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1.	Bunse, Carina, et al. (author) Response of marine bacterioplankton pH homeostasis gene expression to elevated CO2 2016 In: Nature Climate Change. - : Macmillan Publishers Ltd.. - 1758-678X .- 1758-6798. ; 6:5, s. 483-487 Journal article (peer-reviewed)abstract Human-induced ocean acidification impacts marine life. Marine bacteria are major drivers of biogeochemical nutrient cycles and energy fluxes1; hence, understanding their performance under projected climate change scenarios is crucial for assessing ecosystem functioning. Whereas genetic and physiological responses of phytoplankton to ocean acidification are being disentangled2, 3, 4, corresponding functional responses of bacterioplankton to pH reduction from elevated CO2 are essentially unknown. Here we show, from metatranscriptome analyses of a phytoplankton bloom mesocosm experiment, that marine bacteria responded to lowered pH by enhancing the expression of genes encoding proton pumps, such as respiration complexes, proteorhodopsin and membrane transporters. Moreover, taxonomic transcript analysis showed that distinct bacterial groups expressed different pH homeostasis genes in response to elevated CO2. These responses were substantial for numerous pH homeostasis genes under low-chlorophyll conditions (chlorophyll a <2.5 μg l−1); however, the changes in gene expression under high-chlorophyll conditions (chlorophyll a >20 μg l−1) were low. Given that proton expulsion through pH homeostasis mechanisms is energetically costly, these findings suggest that bacterioplankton adaptation to ocean acidification could have long-term effects on the economy of ocean ecosystems.
2.	Villarino, Ernesto, et al. (author) Large-scale ocean connectivity and planktonic body size 2018 In: Nature Communications. - : Springer Science and Business Media LLC. - 2041-1723. ; 9 Journal article (peer-reviewed)abstract Global patterns of planktonic diversity are mainly determined by the dispersal of propagules with ocean currents. However, the role that abundance and body size play in determining spatial patterns of diversity remains unclear. Here we analyse spatial community structure - beta-diversity - for several planktonic and nektonic organisms from prokaryotes to small mesopelagic fishes collected during the Malaspina 2010 Expedition. beta-diversity was compared to surface ocean transit times derived from a global circulation model, revealing a significant negative relationship that is stronger than environmental differences. Estimated dispersal scales for different groups show a negative correlation with body size, where less abundant large-bodied communities have significantly shorter dispersal scales and larger species spatial turnover rates than more abundant small-bodied plankton. Our results confirm that the dispersal scale of planktonic and micro-nektonic organisms is determined by local abundance, which scales with body size, ultimately setting global spatial patterns of diversity.
3.	Alonso-Saez, Laura, et al. (author) Factors controlling the year-round variability in carbon flux through bacteria in a coastal marine system 2008 In: Ecosystems (New York. Print). - : Springer Science and Business Media LLC. - 1432-9840 .- 1435-0629. ; 11:3, s. 397-409 Journal article (peer-reviewed)abstract Data from several years of monthly samplings are combined with a 1-year detailed study of carbon flux through bacteria at a NW Mediterranean coastal site to delineate the bacterial role in carbon use and to assess whether environmental factors or bacterial assemblage composition affected the in situ rates of bacterial carbon processing. Leucine (Leu) uptake rates [as an estimate of bacterial heterotrophic production (BHP)] showed high interannual variability but, on average, lower values were found in winter (around 50 pM Leu(-1) h(-1)) as compared to summer (around 150 pM Leu(-1) h(-1)). Leu-to-carbon conversion factors ranged from 0.9 to 3.6 kgC mol Leu(-1), with generally higher values in winter. Leu uptake was only weakly correlated to temperature, and over a full-year cycle (in 2003), Leu uptake peaked concomitantly with winter chlorophyll a (Chl a) maxima, and in periods of high ectoenzyme activities in spring and summer. This suggests that both low molecular weight dissolved organic matter (DOM) released by phytoplankton, and high molecular weight DOM in periods of low Chl a, can enhance BHP. Bacterial respiration (BR, range 7-48 mu g C l(-1) d(-1)) was not correlated to BHP or temperature, but was significantly correlated to DOC concentration. Total bacterial carbon demand (BHP plus BR) was only met by dissolved organic carbon produced by phytoplankton during the winter period. We measured bacterial growth efficiencies by the short-term and the long-term methods and they ranged from 3 to 42%, increasing during the phytoplankton blooms in winter (during the Chl a peaks), and in spring. Changes in bacterioplankton assemblage structure (as depicted by denaturing gradient gel electrophoresis fingerprinting) were not coupled to changes in ecosystem functioning, at least in bacterial carbon use.
