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Heterotrophic prokaryotes (HP) contribute largely to dissolved organic matter (DOM) processing in the ocean, but they also release diverse organic substances. The bioavailability of DOM released by HP under varying environmental conditions are not fully elucidated. In this study, we investigated the effect of P-limitation on the bioavailability of DOM released by a single bacterial strain and 2 natural HP communities. These HP were grown under P-replete and P-limited conditions, and their DOM (HP-DOM) was used as a substrate for natural prokaryotic communities from a coastal site in the Northwestern Mediterranean Sea. We followed changes in prokaryotic growth, enzymatic activity, diversity and community composition together with the consumption of HP-DOM fluorescence (FDOM). HP-DOM produced under P-replete and P-limited conditions promoted both significant and similar prokaryotic growth. The humic-like fluorescence, commonly considered recalcitrant, was consumed during the incubations when this peak was initially dominating the FDOM pool, and this consumption coincided with higher alkaline phosphatase activity. HP-DOM supported growth of diverse prokaryotic communities and P-driven differences in HP-DOM quality selected for different indicator taxa in the degrading communities. Taken together, our findings emphasize that HP-DOM lability is dependent on both the DOM quality, shaped by P availability, and the composition of the consumers community.

We measured phytoplankton primary production and heterotrophic bacterial activities on microplastics and seawater in the Northwestern Mediterranean Sea during two 3-month spring periods over 2 consecutive years. Microorganisms growing on a 5 mm diameter low density polyethylene films (LDPE; 200 μm thick) faced two contrasting conditions depending on the year. Spring 2018 was characterized by consistent nutrient inputs and bloom development. In spring 2019, nutrient inputs and bloom were low. For the first time, we observed a clear coupling between primary production and heterotrophic prokaryote production on microplastics during both years, but with different intensity between years that reflected the crucial role of the trophic environmental conditions (nutrient supply) in shaping microbial activities on plastics. We found that high primary production on plastics could support the whole (net autotrophy) or the majority of the bacterial carbon demand needed for heterotrophic activities, supplemented by other carbon sources if surrounding waters are highly productive. We propose that microbial activity on plastics influences the microbial community in the surrounding seawater, especially when the environmental conditions are less favorable. An illustrative image of the role of plastics in the environment could be that of an inverter in an electrical circuit that mitigates both positive and negative variations. Our results highlight the potential role of the plastisphere in shaping biogeochemical cycles in the context of increasing amounts of plastic particles in the marine environment.

Marine heterotrophic prokaryotes (HP) play a key role in organic matter processing in the ocean; however, the view of HP as dissolved organic matter (DOM) sources remains underexplored. In this study, we quantified and optically characterized the DOM produced by two single marine bacterial strains. We then tested the availability of these DOM sources to in situ Mediterranean Sea HP communities. Two bacterial strains were used: Photobacterium angustum (a copiotrophic gammaproteobacterium) and Sphingopyxis alaskensis (an oligotrophic alphaproteobacterium). When cultivated on glucose as the sole carbon source, the two strains released from 7% to 23% of initial glucose as bacterial derived DOM (B-DOM), the quality of which (as enrichment in humic or protein-like substances) differed between strains. B-DOM induced significant growth and carbon consumption of natural HP communities, suggesting that it was partly labile. However, B-DOM consistently promoted lower prokaryotic growth efficiencies than in situ DOM. In addition, B-DOM changed HP exoenzymatic activities, enhancing aminopeptidase activity when degrading P. angustum DOM, and alkaline phosphatase activity when using S. alaskensis DOM, and promoted differences in HP diversity and composition. DOM produced by HP affects in situ prokaryotic metabolism and diversity, thus changing the pathways for DOM cycling (e.g. respiration over biomass production) in the ocean.

This study is part of the POLARCSIC platform activities.-- 14 pages, 8 figures, 2 tables, supplementary data https://doi.org/10.1016/j.scitotenv.2022.156921, Phytoplankton-derived organic matter sustains heterotrophic marine life in regions away from terrestrial inputs such as the Southern Ocean. Fluorescence spectroscopy has long been used to characterize the fluorescent organic matter (FOM) pool. However, most studies focus only in the dissolved FOM fraction (FDOM) disregarding the contribution of particles. In order to assess the dynamics and drivers of the dissolved and particulate fractions of FOM, we used a Lagrangian approach to follow the time evolution of phytoplankton proliferations at four different sites in the Southern Ocean and compared the FOM in filtered and unfiltered seawater aliquots. We found that filtration had little effects on FOM visible spectrum fluorescence intensities, implying that most of this signal was due to dissolved fluorophores. On the other hand, protein-like fluorescence was strongly supressed by filtration, with fluorescence of particles accounting for up to 90 % of the total protein-like FOM. Photobleaching was identified as the main driver of visible FDOM composition, which was better described by indices of phytoplankton photoacclimation than by measurements of the incident solar radiation dose. In contrast, protein-like FOM intensity and fractionation were primarily related to abundance, composition and physiological state of phytoplankton proliferations. The chlorophyll a concentration from non-diatom phytoplankton explained 91 % of the particulate protein-like FOM variability. The proportion of protein-like fluorescence found in the dissolved phase was predicted by the combination of potential viral and grazing pressures, which accounted for 51 and 29 % of its variability, respectively. Our results show that comparing FOM measurements from filtered and unfiltered seawater provides relevant information on the taxonomic composition and cell integrity of phytoplankton communities. A better understanding of the commonly overlooked FOM fractionation process is essential for the implementation of in situ fluorescence sensors and will also help us better understand the processes that govern OM cycling in marine systems, The study was funded by the Spanish Government through projects PEGASO (CTM2012-37615), BIOGAPS (CTM2016-81008-R) and DOGMA (PID2020-112653GB-I00). MC-B was supported by a predoctoral fellowship from the Spanish Government (FPU16/01925). [...] We also acknowledge the institutional support of the “Severo Ochoa Centre of Excellence” accreditation (CEX2019-000928-S) from the Spanish government

13 pages, 7 figures, 2 tables, supplementary data https://doi.org/10.1016/j.scitotenv.2019.06.524, Transparent exopolymer particles (TEP) are an abundant class of suspended organic particles, mainly formed by polysaccharides, which play important roles in biogeochemical and ecological processes in the ocean. In this study we investigated horizontal and vertical TEP distributions (within the euphotic layer, including the upper surface) and their short-term variability along with a suite of environmental and biological variables in four distinct regions of the Southern Ocean. TEP concentrations in the surface (4 m) averaged 102.3 ± 40.4 μg XG eq. L and typically decreased with depth. Chlorophyll a (Chl a) concentration was a better predictor of TEP variability across the horizontal (R = 0.66, p < 0.001) and vertical (R = 0.74, p < 0.001) scales than prokaryotic heterotrophic abundance and production. Incubation experiments further confirmed the main role of phytoplankton as TEP producers. The highest surface TEP concentrations were found north of the South Orkney Islands (144.4 ± 21.7 μg XG eq. L), where the phytoplankton was dominated by cryptophytes and haptophytes; however, the highest TEP:Chl a ratios were found south of these islands (153.4 ± 29.8 μg XG eq (μg Chl a), compared to a mean of 79.3 ± 54.9 μg XG eq (μg Chl a) in the whole cruise, in association with haptophyte dominance, proximity of sea ice and high exposure to solar radiation. TEP were generally enriched in the upper surface (10 cm) respect to 4 m, despite a lack of biomass enrichment, suggesting either upward transport by positive buoyancy or bubble scavenging, or higher production at the upper surface by light stress or aggregation. TEP concentrations did not present any significant cyclic diel pattern. Altogether, our results suggest that photobiological stress, sea ice melt and turbulence add to phytoplankton productivity in driving TEP distribution across the Antarctic Peninsula area and Atlantic sector of the Southern Ocean, This research was supported by the Spanish Ministry of Economy and Competitiveness through projects PEGASO (CTM2012-37615) and BIOGAPS (CTM2016-81008-R) to RS. MZ was supported by a FPU predoctoral fellowship (FPU13/04630) from the Spanish Ministry of Education and Culture

