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We apply an innovative approach to time series data of surface chlorophyll from satellite observations with SeaWiFS (Sea‐viewing Wide Field‐of‐view Sensor) to estimate the primary production associated with three major phytoplankton classes (micro‐, nano‐, and picophytoplankton) within the world's oceans. Statistical relationships, determined from an extensive in situ database of phytoplankton pigments, are used to infer class‐specific vertical profiles of chlorophyll a concentration from satellite‐derived surface chlorophyll a. This information is combined with a primary production model and class‐specific photophysiological parameters to compute global seasonal fields of class‐specific primary production over a 10‐year period from January 1998 through December 2007. Microphytoplankton (mostly diatoms) appear as a major contributor to total primary production in coastal upwelling systems (70%) and temperate and subpolar regions (50%) during the spring‐summer season. The contribution of picophytoplankton (e.g., prokaryotes) reaches maximum values (45%) in subtropical oligotrophic gyres. Nanophytoplankton (e.g., prymnesiophytes) provide a ubiquitous, substantial contribution (30–60%). Annual global estimates of class‐specific primary production amount to 15 Gt C yr−1 (32% of total), 20 Gt C yr−1 (44%) and 11 Gt C yr−1 (24%) for micro‐, nano‐, and picophytoplankton, respectively. The analysis of interannual variations revealed large anomalies in class‐specific primary production as compared to the 10‐year mean cycle in both the productive North Atlantic basin and the more stable equatorial Pacific upwelling. Microphytoplankton show the largest range of variability of the three phytoplankton classes on seasonal and interannual time scales. Our results contribute to an understanding and quantification of carbon cycle in the ocean.

We analyzed a large dataset of simultaneous measurements of phytoplankton pigments, spectral specific absorption coefficient for phytoplankton [a*(λ)], and photosynthesis versus irradiance (P versus E) curve parameters to examine the possible relationships between phytoplankton community structure and photophysiological properties at large spatial scales. Data were collected in various regions, mostly covering the trophic gradient encountered in the world's open ocean. The community composition is described in terms of biomass of three phytoplankton classes, determined using specific biomarker pigments. We present a general empirical model that describes the dependence of algal photophysiology on both the community composition and the relative irradiance within the water column (essentially reflecting photoacclimation). The application of the model to the in situ dataset enables the identification of vertical profiles of photophysiological properties for each phytoplankton class. The class-specific a*(λ) obtained are consistent with results from the literature and with previous models developed for small and large cells, both in terms of the absolute values and the vertical patterns. Similarly, for the class-specific P versus E curve parameters, the magnitude and vertical distribution obtained with this method are coherent with previous observations. Large cells (mainly diatoms) may be more efficient in carbon storage than smaller cells, whereas their yield of light absorption is lower. We anticipate that such photophysiological parameterizations can improve primary production models by providing estimates of primary production that are specific to different phytoplankton classes on large scale.

