCM) is ubiquitous during summer, different forms are observed during winter, thus suggesting that factors affecting the vertical distribution of the biomass are complex and highly variable. The [Chl a] spatial distribution in the Mediterranean Sea mimics, on smaller scales, what is observed in the global ocean. As already evidenced by analyzing satellite surface observations, midlatitude- and subtropical-like phytoplankton dynamics coexist in the Mediterranean Sea. Moreover, the Mediterranean DCM variability appears to be characterized by patterns already observed on the global scale.

Understanding the ocean carbon cycle requires a precise assessment of phytoplankton biomass in the oceans. In terms of numbers of observations, satellite data represent the largest available data set. However, as they are limited to surface waters, they have to be merged with in situ observations. Amongst the in situ data, fluorescence profiles constitute the greatest data set available, because fluorometers have operated routinely on oceanographic cruises since the 1970s. Nevertheless, fluorescence is only a proxy of the total chlorophyll a concentration and a data calibration is required. Calibration issues are, however, sources of uncertainty, and they have prevented a systematic and wide range exploitation of the fluorescence data set. In particular, very few attempts to standardize the fluorescence databases have been made. Consequently, merged estimations with other data sources (e.g. satellite) are lacking. We propose a merging method to fill this gap. It consists firstly in adjusting the fluorescence profile to impose a zero chlorophyll a concentration at depth. Secondly, each point of the fluorescence profile is then multiplied by a correction coefficient, which forces the chlorophyll a integrated content measured on the fluorescence profile to be consistent with the concomitant ocean colour observation. The method is close to the approach proposed by Boss et al. (2008) to correct fluorescence data of a profiling float, although important differences do exist. To develop and test our approach, in situ data from three open ocean stations (BATS, HOT and DYFAMED) were used. Comparison of the so-called "satellite-corrected" fluorescence profiles with concomitant bottle-derived estimations of chlorophyll a concentration was performed to evaluate the final error (estimated at 31%). Comparison with the Boss et al. (2008) method, using a subset of the DYFAMED data set, demonstrated that the methods have similar accuracy. The method was applied to two different data sets to demonstrate its utility. Using fluorescence profiles at BATS, we show that the integration of "satellite-corrected" fluorescence profiles in chlorophyll a climatologies could improve both the statistical relevance of chlorophyll a averages and the vertical structure of the chlorophyll a field. We also show that our method could be efficiently used to process, within near-real time, profiles obtained by a fluorometer deployed on autonomous platforms, in our case a bio-optical profiling float. The application of the proposed method should provide a first step towards the generation of a merged satellite/fluorescence chlorophyll a product, as the "satellite-corrected" profiles should then be consistent with satellite observations. Improved climatologies with more consistent satellite and in situ data are likely to enhance the performance of present biogeochemical models.

In vivo fluorescence of Chlorophyll-a (Chl-a) is a potentially useful property to study the vertical distribution of phytoplankton biomass. However the technique is presently not fully exploited as it should be, essentially because of the difficulties in converting the fluorescence signal into an accurate Chl-a concentration. These difficulties arise noticeably from natural variations in the Chl-a fluorescence relationship, which is under the control of community composition as well as of their nutrient and light status. As a consequence, although vertical profiles of fluorescence are likely the most recorded biological property in the open ocean, the corresponding large databases are underexploited. Here with the aim to convert a fluorescence profile into a Chl-a concentration profile, we test the hypothesis that the Chl-a concentration can be gathered from the sole knowledge of the shape of the fluorescence profile. We analyze a large dataset from 18 oceanographic cruises conducted in case-1 waters from the highly stratified hyperoligotrophic waters (surface Chl-a = 0.02 mg m−3) of the South Pacific Gyre to the eutrophic waters of the Benguela upwelling (surface Chl-a = 32 mg m−3) and including the very deep mixed waters in the North Atlantic (Mixed Layer Depth = 690 m). This dataset encompasses more than 700 vertical profiles of Chl-a fluorescence as well as accurate estimations of Chl-a by High Performance Liquid Chromatography (HPLC). Two typical fluorescence profiles are identified, the uniform profile, characterized by a homogeneous layer roughly corresponding to the mixed layer, and the non-uniform profile, characterized by the presence of a Deep Chlorophyll Maximum. Using appropriate mathematical parameterizations, a fluorescence profile is subsequently represented by 3 or 5 shape parameters for uniform or non-uniform profiles, respectively. For both situations, an empirical model is developed to predict the "true" Chl-a concentration from these shape parameters. This model is then used to calibrate a fluorescence profile in Chl-a units. The validation of the approach provides satisfactory results with a median absolute percent deviation of 33 % when comparing the HPLC Chl-a profiles to the Chl-a-calibrated fluorescence. The proposed approach thus opens the possibility to produce Chl-a climatologies from uncalibrated fluorescence profile databases that have been acquired in the past and to which numerous new profiles will be added, thanks to the recent availability of autonomous platforms (profiling floats, gliders and animals) instrumented with miniature fluorometers.