19 pages, 6 figures, 2 tables, supporting Information https://doi.org/10.1002/lno.11944, he Mediterranean Sea is a miniature ocean divided by the Sicily Strait into two basins with a marked west to east trophic gradient and separated of the nearby eastern Atlantic Ocean by the Strait of Gibraltar. Here, we test the hypothesis that these physical and environmental barriers favor the development of specific prokaryotic assemblages, leading to changes in community structure both in the vertical and horizontal spatial scales. By analyzing taxonomic and phylogenetic diversity using amplicon sequence variants (ASVs) of the 16S rRNA gene, we show that there is indeed marked vertical segregation of prokaryotic groups, similar to that found in other areas of the ocean, but also a clear horizontal structuring among the two Mediterranean basins and the adjacent Atlantic waters. Prokaryotic diversity increased with depth and toward the Atlantic, whereas the easternmost stations displayed more phylogenetically diverse phylotypes, despite harboring globally less diverse communities. Basin-indicator taxa (ASVs) accounted for a large fraction of the community (between 50% and 80%) in each of the basins at the surface and bathypelagic layers, being associated with different environmental variables. The existence of biogeographic and environmental barriers in the Mediterranean Sea is likely related to the trophic gradient at the surface and the isolation of water bodies in depth due to the Gibraltar and Sicily straits. Our work highlights the importance of studying microbial regional biogeography and provides the basis for future studies on the impact of this regionalization in the function of Mediterranean Sea prokaryotic communities, Sampling was funded by grant HOTMIX (CTM2011-30010/MAR), and processing also by grants REMEI (CTM2015-70340-R) and MIAU (RTI2018-101025-B-I00) from the Spanish Ministry of Science and Innovation, cofunded with FEDER funds. It has also received support from the Spanish government through the “Severo Ochoa Centre of Excellence” accreditation (CEX2019-000928-S) and the Catalan government through excellence group grant 2017SGR/1568. Laura Gómez-Consarnau was funded by grant OCE1924464 from the United States National Science Foundation (NSF)

Transparent exopolymer particles (TEP) and Coomassie stainable particles (CSP) are gel-like particles, ubiquitous in the ocean, that affect important biogeochemical processes including organic carbon cycling by planktonic food webs. Despite much research on both groups of particles (especially TEP) over many years, whether they exist as distinctly stainable fractions of the same particles or as independent particles, each with different driving factors, remains unclear. To address this question, we examined the temporal dynamics of TEP and CSP over 2 complete seasonal cycles at 2 coastal sites in the Northwestern Mediterranean Sea, the Blanes Bay Microbial Observatory (BBMO) and the L’Estartit Oceanographic Station (EOS), as well as their spatial distribution along a coast-to-offshore transect. Biological, chemical, and physical variables were measured in parallel. Surface concentrations (mean ± standard deviation [SD]) of TEP were 36.7 ± 21.5 µg Xanthan Gum (XG) eq L–1 at BBMO and 36.6 ± 28.3 µg XG eq L–1 at EOS; for CSP, they were 11.9 ± 6.1 µg BSA eq L–1 at BBMO and 13.0 ± 5.9 µg BSA eq L–1 at EOS. Seasonal variability was more evident at EOS, where surface TEP and CSP concentrations peaked in summer and spring, respectively, and less predictable at the shore-most station, BBMO. Vertical distributions between surface and 80 m, monitored at EOS, showed highest TEP concentrations within the surface mixed layer during the stratification period, whereas CSP concentrations were highest before the onset of summer stratification. Phytoplankton were the main drivers of TEP and CSP distributions, although nutrient limitation and saturating irradiance also appeared to play important roles. The dynamics and distribution of TEP and CSP were uncoupled both in the coastal sites and along the transect, suggesting that they are different types of particles produced and consumed differently in response to environmental variability.

15 pages, 8 figures, 2 tables, supplementary data https://doi.org/10.1016/j.dsr.2020.103460, The Variability of chlorophyll-specific phytoplankton light absorption [a*ph (λ)] was examined over depth and time in stratified offshore waters of the North-Western Mediterranean Sea. Coherent water patches were tracked with Lagrangian drifters during two oceanographic cruises in September (late summer) and May (post-spring bloom phase). By simultaneously analysing the phytoplankton absorption and pigment measurements, we explicitly separated the impact of pigment composition from that of pigment packaging on a*ph (λ). We further partitioned the packaging effect by comparing the variation of the packaging index [Qa*(440)], the phytoplankton community size structure (derived from diagnostic pigment analysis), and the chlorophyll-specific beam attenuation by particles [cp* (660)] as an optical index of phytoplankton photophysiology. In the ensemble of cruises, around 50% of the a*ph (440) vertical variation was explained by changes in the pigment composition (ruled by the decrease of photoprotective pigments with depth). The remaining vertical and inter-cruise variation of a*ph (440) was attributed to the packaging effect. We found that differences in the cp*(660) index (most likely indicating changes in the intracellular pigment concentration due to photoacclimation) mainly explained the observed variation in the packaging effect. Differences in cp* (660) were coincident with either the vertical gradient of light availability or the lower mean daily PAR irradiance in the euphotic layer of the September cruise. These explained the stronger packaging and lower a*ph (440) values observed with increasing depth in both cruises, and in September relative to May. On the other hand, differences in the phytoplankton community size structure did not explain the observed patterns in the packing effect. Our results highlight the importance of phytoplankton short-term acclimation to the prevailing light conditions, determining the vertical and temporal variability of a*ph (λ). A better understanding of the a*ph (λ) variability and its main drivers are key to improve different bio-optical applications, This research was funded by the successive Spanish Ministries of Science through projects SUMMER “SUrface Mixing Modulation of the Exposure to solar Radiation” (CTM 2008– 03309/MAR), BIOGAPS (CTM 2016-81008-R) and DOREMI (CTM 2012– 34294), and through the CONICET and CONICET-CSIC scholarships to G. L. Pérez. This is a contribution of the Research Group on Marine Biogeochemistry and Global Change and that of Microbial Diversity and Function in Aquatic Environments, funded by the Catalan Government. With the funding support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S), of the Spanish Research Agency (AEI)

9 pages, 7 figures, 3 tables, Coastal marine ecosystems are strongly influenced by different occasional events, such as intense winds, mixing, rain and river discharges. These events can directly or indirectly cause changes in dissolved organic matter (DOM) quality through a cascade of different biotic and abiotic processes. Changes in DOM quality are often associated with changes in DOM optical properties. Thus, examining the dynamics of chromophoric DOM (CDOM) can provide valuable information about biological and physical processes that have occurred in the ecosystem. Episodic meteorological events, particularly in temperate areas, appear very abruptly and induce very rapid responses; therefore, high time-resolved measurements are needed to capture them. We used a weekly sampling scheme to characterize DOM and nutrient dynamics in the NW Mediterranean coastal station ‘SOLA’. From February 2013 to April 2014, we measured several physical and chemical variables including temperature, salinity, inorganic nutrients, chlorophyll a dissolved organic carbon (DOC), CDOM and fluorescent DOM (FDOM). During this period, two extremely high fresh water intrusions greatly influenced the dynamics of some DOM fractions, in particular the FDOM. Inorganic nutrients and chlorophyll a showed seasonal patterns: A winter period characterized by a high nutrient concentration in surface waters favored the phytoplankton spring bloom; then, summer stratification extended until autumn. This stratification led to nutrient depletion and, consequently, lower chlorophyll a values in the photic zone. The CDOM and FDOM optical active fractions did not follow temporal trends similar to total DOC. This was likely because the potential sources and sinks of these DOM pools are microbial activity and light exposure, and these were acting simultaneously but in opposite directions. Interestingly, DOC exhibited the highest concentrations in summer, coinciding with nutrient and chlorophyll a minima. To explain this mismatch, we propose a sequence of abiotic and biotic phenomena that drive DOC temporal dynamics, This study was supported by the MERMeX/MISTRALS project and is a contribution to international SOLAS program and the projects ANIMA (CTM2015-65720-R) and DOREMI (CTM2012-34294) from the Spanish Ministerio de Economia y Competitividad. E.D. Sánchez-Pérez would like to thank the Consejo Nacional de Ciencia y Tecnologia (CONACyT) for their financial support through a PhD fellowship, With the funding support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S), of the Spanish Research Agency (AEI)

Special issue Microbial Community Structure and Metabolic Networks in Polar Areas.-- 20 pages, 5 figures, 2 tables, supplementary material, We explored how changes of viral abundance and community composition among four contrasting regions in the Southern Ocean relied on physicochemical and microbiological traits. During January–February 2015, we visited areas north and south of the South Orkney Islands (NSO and SSO) characterized by low temperature and salinity and high inorganic nutrient concentration, north of South Georgia Island (NSG) and west of Anvers Island (WA), which have relatively higher temperatures and lower inorganic nutrient concentrations. Surface viral abundance (VA) was highest in NSG (21.50 ± 10.70 × 106 viruses mL−1) and lowest in SSO (2.96 ± 1.48 × 106 viruses mL−1). VA was positively correlated with temperature, prokaryote abundance and prokaryotic heterotrophic production, chlorophyll a, diatoms, haptophytes, fluorescent organic matter, and isoprene concentration, and was negatively correlated with inorganic nutrients (NO3−, SiO42−, PO43−), and dimethyl sulfide (DMS) concentrations. Viral communities determined by randomly amplified polymorphic DNA–polymerase chain reaction (RAPD-PCR) were grouped according to the sampling location, being more similar within them than among regions. The first two axes of a canonical correspondence analysis, including physicochemical (temperature, salinity, inorganic nutrients—NO3−, SiO42−, and dimethyl sulfoniopropionate -DMSP- and isoprene concentrations) and microbiological (chlorophyll a, haptophytes and diatom, and prokaryote abundance and prokaryotic heterotrophic production) factors accounted for 62.9% of the variance. The first axis, temperature-related, accounted for 33.8%; the second one, salinity-related, accounted for 29.1%. Thus, different environmental situations likely select different hosts for viruses, leading to distinct viral communities, The study was supported by the Spanish Ministry of Economy through projects PEGASO (CTM2012-37615) to R. Simó, and BIO-NUC (CGL2013-49020-R) to M. Dall’Osto, With the funding support of the ‘Severo Ochoa Centre of Excellence’ accreditation (CEX2019-000928-S), of the Spanish Research Agency (AEI)