The deep chlorophyll maximum (DCM) is a ubiquitous feature of phytoplankton vertical distribution in stratified waters that is relevant to our understanding of the mechanisms that underpin the variability in photoautotroph ecophysiology across environmental gradients and has implications for remote sensing of aquatic productivity. During the PEACETIME (Process studies at the air-sea interface after dust deposition in the Mediterranean Sea) cruise, carried out from 10 May to 11 June 2017, we obtained 23 concurrent vertical profiles of phytoplankton chlorophyll a, carbon biomass and primary production, as well as heterotrophic prokaryotic production, in the western and central Mediterranean basins. Our main aims were to quantify the relative role of photoacclimation and enhanced growth as underlying mechanisms of the DCM and to assess the trophic coupling between phytoplankton and heterotrophic prokaryotic production. We found that the DCM coincided with a maximum in both the biomass and primary production but not in the growth rate of phytoplankton, which averaged 0.3 d−1 and was relatively constant across the euphotic layer. Photoacclimation explained most of the increased chlorophyll a at the DCM, as the ratio of carbon to chlorophyll a (C:Chl a) decreased from ca. 90–100 (g:g) at the surface to 20–30 at the base of the euphotic layer, while phytoplankton carbon biomass increased from ca. 6 mg C m−3 at the surface to 10–15 mg C m−3 at the DCM. As a result of photoacclimation, there was an uncoupling between chlorophyll a-specific and carbon-specific productivity across the euphotic layer. The ratio of fucoxanthin to total chlorophyll a increased markedly with depth, suggesting an increased contribution of diatoms at the DCM. The increased biomass and carbon fixation at the base of the euphotic zone was associated with enhanced rates of heterotrophic prokaryotic activity, which also showed a surface peak linked with warmer temperatures. Considering the phytoplankton biomass and turnover rates measured at the DCM, nutrient diffusive fluxes across the nutricline were able to supply only a minor fraction of the photoautotroph nitrogen and phosphorus requirements. Thus the deep maxima in biomass and primary production were not fuelled by new nutrients but likely resulted from cell sinking from the upper layers in combination with the high photosynthetic efficiency of a diatom-rich, low-light acclimated community largely sustained by regenerated nutrients. Further studies with increased temporal and spatial resolution will be required to ascertain if the peaks of deep primary production associated with the DCM persist across the western and central Mediterranean Sea throughout the stratification season.

One pathway by which the oceans influence climate is via the emission of sea spray that may subsequently influence cloud properties. Sea spray emissions are known to be dependent on atmospheric and oceanic physicochemical parameters, but the potential role of ocean biology on sea spray fluxes remains poorly characterized. Here we show a consistent significant relationship between seawater nanophytoplankton cell abundances and sea-spray derived Cloud Condensation Nuclei (CCN) number fluxes, generated using water from three different oceanic regions. This sensitivity of CCN number fluxes to ocean biology is currently unaccounted for in climate models yet our measurements indicate that it influences fluxes by more than one order of magnitude over the range of phytoplankton investigated.

The current paradigm holds that cyanobacteria, which evolved oxygenic photosynthesis more than 2 billion years ago, are still the major light harvesters driving primary productivity in open oceans. Here we show that tiny unicellular eukaryotes belonging to the photosynthetic lineage of the Haptophyta are dramatically diverse and ecologically dominant in the planktonic photic realm. The use of Haptophyta-specific primers and PCR conditions adapted for GC-rich genomes circumvented biases inherent in classical genetic approaches to exploring environmental eukaryotic biodiversity and led to the discovery of hundreds of unique haptophyte taxa in 5 clone libraries from subpolar and subtropical oceanic waters. Phylogenetic analyses suggest that this diversity emerged in Paleozoic oceans, thrived and diversified in the permanently oxygenated Mesozoic Panthalassa, and currently comprises thousands of ribotypic species, belonging primarily to low-abundance and ancient lineages of the \"rare biosphere.\" This extreme biodiversity coincides with the pervasive presence in the photic zone of the world ocean of 19'-hexanoyloxyfucoxanthin (19-Hex), an accessory photosynthetic pigment found exclusively in chloroplasts of haptophyte origin. Our new estimates of depth-integrated relative abundance of 19-Hex indicate that haptophytes dominate the chlorophyll a-normalized phytoplankton standing stock in modern oceans. Their ecologic and evolutionary success, arguably based on mixotrophy, may have significantly impacted the oceanic carbon pump. These results add to the growing evidence that the evolution of complex microbial eukaryotic cells is a critical force in the functioning of the biosphere.