The cores of the subtropical anticyclonic gyres are characterized by their oligotrophic status and minimal chlorophyll concentration, compared to that of the whole ocean. These zones are unambiguously detected by space borne ocean color sensors thanks to their typical spectral reflectance, which is that of extremely clear and deep blue waters. Not only the low chlorophyll (denoted [Chl]) level, but also a reduced amount of colored dissolved organic matter (CDOM or "yellow substance") account for this clarity. The oligotrophic waters of the North and South Pacific gyres, the North and South Atlantic gyres, and the South Indian gyre have been comparatively studied with respect to both [Chl] and CDOM contents, by using 10-year data (1998–2007) of the Sea-viewing Wide field-of-view Sensor (SeaWiFS, NASA). Albeit similar these oligotrophic zones are not identical regarding their [Chl] and CDOM contents, as well as their seasonal cycles. According to the zone, the averaged [Chl] value varies from 0.026 to 0.059 mg m−3, whereas the ay(443) average (the absorption coefficient due to CDOM at 443 nm) is between 0.0033 and 0.0072 m−1. The CDOM-to-[Chl] relative proportions also differ between the zones. The clearest waters, corresponding to the lowest [Chl] and CDOM concentrations, are found near Easter Island and near Mariana Islands in the western part of the North Pacific Ocean. In spite of its low [Chl], the Sargasso Sea presents the highest CDOM content amongst the six zones studied. Except in the North Pacific gyre (near Mariana and south of Hawaii islands), a conspicuous seasonality appears to be the rule in the other 4 gyres and affects both [Chl] and CDOM; both quantities vary in a ratio of about 2 (maximum-to-minimum). Coinciding [Chl] and CDOM peaks occur just after the local winter solstice, which is also the period of the maximal mixed layer depth in these latitudes. It is hypothesized that the vertical transport of unbleached CDOM from the subthermocline layers is the main process enhancing the CDOM concentration within the upper layer in winter. In summer, the CDOM experiences its minimum which is delayed with respect to the [Chl] minimum; apparently, the solar photo-bleaching of CDOM is a slower process than the post-bloom algal Chl decay. Where they exist, the seasonal cycles are repeated without notable change from year to year. Long term (10 y) trends have not been detected in these zones. These oligotrophic gyres can conveniently be used for in-flight calibration and comparison of ocean color sensors, provided that their marked seasonal variations are accounted for.

BIOSOPE cruise covered an oceanographic transect through the centre of the South Pacific Gyre (SPG) from the Marquesas archipelago to the Peru-Chile upwelling (PCU). Water samples from 6 depths in the euphotic zone were collected at 20 stations. The concentrations of suspended calcite particles, coccolithophores cells and detached coccoliths were estimated together with size and weight using an automatic polarizing microscope, a digital camera, and a collection of softwares performing morphometry and pattern recognition. Some of these softwares are new and described here for the first time. The coccolithophores standing stocks were usually low and reached maxima west of the PCU. The coccoliths of Emiliania huxleyi, Gephyrocapsa spp. and Crenalithus spp. (Order Isochrysidales) represented more than 30% of all the suspended calcite particles detected in the size range 0.1–46 μm (22% of PIC in term of calcite weight). These species grew preferentially in the Chlorophyll maximum zone. In the SPG their maximum cell concentrations were recorded between depth of 150 and 200 m, which is unusually deep for these taxa. The weight of coccoliths and coccospheres were correlated to their size. Large and heavy coccoliths and coccospheres were found in regions with relatively high fertility in the Marquises Island and in the PCU. Small and light coccoliths and coccospheres were found west of the PCU. This distribution is strongly related to ocean chemistry in particular to alkalinity and to carbonate ions concentration. The biotic (coccolithophores production) influence on calcification is mainly driven at the local scale (depth) whereas the abiotic (carbonate chemistry) plays its most important role at the regional (horizontal) level. Here 94% of the variability of coccolith and coccosphere weight can be explained by a change in 7 environmental variables.