11 pages, 6 figures, 2 tables, Transparent Exopolymer Particles (TEPs) are a subclass of organic particles with high impact in biogeochemical and ecological processes, such as the biological carbon pump, air-sea interactions, or the microbial loop. However, the complexity in production and consumption makes TEP dynamics hardly predictable, calling for the need of descriptive studies about the in situ dynamics of these particles. We followed monthly TEP dynamics and combined them with a dataset of environmental variables during three years in a coastal site of the oligotrophic North Western Mediterranean (Blanes Bay). TEP concentration, ranging from 11.3 to 289.1 μg XG eq L (average 81.7 ± 11.7 μg XG eq L), showed recurrent peaks in early summer (June–July). TEP were temporally disconnected from chlorophyll a maxima, that occurred in late winter and early spring (maxima 1.21 μg L), but they were significantly related to the abundance of specific phytoplankton groups (diatoms and dinoflagellates) and also coincided with periods of low nutrient concentrations. The fraction of particulate organic carbon in the form of TEP (the TEP:POC and TEP:PM ratios) were also highest in early summer, indicating that TEP-enriched particles of low density accumulate in surface waters during stratified periods. We hypothesize that the accumulation of these particles affects the microbial food web by enhancing the activity of specific prokaryotic extracellular enzymes (esterase, β-glucosidase and alkaline phosphatase) and promoting the abundance of heterotrophic nanoflagellates, This work was funded by projects STORM (CTM2009-09352/MAR), PEGASO (CTM2012-37615), ANIMA (CTM2015-65720-R) and REMEI (CTM2015-70340-R) of the Spanish Ministry of Science or Economy. [...] EOR was supported by a Juan de la Cierva Fellowship

12 pages, 5 tables, 6 figures, Transparent exopolymer particles (TEP) play a key role in ocean carbon export and structuring microbial habitats, but information on their distribution across different ocean basins and depths is scarce, particularly in the dark ocean. We measured TEP vertical distribution from the surface to bathypelagic waters in an east-to-west transect across the Mediterranean Sea (MedSea) and the adjacent North East Atlantic Ocean (NEA), and explored their physical and biological drivers. TEP ranged from 0.6 to 81.7 µg XG eq L−1, with the highest values in epipelagic waters above the deep chlorophyll maximum, and in areas near the Gibraltar and Sicily Straits. TEP were significantly related to particulate organic carbon (POC) in all basins and depth layers (epipelagic vs. deep), but the contribution of TEP to POC was higher in the NEA (85%, 79% and 67% in epi-, meso- and bathypelagic waters, respectively) than in the MedSea (from 53% to 62% in epipelagic waters, and from 45% to 48% in meso- and bathypelagic waters), coinciding with higher carbon to nitrogen particulate organic matter ratios in the NEA. The TEP connectivity between epipelagic waters and mesopelagic waters was less straightforward than between mesopelagic waters and bathypelagic waters, with a 23% and 55% of the variance in the relationship between layers explained respectively. Prokaryotes were found to be a likely net source of TEP as inferred by the significant direct relationship observed between prokaryotic heterotrophic abundance and TEP. This assumption was confirmed using experimental incubations, where prokaryotes produced TEP in concentrations ranging from 0.7 (Western Mediterranean, bathypelagic) to 232 (Western Mediterranean, mesopelagic) µg XG eq. L−1 day−1, This work was funded by project HOTMIX (CTM2011-30010/MAR) of the Spanish Ministry of Economy and Innovation, co-financed with FEDER funds. Financial support for the writing of this manuscript was also provided by project FLUXES (CTM2015-69392-C3) to XAAS, MFM and JA, REMEI (CTM2015-70340-R) to JMG, and MNat Scientific Unit of Excellence (UCE.PP2017.03) to IR

17 pages, 5 figures, 4 tables, Transparent exopolymer particles (TEPs) are a class of gel particles, produced mainly by microorganisms, which play important roles in biogeochemical processes such as carbon cycling and export. TEPs (a) are colonized by carbon-consuming microbes; (b) mediate aggregation and sinking of organic matter and organisms, thereby contributing to the biological carbon pump; and (c) accumulate in the surface microlayer (SML) and affect air–sea gas exchange. The first step to evaluate the global influence of TEPs in these processes is the prediction of TEP occurrence in the ocean. Yet, little is known about the physical and biological variables that drive their abundance, particularly in the open ocean. Here we describe the horizontal TEP distribution, along with physical and biological variables, in surface waters along a north–south transect in the Atlantic Ocean during October–November 2014. Two main regions were separated due to remarkable differences: the open Atlantic Ocean (OAO, n=30), and the Southwestern Atlantic Shelf (SWAS, n=10). TEP concentration in the entire transect ranged 18.3–446.8 µg XG eq L−1 and averaged 117.1±119.8 µg XG eq L−1, with the maximum concentrations in the SWAS and in a station located at the edge of the Canary Coastal Upwelling (CU), and the highest TEP to chlorophyll a (TEP:Chl a) ratios in the OAO (183±56) and CU (1760). TEPs were significantly and positively related to Chl a and phytoplankton biomass, expressed in terms of C, along the entire transect. In the OAO, TEPs were positively related to some phytoplankton groups, mainly Synechococcus. They were negatively related to the previous 24 h averaged solar irradiance, suggesting that sunlight, particularly UV radiation, is more a sink than a source for TEP. Multiple regression analyses showed the combined positive effect of phytoplankton and heterotrophic prokaryotes (HPs) on TEP distribution in the OAO. In the SWAS, TEPs were positively related to high nucleic acid-containing prokaryotic cells and total phytoplankton biomass, but not to any particular phytoplankton group. Estimated TEP–carbon constituted an important portion of the particulate organic carbon pool in the entire transect (28 %–110 %), generally higher than the phytoplankton and HP carbon shares, which highlights the importance of TEPs in the cycling of organic matter in the ocean, This research was funded by the Spanish Ministry of Economy and Competitiveness through projects PEGASO (CTM2012–37615) and BIOGAPS (CTM2016-81008-R) to Rafel Simó. Marina Zamanillo was supported by a FPU predoctoralfellowship from the Spanish Ministry of Education and Culture. Eva Ortega-Retuerta was supported by a Marie Curie Actions Intra-European Fellowship (H2020-MSCA-IF-2015-703991)

23 pages, 11 figures, 7 tables, supplementary material https://dx.doi.org/10.3989/scimar.04866.10A, [EN] The relationships between the structure of the phytoplankton community and the bio-optical properties of surface waters were studied during the TransPEGASO cruise along a transect across the Atlantic Ocean that covered seven biogeographical provinces, from the Alborán Sea (SW Mediterranean) to the Patagonian Shelf. We characterized the composition of the phytoplankton community by means of high-performance liquid chromatography and CHEMTAX pigment analyses applied to whole water and two filtration size classes (0.5 mg m–3) with a single Mediterranean (MEDI) sample and those from the southwestern Atlantic Shelf (SWAS). According to CHEMTAX, the most important taxa in the tropical and subtropical Atlantic were Prochlorococcus, haptophytes and Synechoccoccus, while the MEDI and SWAS were dominated by diatoms and haptophytes. Both the VU and HI algorithms, which are based on pigment composition or Chl a concentration, predicted for SWAS a high proportion of nano- and microphytoplankton, while the SFF indicated dominance of the, [ES] Durante la campaña TransPEGASO, realizada a lo largo de un transecto a través del Océano Atlántico que cubrió siete provincias biogeográficas, desde el mar de Alborán (Mediterráneo SO) hasta la Plataforma Patagónica, se estudiaron las relaciones entre la estructura de la comunidad fitoplanctónica y las propiedades bio-ópticas del agua. La composición del fitoplancton en muestras de agua entera y de dos fracciones de tamaño (0.5 mg m ) con una sola muestra mediterránea (MEDI) y las de la plataforma patagónica, en el sudoeste del Atlántico (SWAS). Según CHEMTAX, los taxones más importantes en el Atlántico tropical y subtropical fueron Prochlorococcus, haptofitos y Synechoccoccus, mientras que las provincias MEDI y SWAS estuvieron dominadas por diatomeas y haptofitos. Tanto los algoritmos VU como los HI, que se basan en la composición de pigmentos o en la concentración de Chl a, predijeron para SWAS una alta proporción de nano y microfitoplancton, mientras que la SFF indicó un dominio de la clase de tamaño, Por otra parte, los resultados de CHEMTAX indicaron que, en promedio, la contribución de las diatomeas era importante en esta provincia. Sin embargo, en varias estaciones de SWAS para las que CHEMTAX estimaba una elevada contribución de diatomeas, las observaciones microscópicas encontraron solamente escasas concentraciones de células de diatomeas de tamaño nano- o microplanctónico. Esta discrepancia parece deberse a la presencia, confirmada por microscopía electrónica de barrido, de pequeñas células (, The cruise TransPEGASO was funded by the Spanish Ministry of Economía and Competitividad through project PEGASO (CTM2012-37615). S.N. was supported with a doctoral fellowship from the National Council of Technological and Scientific Development (CNPq) of Brazil