Phytoplankton are composed of diverse taxonomical groups, which are manifested as distinct morphology, size and pigment composition. These characteristics, modulated by their physiological state, impact their light absorption and scattering, allowing them to be detected with ocean color satellite radiometry. There is a growing volume of literature describing satellite algorithms to retrieve information on phytoplankton composition in the ocean. This synthesis provides a review of current methods and a simplified comparison of approaches. The aim is to provide an easily comprehensible resource for non-algorithm developers, who desire to use these products, thereby raising the level of awareness and use of these products and reducing the boundary of expert knowledge needed to make a pragmatic selection of output products with confidence. The satellite input and output products, their associated validation metrics, as well as assumptions, strengths and limitations of the various algorithm types are described, providing a framework for algorithm organization to assist users and inspire new aspects of algorithm development capable of exploiting the higher spectral, spatial and temporal resolutions from the next generation of ocean color satellites.

Pigment-based phytoplankton community composition and primary production were investigated for the first time in the Red Sea in February-April 2015 to demonstrate how the strong south to north environmental gradients determine phytoplankton community structure in Red Sea offshore regions (along the central axis). Taxonomic pigments were used as size group markers of pico, nano- and microphytoplankton. Phytoplankton primary production rates associated with the three phytoplankton groups (pico-, nano- and microphytoplankton) were estimated using a bio-optical model. Pico- (Synechococcus and Prochlorococcus sp) and Nanophytoplankton (Prymnesiophytes and Pelagophytes) were the dominant size groups and contributed to 49% and 38%, respectively, of the phytoplankton biomass. Microphytoplankton (diatoms) contributed to 13% of the phytoplankton biomass within the productive layer (1.5 Zeu). Sub-basin and mesoscale structures (cyclonic eddy and mixing) were exceptions to this general trend. In the southern Red Sea, diatoms and picophytoplankton contributed to 27% and 31% of the phytoplankton biomass, respectively. This result induced higher primary production rates (430 ± 50 mgC m-2 d-1) in this region (opposed to CRS and NRS). The cyclonic eddy contained the highest microphytoplankton proportion (45% of TChla) and the lowest picophytoplankton contribution (17% of TChla) while adjacent areas were dominated by pico- and nanophytoplankton. We estimated that the cyclonic eddy is an area of enhanced primary production, which is up to twice those of the central part of the basin. During the mixing of the water column in the extreme north of the basin, we observed the highest TChla integrated (40 mg m-2) and total primary production rate (640 mgC m-2 d-1) associated with the highest nanophytoplankton contribution (57% of TChla). Microphytoplankton were a major contributor to total primary production (54%) in the cyclonic eddy. The contribution of picophytoplankton (Synechococcus and Prochlorococcus sp) reached maximum values (49%) in the central Red Sea. Nanophytoplankton seem to provide a ubiquitous substantial contribution (30-56%). Our results contribute to providing new insights on the spatial distribution and structure of phytoplankton groups. An understanding and quantification of the carbon cycle in the Red Sea was made based on estimates of primary production associated with pico-, nano- and microphytoplankton.

Monitoring the spatio-temporal variations of surface chlorophyll-a concentration (Chl, a proxy of phytoplankton biomass) greatly benefited from the availability of continuous and global ocean color satellite measurements from 1997 onward. These two decades of satellite observations are however still too short to provide a comprehensive description of Chl variations at decadal to multi-decadal timescales. This paper investigates the ability of a machine learning approach (a non-linear statistical approach based on Support Vector Regression, hereafter SVR) to reconstruct global spatio-temporal Chl variations from selected surface oceanic and atmospheric physical parameters. With a limited training period (13 years), we first demonstrate that Chl variability from a 32-years global physical-biogeochemical simulation can generally be skillfully reproduced with a SVR using the model surface variables as input parameters. We then apply the SVR to reconstruct satellite Chl observations using the physical predictors from the above numerical model and show that the Chl reconstructed by this SVR more accurately reproduces some aspects of observed Chl variability and trends compared to the model simulation. This SVR is able to reproduce the main modes of interannual Chl variations depicted by satellite observations in most regions, including El Niño signature in the tropical Pacific and Indian Oceans. In stark contrast with the trends simulated by the biogeochemical model, it also accurately captures spatial patterns of Chl trends estimated by satellite data, with a Chl increase in most extratropical regions and a Chl decrease in the center of the subtropical gyres, although the amplitude of these trends are underestimated by half. Results from our SVR reconstruction over the entire period (1979–2010) also suggest that the Interdecadal Pacific Oscillation drives a significant part of decadal Chl variations in both the tropical Pacific and Indian Oceans. Overall, this study demonstrates that non-linear statistical reconstructions can be complementary tools to in situ and satellite observations as well as conventional physical-biogeochemical numerical simulations to reconstruct and investigate Chl decadal variability.