The distribution of lipid biomarkers and their stable carbon isotope composition was investigated on suspended particles from different contrasting trophic environments at six sites in the South East Pacific. High algal biomass with diatom-related lipids (24-methylcholesta-5,24(28)-dien-3β-ol, C25 HBI alkenes, C16:4 FA, C20:5 FA) was characteristic in the upwelling zone, whereas haptophyte lipids (long-chain (C37-C39) unsaturated ketones) were proportionally most abundant in the nutrient-poor settings of the centre of the South Pacific Gyre and on its easter edge. The dinoflagellate–sterol, 4α-23,24-trimethylcholest-22(E)-en-3β-ol, was a minor contributor in all of the studied area and the cyanobacteria-hydrocarbon, C17n-alkane, was at maximum in the high nutrient low chlorophyll regime of the subequatorial waters near the Marquesas archipelago. The taxonomic and spatial variability of the relationships between carbon photosynthetic fractionation and environmental conditions for four specific algal taxa (diatoms, haptophytes, dinoflagellates and cyanobacteria) was also investigated. The carbon isotope fractionation factor (εp) of the 24-methylcholesta-5,24(28)-dien-3β-ol diatom marker, varied over a range of 16% along the different trophic systems. In contrast, εp of dinoflagellate, cyanobacteria and alkenone markers varied only by 7–10‰. The low fractionation factors and small variations between the different phytoplankton markers measured in the upwelling area likely reveals uniformly high specific growth rates within the four phytoplankton taxa, and/or that transport of inorganic carbon into phytoplankton cells may not only occur by diffusion but also by other carbon concentrating mechanisms (CCM). In contrast, in the oligotrophic zone, i.e. gyre and eastgyre, relatively high εp values, especially for the diatom marker, indicate diffusive CO2 uptake by the eukaryotic phytoplankton. At these nutrient-poor sites, the lower εp values for haptophytes, dinoflagellates and cyanobacteria indicate higher growth rates or major differences on the carbon uptake mechanisms compared to diatoms.

Due to the low atmospheric input of phosphate into the open ocean, it is one of the key nutrients that could ultimately control primary production and carbon export into the deep ocean. The observed trend over the last 20 years has shown a decrease in the dissolved inorganic phosphate (DIP) pool in the North Pacific gyre, which has been correlated to the increase in di-nitrogen (N2) fixation rates. Following a NW-SE transect, in the Southeast Pacific during the early austral summer (BIOSOPE cruise), we present data on DIP, dissolved organic phosphate (DOP) and particulate phosphate (PP) pools along with DIP turnover times (TDIP) and N2 fixation rates. We observed a decrease in DIP concentration from the edges to the centre of the gyre. Nevertheless the DIP concentrations remained above 100 nmol L−1 and T DIP was more than 6 months in the centre of the gyre; DIP availability remained largely above the level required for phosphate limitation to occur and the absence of Trichodesmium spp and low nitrogen fixation rates were likely to be controlled by other factors such as temperature or iron availability. This contrasts with recent observations in the North Pacific Ocean at the ALOHA station and in the western Pacific Ocean at the same latitude (DIAPALIS cruises) where lower DIP concentrations (−1) and T DIP 2 fixation rates and possibly carbon dioxide sequestration, if the primary ecophysiological controls, temperature and/or iron availability, were alleviated.

The optical properties of Case 1 waters have been empirically related to the chlorophyll concentration, [Chl], historically used as an index of the trophic state and of the abundance of the biological materials. The well-known natural variability around the mean statistical relationships is here examined by comparing the apparent optical properties (spectral downward irradiance attenuation and reflectance) as a function of [Chl] in two Case 1 environments, the Pacific and Mediterranean waters. These oceanic zones apparently represent two extremes of the possible bio-optical variability range around the mean. The systematic deviations, in both directions with respect to the average laws, mainly result from the differing contents in non-algal detrital materials and dissolved colored substance for a given [Chl] level. These contents are higher than the average in the Mediterranean Sea, and lower in the Pacific Ocean, respectively. These divergences between the two water bodies, detectable in the visible spectral domain, are considerably accentuated in the UV domain. The bio-optical properties in this spectral domain (310–400 nm) are systematically explored. They are more varying for a given [Chl] than those in the visible domain. Their prediction based on the sole [Chl] index is thus problematic, although it is probably possible on a regional scale if reliable field data are available. It does not seem, however, that ubiquitous relationships exist for this spectral domain for all Case 1 waters at global scale.