Transparent exopolymer particles (TEP) are a class of gel particles produced mainly by microorganisms. TEP play an important role in the ocean carbon cycle, affect sea–air gas exchange and contribute to organic aerosols. The first step to evaluate the TEP influence in these processes is the prediction of TEP occurrence in the ocean. Yet, little is known about the physical and biological variables that control their abundance, particularly in the open ocean. Here we describe horizontal TEP distribution in the surface waters along a North–South transect in the Atlantic Ocean during October–November 2014. Physical and biological variables were run in parallel. Two main regions were separated due to remarkable differences; the open Atlantic Ocean (OAO, n = 30), and the Southwestern Atlantic Shelf (SWAS, n = 10). TEP concentration in the entire transect ranged from 18.3 to 446.8 µg XG eq L−1 and averaged 117.1 ± 119.8 µg XG eq L−1, with the maximum concentrations in the edge of the Canary Coastal Upwelling (CU, n = 1) and the SWAS, but with the highest TEP to chlorophyll a (TEP : Chl a) ratios at the OAO (CU excluded, average 183 ± 56) and CU (1760.4). TEP were significantly and positively related to Chl a and phytoplankton biomass, expressed in terms of C, along the entire transect. In the OAO, TEP were positively related to some phytoplankton groups, mainly to Synechococcus, and negatively related to the previous 24–hours averaged solar radiation, suggesting the predominance of TEP breaking above the induction of TEP production by UV radiation. Multiple regression analyses showed the combined positive effect of phytoplankton and heterotrophic prokaryotes (HP) on TEP distribution in this region. In the SWAS, TEP were positively related to high nucleic acid prokaryotic cells (HNA) and total phytoplankton biomass, but not with any particular phytoplankton group. TEP constituted an important portion of the particulate organic carbon (POC) pool in the entire transect (28.1–109.8 %), and was generally higher than the phytoplankton and HP fraction, highlighting the importance of TEP in the cycle of organic matter in the ocean.

14 pages, 5 figures, 2 tables, supporting information https://dx.doi.org/10.1111/mec.14421, Biotic and abiotic particles shape the microspatial architecture that defines the microbial aquatic habitat, being particles highly variable in size and quality along oceanic horizontal and vertical gradients. We analysed the prokaryotic (bacterial and archaeal) diversity and community composition present in six distinct particle size classes ranging from the pico- to the microscale (0.2 to 200 μm). Further, we studied their variations along oceanographic horizontal (from the coast to open oceanic waters) and vertical (from the ocean surface into the meso- and bathypelagic ocean) gradients. In general, prokaryotic community composition was more variable with depth than in the transition from the coast to the open ocean. Comparing the six size-fractions, distinct prokaryotic communities were detected in each size-fraction, and whereas bacteria were more diverse in the larger size-fractions, archaea were more diverse in the smaller size-fractions. Comparison of prokaryotic community composition among particle size-fractions showed that most, but not all, taxonomic groups have a preference for a certain size-fraction sustained with depth. Species sorting, or the presence of diverse ecotypes with distinct size-fraction preferences, may explain why this trend is not conserved in all taxa, The present work was possible thanks to the Generalitat de Catalunya Quality Research Group 2014SGR/1179 and projects STORM (CTM2009-09352), ADEPT (CTM2011-23458), NEMO (CTM2011-13137-E), ANIMA (CTM2015-65720-R) and REMEI (CTM2015-70340-R) funded by the former Ministry of Science and Innovation and the Ministry of Economy and Competitiveness. M. Mestre was supported by a JAE-Predoc grant, a Deutscher Akademischer Austausch Dienst (DAAD) grant, and by the Ministry of Labour, Employment and Social Security. The Leibniz society and the MIBI group at IGB are thanked for funding and technical as well as scientific support, respectively. I. Ferrera was funded by a postdoctoral fellowship from the Leibniz society and the IGB. E. Borrull benefited from a FPI predoctoral fellowship from the Spanish Ministry of Economy and Competitiveness. HPG was funded by DFG (German Science Foundation) projects Aquameth (GR1540-21/1), MicroPrime (GR1540-23/1) and Aggregates (GR1540-28/1)

The omnipresence of chromophoric dissolved organic matter (CDOM) in the open ocean enables its use as a tracer for biochemical processes throughout the global overturning circulation. We made an inventory of CDOM optical properties, ideal water age (τ), and apparent oxygen utilization (AOU) along the Atlantic, Indian, and Pacific Ocean waters sampled during the Malaspina 2010 expedition. A water mass analysis was applied to obtain intrinsic, hereinafter archetypal, values of τ, AOU, oxygen utilization rate (OUR), and CDOM absorption coefficients, spectral slopes and quantum yield for each one of the 22 water types intercepted during this circumnavigation. Archetypal values of AOU and OUR have been used to trace the differential influence of water mass aging and aging rates, respectively, on CDOM variables. Whereas the absorption coefficient at 325 nm (a325) and the fluorescence quantum yield at 340 nm (Φ340) increased, the spectral slope over the wavelength range 275–295 nm (S275–295) and the ratio of spectral slopes over the ranges 275–295 nm and 350–400 nm (SR) decreased significantly with water mass aging (AOU). Combination of the slope of the linear regression between archetypal AOU and a325 with the estimated global OUR allowed us to obtain a CDOM turnover time of 634 ± 120 years, which exceeds the flushing time of the dark ocean (>200 m) by 46%. This positive relationship supports the assumption of in situ production and accumulation of CDOM as a by-product of microbial metabolism as water masses turn older. Furthermore, our data evidence that global-scale CDOM quantity (a325) is more dependent on aging (AOU), whereas CDOM quality (S275–295, SR, Φ340) is more dependent on aging rate (OUR).

12 pages, 6 figures, 3 tables, supplementary material http://journal.frontiersin.org/article/10.3389/fmicb.2016.02159/full#supplementary-material, Transparent Exopolymer Particles (TEP) are relevant in particle and carbon fluxes in the ocean, and have economic impact in the desalination industry affecting reverse osmosis membrane fouling. However, general models of their occurrence and dynamics are not yet possible because of the poorly known co-variations with other physical and biological variables. Here, we describe TEP distributions in the NW Mediterranean Sea during late spring 2012, along perpendicular and parallel transects to the Catalan coast. The stations in the parallel transect were sampled at the surface, while the stations in the perpendicular transect were sampled from the surface to the bathypelagic, including the bottom nepheloid layers. We also followed the short-term TEP dynamics along a 2-day cycle in offshore waters. TEP concentrations in the area ranged from 4.9 to 122.8 and averaged 31.4 ± 12.0 μg XG eq L−1. The distribution of TEP measured in transects parallel to the Catalan Coast correlated those of chlorophyll a (Chla) in May but not in June, when higher TEP-values with respect to Chla were observed. TEP horizontal variability in epipelagic waters from the coast to the open sea also correlated to that of Chla, O2 (that we interpret as a proxy of primary production) and bacterial production (BP). In contrast, the TEP vertical distributions in epipelagic waters were uncoupled from those of Chla, as TEP maxima were located above the deep chlorophyll maxima. The vertical distribution of TEP in the epipelagic zone was correlated with O2 and BP, suggesting combined phytoplankton (through primary production) and bacterial (through carbon reprocessing) TEP sources. However, no clear temporal patterns arose during the 2-day cycle. In meso- and bathypelagic waters, where phytoplanktonic sources are minor, TEP concentrations (10.1 ± 4.3 μg XG eq l−1) were half those in the epipelagic, but we observed relative TEP increments coinciding with the presence of nepheloid layers. These TEP increases were not paralleled by increases in particulate organic carbon, indicating that TEP are likely to act as aggregating agents of the mostly inorganic particles present in these bottom nepheloid layers, This work was funded by projects funded by the Spanish Ministry of Science STORM (CTM2009-09352/MAR), SUMMER (CTM2008-03309/MAR), DOREMI (CTM2012-34294), REMEI (CTM2015-70340-R), ANIMA (CTM2015-65720-R), PEGASO (CTM2012-37615), and Grup consolidat de Recerca de la Generalitat de Catalunya (2014SGR/1179)