In-situ fluorescence is a widely used method to estimate the chlorophyll-a (Chla) concentration, a proxy of the phytoplankton biomass. With the emergence of autonomous platforms such as BioGeoChemical-Argo (BGC-Argo) profiling floats, its use has expanded to global scale observations. However, the relationship between in-situ fluorescence and Chla may vary significantly, leading to major discrepancies between oceanic regions. This study aims to investigate the main sources of the natural variability in the in-situ fluorescence signal in the global open ocean, specifically the influence of the phytoplankton community composition. In this view, we analyzed a combination of three datasets comprising concomitant measurements of in-situ fluorescence, pigment concentrations and phytoplankton absorption spectra. Two datasets cover several contrasted bioregions of the global ocean whereas the third one consists of a regional time series in the northwestern Mediterranean Sea, which allows to examine the effect of phytoplankton community composition on the fluorescence signal on the global, seasonal and vertical scales. We studied the variability of the two major drivers of the natural variability of the fluorescence process, i.e. the light absorption and the fluorescence quantum yield of phytoplankton, in regards of the variability of the pigment composition of the communities. The community composition correlates substantially with the Chla-to-fluorescence ratio, with high fluorescence values associated with phytoplankton communities dominated by large cells. This trend may be explained by the combined effects of the community composition on the phytoplankton absorption coefficient and the fluorescence quantum yield, and is consistently observed globally, seasonally and vertically. Non-photosynthetic pigments also appear to play a critical role in oligotrophic surface waters, leading to a reduction of the quantum yield of fluorescence. The results indicate that the phytoplankton community composition plays a key role in the relationship between the in-situ fluorescence signal and Chla concentration. Therefore, we suggest that taking into account the composition of phytoplankton communities in the retrieval of the Chla concentration from current in-situ fluorometers, those mounted on BGC-Argo floats in particular, would lead to a better estimation of the phytoplankton biomass on a wide range of spatial and temporal scales.

Since 2012, an array of 105 Biogeochemical-Argo (BGC-Argo) floats has been deployed across the world's oceans to assist in filling observational gaps that are required for characterizing open-ocean environments. Profiles of biogeochemical (chlorophyll and dissolved organic matter) and optical (single-wavelength particulate optical backscattering, downward irradiance at three wavelengths, and photosynthetically available radiation) variables are collected in the upper 1000 m every 1 to 10 days. The database of 9837 vertical profiles collected up to January 2016 is presented and its spatial and temporal coverage is discussed. Each variable is quality controlled with specifically developed procedures and its time series is quality-assessed to identify issues related to biofouling and/or instrument drift. A second database of 5748 profile-derived products within the first optical depth (i.e., the layer of interest for satellite remote sensing) is also presented and its spatiotemporal distribution discussed. This database, devoted to field and remote ocean color applications, includes diffuse attenuation coefficients for downward irradiance at three narrow wavebands and one broad waveband (photosynthetically available radiation), calibrated chlorophyll and fluorescent dissolved organic matter concentrations, and single-wavelength particulate optical backscattering. To demonstrate the applicability of these databases, data within the first optical depth are compared with previously established bio-optical models and used to validate remotely derived bio-optical products. The quality-controlled databases are publicly available from the SEANOE (SEA scieNtific Open data Edition) publisher at https://doi.org/10.17882/49388 and https://doi.org/10.17882/47142 for vertical profiles and products within the first optical depth, respectively.