Transparent Exopolymer Particles (TEP) are relevant in particle and carbon fluxes in the ocean, and have economic impact in the desalination industry affecting reverse osmosis membrane fouling. However, general models of their occurrence and dynamics are not yet possible because of the poorly known co-variations with other physical and biological variables. Here, we describe TEP distributions in the NW Mediterranean Sea during late spring 2012, along perpendicular and parallel transects to the Catalan coast. The stations in the parallel transect were sampled at the surface, while the stations in the perpendicular transect were sampled from the surface to the bathypelagic, including the bottom nepheloid layers. We also followed the short-term TEP dynamics along a 2-day cycle in offshore waters. TEP concentrations in the area ranged from 4.9 to 122.8 and averaged 31.4 ± 12.0 μg XG eq L−1. The distribution of TEP measured in transects parallel to the Catalan Coast correlated those of chlorophyll a (Chla) in May but not in June, when higher TEP-values with respect to Chla were observed. TEP horizontal variability in epipelagic waters from the coast to the open sea also correlated to that of Chla, O2 (that we interpret as a proxy of primary production) and bacterial production (BP). In contrast, the TEP vertical distributions in epipelagic waters were uncoupled from those of Chla, as TEP maxima were located above the deep chlorophyll maxima. The vertical distribution of TEP in the epipelagic zone was correlated with O2 and BP, suggesting combined phytoplankton (through primary production) and bacterial (through carbon reprocessing) TEP sources. However, no clear temporal patterns arose during the 2-day cycle. In meso- and bathypelagic waters, where phytoplanktonic sources are minor, TEP concentrations (10.1 ± 4.3 μg XG eq l−1) were half those in the epipelagic, but we observed relative TEP increments coinciding with the presence of nepheloid layers. These TEP increases were not paralleled by increases in particulate organic carbon, indicating that TEP are likely to act as aggregating agents of the mostly inorganic particles present in these bottom nepheloid layers.

Corrigendum: Turnover time of fluorescent dissolved organic matter in the dark global ocean

Marine dissolved organic matter (DOM) is one of the largest reservoirs of reduced carbon on Earth. In the dark ocean (200 m), most of this carbon is refractory DOM. This refractory DOM, largely produced during microbial mineralization of organic matter, includes humic-like substances generated in situ and detectable by fluorescence spectroscopy. Here we show two ubiquitous humic-like fluorophores with turnover times of 435±41 and 610±55 years, which persist significantly longer than the ~350 years that the dark global ocean takes to renew. In parallel, decay of a tyrosine-like fluorophore with a turnover time of 379±103 years is also detected. We propose the use of DOM fluorescence to study the cycling of resistant DOM that is preserved at centennial timescales and could represent a mechanism of carbon sequestration (humic-like fraction) and the decaying DOM injected into the dark global ocean, where it decreases at centennial timescales (tyrosine-like fraction).

Global warming in the Arctic Ocean may result in changes to the stock and dynamics of nutrients that drive the activity of heterotrophic prokaryotes, a key component of the microbial food web. We performed 14 enrichment experiments during summer in the Beaufort and Chukchi Seas (Western Arctic Ocean), with C (acetate and/or glucose), N (nitrate and/or ammonium) and P (phosphate) amendments. In 8 out of 11 experiments performed with surface waters, prokaryotic heterotrophic production was limited by N, either alone (5 experiments) or in co-limitation with C (3 experiments). This contrasted with the experiments performed using waters from the chlorophyll maxima, where N was never limiting. Diversity analyses (DNA- and RNA-based fingerprinting) suggest that limitation was not restricted to specific operational taxonomic units but common to the different groups. This is the first report of N limitation of prokaryotic heterotrophic production in Arctic surface waters. This control by N may gain importance in future scenarios of higher productivity in the area.

Transparent exopolymer particles (TEP) compose an important pool of particulate organic matter (POM) in aquatic systems. However, no studies of TEP contribution to C export to sediment exist for freshwaters. We quantify the contribution of TEP to C sinking fluxes in an oligotrophic reservoir (Quentar, Southern Spain) by monitoring TEP in the water column and TEP, particulate organic carbon (POC) and dry weight in sedimentation traps. TEP sinking fluxes ranged from 0.73 to 183.23 mg C m−2 day−1 and from 0.51 to 177.04 mg C m−2 day−1 at the surface and at the bottom layer, respectively. These values represent that, over an annual basis, 5.59 Ton TEP-C (over 61.32 Ton POC) are exported, on an average, from the water column to the sediment of Quentar reservoir. TEP concentrations (average = 48.0 μg XG eq l−1) were lower than the scarce data reported for freshwaters. No significant relationships between TEP and Chl a concentrations or BA were observed. Average value for daily sedimentation flux (6.63 g Dry Weight m−2 day−1) in the study reservoir was higher than that documented for low productive natural aquatic ecosystems as a consequence of the high amount of allochthonous material input characterizing reservoirs. TEP contributed to C export to sediment with a value that range from 0.02 to 31%. Our results show that even in man-made systems, which are predominantly controlled by allochthonous inputs, TEP may be relevant for explaining POM settling fluxes.

Bacterioplankton are a heterogenous community composed of cells with different physiological states. The consideration of the active fraction of bacterioplankton as a potential factor affecting the strength of the relationship between bacteria and phytoplankton in the Southern Ocean was evaluated in waters around the Antarctic Peninsula. We estimated active bacterioplankton from uptake of 3H-Leucine (bacterial production [BP]) and using vital stains to estimate their proportion within the total bacterioplankton community, based on their relative nucleic acid content (high [HNA] vs. low [LNA]), and by nucleic acid double staining (NADS), based on their membrane permeability. Then we performed a comparative analysis between total and active bacterioplankton and chlorophyll a (chl a) in this area. Staining with NADS suggested that 61% of all bacteria were viable, a higher proportion of the total bacterial community than previously reported for the Southern Ocean. HNA bacteria comprised 45% of all bacteria, indicating that 16% of bacteria may be viable but with LNA. BP was more strongly related to abundance of LNA cells than NADS-viable or HNA bacteria. The relationship between chl a and bacterial abundance (BA) did not increase when considering the abundance of HNA or NADS-viable cells alone, showing that viability/activity of stains did not enhance the linkage between BA and phytoplankton biomass in the Southern Ocean. In contrast, the relationship between chl a and BP was stronger than those reported in the literature, suggesting that, in this region, BP is closely dependent on phytoplankton. © Inter-Research 2008.

Sunlight reacts with dissolved organic matter (DOM) modifying its availability as bacterial substrate. We assessed the impact of DOM photoproducts and mineral nutrient supply on bacterial growth in seven inland waters from the South of Spain, where DOM is characterized by low chromophoric content and long residence time. Factorial experiments were performed with presence vs absence of DOM photoproducts and mineral nutrient supply. In six of the seven experiments, we found a significant and negative effect of DOM photoproducts on bacterial growth and a significant and positive effect of mineral nutrient supply. The interaction of these two factors leaded to a compensation of negative effects of photoproducts by availability of mineral nutrients. Dissolved organic matter diagenetic status and the ionic environment where organic carbon is dissolved can be influencing bacterial DOM processing.

Special issue More from NASA’s ICESCAPE (Impacts of Climate on EcoSystems and Chemistry of the Arctic Pacific Environment) program, Part 1.-- 9 pages, 7 figures, 2 tables, Colored dissolved organic matter (CDOM), a significant fraction of dissolved organic carbon (DOC), plays various roles in physical and biogeochemical processes in natural waters. In the Arctic Ocean, CDOM is abundant because of major input by large rivers. To better understand the processes that drive variations in CDOM, light absorption coefficients of CDOM [aCDOM(λ), m-1] were extensively documented together with temperature, salinity, chlorophyll a, nitrate concentrations, and bacterial production (BP) and abundance (BA) in the Western Arctic Ocean (WAO) from early to late summer as part of the MALINA and the ICESCAPE expeditions. The data set covered contrasting situations, from bloom to post-bloom conditions and from river-influenced to oceanic water masses. While CDOM photobleaching occurred in the surface layer (, This study was conducted as part of the MALINA Scientific Program funded by ANR (Agence nationale de la recherche), INSU-CNRS (Institut national des sciences de l’univers—Centre national de la recherche scientifique), CNES (Centre national d’études spatiales) and ESA (European Space Agency). This research was also supported by the Ocean Biology and Biogeochemistry Program and the Cryosphere Science Program of the National Aeronautic and Space Administration (NNX10AF42G to K. Arrigo). We also thank a joint contribution to the research programs of UMI Takuvik, ArcticNet (Network Centres of Excellence of Canada) and the Canada Excellence Research Chair in Remote Sensing of Canada’s New Arctic Frontier

8 pages, 6 figures, 5 tables, We have measured both prokaryotic heterotrophic production (PHP) and respiration (R), then providing direct estimates of prokaryotic growth efficiencies (PGE), in the upper mesopelagic zone (300-600m) of the South Atlantic and Indian Oceans. Our results show that in situ R ranged 3-fold, from 87 to 238μmolCm-3d-1. In situ PHP rates were much lower but also more variable than R (ranging from 0.3 to 9.1μmolCm-3d-1). The derived in situ PGE values were on average ~1.4% (from 0.3% to 3.7%), indicating that most of the organic substrates incorporated by prokaryotes were respired instead of being used for growth. Together with the few previous studies on PGE published before for the Atlantic Ocean and Mediterranean Sea, our findings support the hypothesis that the global mesopelagic zone represents a key remineralization site for export production in the open ocean. We also found a strong correlation between R and PGE with temperature across a gradient ranging from 8.7 to 14.9°C. The derived Q10 value of 3.7 suggests that temperature variability in the mesopelagic zone plays a significant role in the remineralization of organic matter. © 2014 Elsevier Ltd., This paper is a contribution to the INGENIO 2010 CONSOLIDER program (CDS2008-00077). IPM was supported by a JAE Pre-doc fellowship, from the Spanish National Research Council (CSIC) and the BBVA Foundation, Spain. IPM, JA and JMG were partly supported by Project HotMix (CTM2011-30010-C02/MAR)

Catalá, Teresa S. ... et. al.-- 8 pages, 4 figures, 1 table, supplementary information http://www.nature.com/ncomms/2015/150129/ncomms6986/full/ncomms6986.html#/supplementary-information, Marine dissolved organic matter (DOM) is one of the largest reservoirs of reduced carbon on Earth. In the dark ocean (>200 m), most of this carbon is refractory DOM. This refractory DOM, largely produced during microbial mineralization of organic matter, includes humic-like substances generated in situ and detectable by fluorescence spectroscopy. Here we show two ubiquitous humic-like fluorophores with turnover times of 435±41 and 610±55 years, which persist significantly longer than the ∼350 years that the dark global ocean takes to renew. In parallel, decay of a tyrosine-like fluorophore with a turnover time of 379±103 years is also detected. We propose the use of DOM fluorescence to study the cycling of resistant DOM that is preserved at centennial timescales and could represent a mechanism of carbon sequestration (humic-like fraction) and the decaying DOM injected into the dark global ocean, where it decreases at centennial timescales (tyrosine-like fraction). © 2015 Macmillan Publishers Limited. All rights reserved, This study was financed by the Malaspina 2010 circumnavigation expedition (grant number CSD2008–00077). C.R.-C. acknowledges funding through a Beatriu de Pinos postdoctoral fellowship from the Generalitat de Catalunya. M.N.-C. was funded by the CSIC Program ‘Junta para la Ampliación de Estudios’ cofinanced by the ESF

The Iberian Peninsula is close to the Saharan Desert, which is the biggest source of atmospheric aerosols of the World. Currently, it is recognized that atmospheric deposition of aerosols over ecosystems is a significant source not only of elements with gaseous phases but also of rock-derived ones. In the last years we have been quantifying the atmospheric flux of elements and substances of biogeochemical interest on the aquatic ecosystems of the South Iberian Peninsula, and their impact on their functioning and structure. The results we are obtaining indicate that atmospheric contribution of P and Ca are essential to explain the functioning of high mountain lakes, and that atmospheric input of organic matter partially supports the pelagic food web of these ecosystems. In this article we offer a summary of some of the results obtained to date. La Península Ibérica está próxima al Desierto del Sahara que es la mayor fuente de aerosoles atmosféricos del Planeta. Actualmente, se reconoce que la deposición de aerosoles sobre los ecosistemas es una entrada significativa no sólo de elementos con fases gaseosas sino, también, de elementos derivados de rocas. En los últimos años hemos estado cuantificado el flujo atmosférico de elementos y sustancias de interés biogeoquímico sobre los ecosistemas acuáticos del sur de la Península Ibérica y el impacto sobre su funcionamiento y estructura. Los resultados que estamos obteniendo indican que los aportes atmosféricos de P y Ca son esenciales para explicar el funcionamiento de los lagos de alta montaña y que las entradas atmosféricas de materia orgánica sostienen parcialmente las redes tróficas pelágicas de estos ecosistemas. En este artículo ofrecemos un resumen de algunos de los resultados obtenidos hasta ahora.

Blanchet, Marine ... et. al.-- Special issue Microbial Ecology of the Continental and Coastal Environments.-- 16 pages, 8 figures, 2 tables, electronic supplementary material http://dx.doi.org/10.1007/s11356-014-3848-x, Spatial increases and temporal shifts in outbreaks of gelatinous plankton have been observed over the past several decades in many estuarine and coastal ecosystems. The effects of these blooms on marine ecosystem functioning and particularly on the dynamics of the heterotrophic bacteria are still unclear. The response of the bacterial community from a Mediterranean coastal lagoon to the addition of dissolved organic matter (DOM) from the jellyfish Aurelia aurita, corresponding to an enrichment of dissolved organic carbon (DOC) by 1.4, was assessed for 22 days in microcosms (8 l). The high bioavailability of this material led to (i) a rapid mineralization of the DOC and dissolved organic nitrogen from the jellyfish and (ii) the accumulation of high concentrations of ammonium and orthophosphate in the water column. DOM from jellyfish greatly stimulated heterotrophic prokaryotic production and respiration rates during the first 2 days; then, these activities showed a continuous decay until reaching those measured in the control microcosms (lagoon water only) at the end of the experiment. Bacterial growth efficiency remained below 20 %, indicating that most of the DOM was respired and a minor part was channeled to biomass production. Changes in bacterial diversity were assessed by tag pyrosequencing of partial bacterial 16S rRNA genes, DNA fingerprints, and a cultivation approach. While bacterial diversity in control microcosms showed little changes during the experiment, the addition of DOM from the jellyfish induced a rapid growth of Pseudoalteromonas and Vibrio species that were isolated. After 9 days, the bacterial community was dominated by Bacteroidetes, which appeared more adapted to metabolize high-molecular-weight DOM. At the end of the experiment, the bacterial community shifted toward a higher proportion of Alphaproteobacteria. Resilience of the bacterial community after the addition of DOM from the jellyfish was higher for metabolic functions than diversity, suggesting that jellyfish blooms can induce durable changes in the bacterial community structure in coastal lagoons. © 2014, Springer-Verlag Berlin Heidelberg, This study was supported by the CNRS-EC2CO program through the GELPROC project (2010–2012): “Interactions between gelatinous plankton and dynamics of prokaryotic communities.”

Special issue The Phytoplankton Megabloom beneath Arctic Sea Ice: Results from the ICESCAPE Program.-- 11 pages, 6 figures, 5 tables, 1 appendix supplementary data https://doi.org/10.1016/j.dsr2.2014.03.015, The activity of heterotrophic bacterioplankton and their response to changes in primary production in the Arctic Ocean is essential to understand biogenic carbon flows in the area. In this study, we explored the patterns of bacterial abundance (BA) and bacterial production (BP) in waters coinciding with a massive under-ice phytoplankton bloom in the Chukchi Sea in summer 2011, where chlorophyll a (chl a) concentrations were up to 38.9mgm-3. Contrary to our expectations, BA and BP did not show their highest values coinciding with the bloom. In fact, bacterial biomass was only 3.5% of phytoplankton biomass. Similarly, average DOC values were similar inside (average 57.2±3.1μM) and outside (average 64.3±4.8μM) the bloom patch. Regression analyses showed relatively weak couplings, in terms of slope values, between chl a or primary production and BA or BP. Multiple regression analyses indicated that both temperature and chl a explained BA and BP variability in the Chukchi Sea. This temperature dependence was confirmed experimentally, as higher incubation temperatures (6.6°C vs. 2.2°C) enhanced BA and BP, with Q10 values of BP up to 20.0. Together, these results indicate that low temperatures in conjunction with low dissolved organic matter release can preclude bacteria to efficiently process a higher proportion of carbon fixed by phytoplankton, with further consequences on the carbon cycling in the area. © 2014 Elsevier Ltd., This work was supported by the Ocean Biology and Biogeochemistry Program and the Cryosphere Science Program of the National Aeronautic and Space Administration (NNX10AF42G to K. Arrigo) and by the French National Research Agency, under the Grant no. ANR-BLAN08-1 310980 to the MALINA project, the LEFE-CYBER and CNES TOSCA programmes, ArcticNet, and the European Space Agency

Forest, A. ...et al. -- 34 pages, 19 figures, 5 tables, A better understanding of how environmental changes affect organic matter fluxes in Arctic marine ecosystems is sorely needed. Here we combine mooring times series, ship-based measurements and remote sensing to assess the variability and forcing factors of vertical fluxes of particulate organic carbon (POC) across the Mackenzie Shelf in 2009. We developed a geospatial model of these fluxes to proceed to an integrative analysis of their determinants in summer. Flux data were obtained with sediment traps moored around 125 m and via a regional empirical algorithm applied to particle size distributions (17 classes from 0.08–4.2 mm) measured by an Underwater Vision Profiler 5. The low fractal dimension (i.e., porous, fluffy particles) derived from the algorithm (1.26 ± 0.34) and the dominance (~ 77%) of rapidly sinking small aggregates (< 0.5 mm) in total fluxes suggested that settling material was the product of recent aggregation processes between marine detritus, gel-like substances, and ballast minerals. Modeled settling velocity of small and large aggregates was, respectively, higher and lower than in previous studies within which a high fractal dimension (i.e., more compact particles) was consequential of deep-trap collection (~400–1300 m). Redundancy analyses and forward selection of abiotic/biotic parameters, linear trends, and spatial structures (i.e., principal coordinates of neighbor matrices, PCNM) were conducted to partition the variation of the 17 POC flux size classes. Flux variability was explained at 69.5% by the addition of a temporal trend, 7 significant PCNM, and 9 biophysical variables. The first PCNM canonical axis (44.5% of spatial variance) reflected the total magnitude of POC fluxes through a shelf-basin gradient controlled by bottom depth and sea ice concentration (p < 0.01). The second most important spatial structure (5.0%) corresponded to areas where shelf break upwelling is known to occur under easterlies and where phytoplankton was dominated by diatoms. Among biophysical parameters, bacterial production and northeasterly wind (upwelling-favorable) were the two strongest corollaries of POC fluxes (r2 cum. = 0.37). Bacteria were correlated with small aggregates, while northeasterly wind was associated with large size classes (> 1 mm ESD), but these two factors were weakly related with each other. Copepod biomass was overall negatively correlated (p < 0.05) with vertical POC fluxes, implying that metazoans acted as regulators of export fluxes, even if their role was minor given that our study spanned the onset of diapause. Our results demonstrate that on interior Arctic shelves where productivity is low in mid-summer, localized upwelling zones (nutrient enrichment) may result in the formation of large filamentous phytoaggregates that are not substantially retained by copepod and bacterial communities, This work would not have been possible without the professional and enthusiastic assistance of the officers and crew members of the CCGS Amundsen. We express gratitude to L. Prieur and C. Marec for their help in the deployment of the CTD rosette and for the onboard processing of UVP5 data. We thank J. Martin, J. Gagnon, A. Mignot, and M. Gosselin for sharing the chlorophyll data in order to post-calibrate the fluorometer. We thank P. Guillot for the validation of physical data. We thank M. Fortier, K. L´evesque, and J. Ehn for the organization of the fieldwork and workshops and for support at sea. We are grateful to three anonymous reviewers for insightful comments that improved the initial manuscript. This study was conducted as part of the Malina Scientific Program funded by ANR (Agence nationale de la recherche), INSU-CNRS (Institut national des sciences de l’univers – Centre national de la recherche scientifique), CNES (Centre national d’´etudes spatiales) and ESA (European Space Agency). Additional support from ArcticNet (a Network of Centres of Excellence of Canada) and from the ArcticNet-Imperial Oil Research Collaboration was welcomed and appreciated. The IAEA is grateful to the Government of the Principality of Monaco for the support provided to its Environment Laboratories. This work is a joint contribution to the Malina Project and to the research programs of Qu´ebec-Oc´ean, ArcticNet, the Takuvik Joint Universit´e Laval/CNRS Laboratory, the Arctic in Rapid Transition (ART) Initiative, to the Canada Research Chair on the Response of Marine Arctic Ecosystems to Climate Warming, and to the Canada Excellence Research Chair (CERC) in Remote Sensing of Canada’s New Arctic Frontier

Most eukaryotic phytoplankton require vitamin B12 to grow. However, the cycling of this organic growth factor has received substantially less attention than other bioactive substances such as trace metals in the marine environment. This is especially true in the Mediterranean Sea, where direct measurements of dissolved vitamins have never been reported. We report here the first direct measurements of dissolved vitamin B12 across longitudinal gradients in Mediterranean waters. The range of vitamin B12 concentrations measured over the whole transect was 0.5-6.2 pM, which is slightly higher than the range (undetectable-4 pM) of ambient concentrations measured in other open ocean basins in the Pacific and Atlantic oceans. The concentrations measured in the western basin were significantly higher (p < 0.05) than those of the eastern basin. They were positively correlated with chlorophyll concentrations in the most western part of the basin, and did not show any significant correlation with any other biological variables in other regions of the sampling transect. © 2013 Bonnet, Tovar-Sánchez, Panzeca, Duarte, Ortega-Retuerta and Sañudo-Wilhelmy.

12 pages, 6 figures, 6 tables, The Mediterranean reservoirs receive frequent atmospheric Saharan dust inputs with soil-derived organic components mostly during the stratification periods, when run-off inputs are particularly limited. Here, we quantified and optically characterized the water-soluble organic carbon (WSOC) of the (dry and wet) atmospheric deposition in collectors placed near three reservoirs from the western Mediterranean Basin. In addition, we determined the WSOC contribution to the pool of dissolved organic carbon (DOC) in the reservoirs and the influence of dust-derived chromophoric organic components on the water transparency during their stratification periods. We found synchronous dynamics in the WSOC atmospheric inputs among the three collectors and in the DOC concentrations among the three reservoirs. The DOC concentrations and the WSOC atmospheric inputs were positive and significantly correlated in the most oligotrophic reservoir (Quéntar) and in the reservoir with the highest ratio of surface area to mixing water depth (Cubillas). Despite these correlations, WSOC atmospheric inputs represented less than 10% of the total DOC pool, suggesting that indirect effects of dust inputs on reservoir DOC may also promote these synchronous patterns observed in the reservoirs. Chromophoric components from dust inputs can significantly reduce the water transparency to the ultraviolet radiation (UVR). The depths where UVR at λ = 320 nm was reduced to ten percent of surface intensity (Z10%) decreased 27 cm in Béznar, 49 cm in Cubillas, and 69 cm in Quéntar due to the dust inputs. Therefore, the increasing dust export to the atmosphere may have consequences for the water transparency of aquatic ecosystems located under the influence of the global dust belt, This work was funded by the Spanish Ministry of Science and Technology (DISPAR, CGL2005-00076 to IR and CGL2008-06101/BOS to IdV) and by the Spanish Ministry of Education and Science (CICYT grant REN2003-03038 to RM-B)

Heterotrophic bacteria contribute significantly to food web structure and biogeochemical cycling in the ocean. In oligotrophic regions such as the Mediterranean Sea, the functioning and productivity of the pelagic ecosystem largely depends on the activity of heterotrophic bacteria which are at the center of the balance between exported and mineralized carbon. As a result of pioneer studies developing techniques to assess bacterial production and abundance, these parameters have been measured in the Mediterranean Sea since the late 1980s. Much more recently, thanks to the advances in molecular methods for exploring microbial biodiversity, an understanding of the biogeochemical implications of the assemblage's taxonomic composition begins to emerge. Today, research efforts on the role of heterotrophic bacteria in the Mediterranean Sea face new challenges related to understanding and foreseeing the probable responses to changing natural and anthropogenic forcing. This chapter aims at reviewing the current state of knowledge on the variability of bacterial abundance, activity and community structure at both spatial and temporal scales in the pelagic waters of the Mediterranean Sea. In addition, the response of heterotrophic bacteria to major human driven changes is evoked. Finally, future directions dealing with key questions and new tools related to research on heterotrophic bacteria are proposed.

14 pages, 6 figures, 5 tables, During August 2009, measurements of bacterial abundance and nucleic acid content were made along with production and respiration in coastal waters of the Beaufort Sea (Arctic Ocean), an area influenced by the Mackenzie River inflow. The main purpose was to evaluate bacterial organic carbon processing with respect to local sources, mainly primary production and river inputs. Bacterial production and abundance generally decreased from river to offshore waters and from surface to deep waters. In contrast, the percentage of high nucleic acid bacteria was higher in deep waters rather than in surface or river waters. Statistical analyses indicated that bacterial production was primarily controlled by temperature and the availability of labile organic matter, as indicated by total dissolved amino acid concentrations. Direct comparisons of bacterial carbon demand and primary production indicated net heterotrophy was common in shelf waters. Net autotrophy was observed at stations in the Mackenzie River plume, suggesting that the carbon fixed in plume waters helped fuel net heterotrophy in the Beaufort Sea margin, This work was supported by the French National Research Agency, under the grant no. ANR-BLAN08-1 310980 to the MALINA project, the LEFE-CYBER and CNES TOSCA programmes, and the European Space Agency. W.H.J was supported by a University of West Florida faculty scholarly and creative activity award. R. B. was supported by the US NSF (grant no 0713915)

Chromophoric dissolved organic matter (CDOM) plays a key role regulating light attenuation in the ocean. This optically reactive pool of organic matter is driven by several physical and biological processes such as photobleaching, photohumification, and biogeneration, that act as primary sinks and sources of CDOM. In this study, we described the geographical and vertical distribution of CDOM in the Antarctic Peninsula area (Southern Ocean), and assessed its potential driving factors, with special emphasis on CDOM photoreactivity. CDOM values were between the detection limit and 2.17 m- 1 at 325 nm and between the detection limit and 0.76 m- 1 at 443 nm (average a325 = 0.36 ± 0.02 m- 1, average a443 = 0.11 ± 0.01 m- 1), with the highest values inside Deception Island in 2004, and the lowest in the Eastern Bransfield Strait. In Bellingshausen Sea, CDOM was higher below the mixed layer suggesting a significant role of photobleaching. By contrast in the Weddell Sea maximum values were found within the mixed layer. In the Weddell Sea, a positive correlation between CDOM and both chlorophyll a and bacterial production and a negative correlation with salinity suggest a biological source of CDOM likely associated to ice melting. Salinity was also negatively related to the spectral slopes from 275 to 295 nm, considered a good proxy for DOM molecular weight. The experimental results demonstrate the photoreactive nature of CDOM, with half lives from 2.1 to 5.1 days due to photobleaching in the upper layer and duplication times from 4.9 to 15.7 days due to photohumification, that highlight the highly dynamic nature of CDOM in this area. © 2009 Elsevier B.V. All rights reserved., This work was funded by the Spanish Ministry of Science and Technology (ICEPOS, REN2002-04165-CO3-02 to CD and DISPAR, CGL2005-00076 to IR).

Chromophoric dissolved and detrital organic matter (CDM), the optically active fraction of organic matter, affects significantly the underwater light environment and interferes with ocean color algorithms. Here, we studied the distribution and dynamics of CDM in waters around the Antarctic Peninsula, Southern Ocean, using remotely sensed data in austral summers from 1997 to 2005. First, we validated the global semi-analytic algorithm Garver-Siegel-Maritorena (GSM) by comparing simultaneous field and satellite measurements of CDM. These comparisons confirmed the validity of CDM satellite measurements obtained by the GSM algorithm (r2=0.74, slope value=1.01±0.16, n=15).We found a higher (20%) contribution of detrital particles to the CDM signal compared to other studies in lower latitudes (average 12%). Patches of higher CDM were observed in coastal areas and zones with recent ice melting. The seasonal variability of CDM, with maximum values at the end of austral summer, appeared to be ultimately controlled by the dynamics of ice, both directly and indirectly through the growth of phytoplankton and other organisms which are potential sources of CDM. At an interannual timescale, CDM dynamics may be driven by climatic forcing such as the Antarctic Oscillation. © 2010 Elsevier B.V., This work was funded by the Spanish Ministry of Science and Technology (ICEPOS, REN2002-04165-CO3-02 to CMD and DISPAR, CGL2005- 00076 to IR) E. O.-R. was supported by fellowships from the Spanish Ministry of Science and Education and the University of Granada.

The study of transparent exopolymer particles (TEP) in the Mediterranean Sea is particularly relevant as they can be promoters of mucilage events, a frequent phenomenon there. We assessed the influence of bacterioplankton on TEP distribution and dynamics across the west-east axis of the Mediterranean Sea. We performed an extensive study of TEP, dissolved carbohydrates, and their relationships with bacterial abundance and bacterial production (BP). A significant and positive relationship was observed between BP and TEP in the study region (r2 = 0.51, p < 0.001). The direct release of TEP by bacteria was experimentally corroborated using regrowth cultures where increases in TEP tracked bacterial growth in abundance and production. These TEP increases were positively related to the increases in BP (r2 = 0.78, p < 0.05). The consistency (similar slopes) of the regression lines between BP and TEP in natural conditions and between the increases of BP and TEP in the experiments underlines the relevant role of bacteria in the formation of TEP in this area. © 2010 Springer Science+Business Media, LLC., This work was funded by the Spanish Ministry of Science and Technology (THRESHOLDS complementary action ref. CTM2005-24238-E to CMD and DISPAR CGL2005-00076 to IR). E.O.-R. was supported by fellowships from the Spanish Ministry of Science and Education and the University of Granada.

Transparent exopolymer particles (TEP) are the most ubiquitous gel particles in the ocean and form abiotically from dissolved precursors. Although these particles can accumulate at the ocean surface, being thus exposed to intense sunlight, the role of solar radiation for the assembly and degradation of TEP is unknown. In this study, we experimentally determined the effects of visible and ultraviolet B (UVB) radiation on (1) TEP degradation (photolysis experiments), (2) TEP assembly from dissolved polymers (photoinhibition experiments) and (3) TEP release by microorganisms. Solar radiation, particularly in the UVB range, caused significant TEP photolysis, with loss rates from 27 to 34% per day. Dissolved polysaccharides did not increase in parallel. No TEP were formed under UVB, visible or dark conditions, indicating that light does not promote TEP assembly. UVB radiation enhanced TEP release by microorganisms, possibly due to cell deaths, or as a protective measure. Increases in UVB may lead to enhanced TEP photolysis in the ocean, with further consequences for TEP dynamics and, ultimately, sea-air gas exchange., This work was funded by Alfred Wegener Institute for Polar and Marine Research and by the Spanish Ministry of Science and Technology (DISPAR, CGL2005-00076 to IR). E. O.-R. was supported by fellowships of the the Spanish Ministry of Science and Education and University of Granada. The publication of this article is financed by CNRS-INSU.

11 páginas, 5 tablas, 6 figuras., We assessed the effects of Saharan dust inputs of particulate matter (PM), total phosphorus (TP), total nitrogen, and water soluble organic carbon (WSOC) on bacterial abundance (BA) in two alpine lakes and two reservoirs in the Mediterranean region. We also experimentally assessed the effects of dust inputs on bacterial activity and community composition and explored the presence of airborne bacteria. We found synchronous BA dynamics at least in one of the study years for each corresponding pair of ecosystems, suggesting an external control. The link between BA dynamics and inputs of PM, WSOC, or TP occurred only in those ecosystems with severe P-limitation and low dissolved organic carbon. The response was most intense in the most P-limited ecosystem. Dust addition had a significant positive effect on bacterial growth and abundance, but not on richness, diversity, or composition of the indigenous bacterial assemblages. We also obtained experimental evidence that some airborne bacteria could develop in oligotrophic waters by observing the growth of gamma-proteobacteria, a group poorly represented in natural aquatic environments., This research was supported by the projects ECOSENSOR (Fundación BBVA, BIOCON04/009) and MICROBIOGEOGRAPHY (080-2007) to IR, REN03-03038 (Ministerio de Ciencia y Tecnologı´a) to RMB, and AERBAC (079-2007) to EOC, FPU (Formacio´n del Profesorado Universitario), and FPI (Formacio´n del Personal Investigador) grants from Spanish Government to E. Ortega-Retuerta, E. Pulido-Villena, and O. Romera.

Chromophoric dissolved organic matter (CDOM), the optically active fraction of dissolved organic matter, is primarily generated by pelagic organisms in the open ocean. In this study, we experimentally determined the quantity and spectral quality of CDOM generated by bacterioplankton using two different substrates (with and without photoproducts) and by Antarctic krill Euphausia superba and evaluated their potential contributions to CDOM dynamics in the peninsular region of the Southern Ocean. CDOM was generated by bacteria in all experiments, and the presence of photoproducts influenced both the quantity and the spectral quality of the resultant CDOM. We confirmed a direct link between bacterial production and CDOM generation, which yielded in situ CDOM duplication times from 31 to 33 d. Antarctic krill as a direct source of CDOM was also confirmed experimentally. We estimated that CDOM generation by krill would lead to CDOM duplication times from 0.48 to 0.80 d within krill swarms. Our findings highlight the potential significance of bacteria and Antarctic krill swarms in the generation of CDOM and underscore the dynamic nature of CDOM in this area.