4.	Baltar, Federico, et al. (author) High dissolved extracellular enzymatic activity in the deep Central Atlantic Ocean 2010 In: Aquatic Microbial Ecology. - : Inter-Research Science Center. - 0948-3055 .- 1616-1564. ; 58:3, s. 287-302 Journal article (peer-reviewed)abstract The distribution of prokaryotic abundance (PA), prokaryotic heterotrophic production (PHP), and suspended particulate organic material (POM), as well as total and dissolved (operationally defined as passing through 0.2 mu m pore size filters) potential extracellular enzymatic activities (EEA; alpha- and beta-glucosidase [AGase and BGase], leucine aminopeptidase [LAPase], and alkaline phosphatase [APase]) were determined in the meso- and bathypelagic waters of the (sub)tropical Atlantic along an eastern zonal transatlantic transect and a western N-S transect. Significant differences between both transects were found for POM concentration but not for PA, PHP (except in the subsurface and oxygen minimum layer), and dissolved and total EEA. PHP decreased by 3 orders of magnitude from the lower euphotic zone to bathypelagic waters, while PA and cell-specific PHP decreased only by 1 and 2 orders of magnitude, respectively. The proportion of the dissolved to the total EEA was high in the dark ocean for all the enzymes, ranging from 54 to 100, 56 to 100, 65 to 100 and 57 to 97 % for AGase, BGase, LAPase and APase, respectively. The kinetic parameters (V-max, and K-m) of both the dissolved and total fractions of LAPase and APase were very similar throughout the water column, suggesting a similar origin for both dissolved and particulate EEA. Significant correlations of both dissolved and total EEA were found with prokaryotic metabolism and the POM pool. Based on the previous notion that the fraction of dissolved EEA is higher in particle-attached than in free-living microbes, our results suggest that microbial activity in the dark ocean occurs mainly on colloidal and particulate material. This is in agreement with recent genomic evidence. However, these colloidal and particulate materials are prone to disruption during the sampling process. Hence, more selective sampling techniques are needed to specifically collect these deep-water aggregates that probably represent hotspots of microbial activity in the deep ocean.
5.	Baltar, Federico, et al. (author) Prokaryotic extracellular enzymatic activity in relation to biomass production and respiration in the meso- and bathypelagic waters of the (sub)tropical Atlantic 2009 In: Environmental Microbiology. - : Wiley. - 1462-2912 .- 1462-2920. ; 11:8, s. 1998-2014 Journal article (peer-reviewed)abstract P>Prokaryotic extracellular enzymatic activity, abundance, heterotrophic production and respiration were determined in the meso- and bathypelagic (sub)tropical North Atlantic. While prokaryotic heterotrophic production (PHP) decreased from the lower euphotic layer to the bathypelagic waters by two orders of magnitude, prokaryotic abundance and cell-specific PHP decreased only by one order of magnitude. In contrast to cell-specific PHP, cell-specific extracellular enzymatic activity (alpha- and beta-glucosidase, leucine aminopeptidase, alkaline phosphatase) increased with depth as did cell-specific respiration rates. Cell-specific alkaline phosphatase activity increased from the intermediate water masses to the deep waters up to fivefold. Phosphate concentrations, however, varied only by a factor of two between the different water masses, indicating that phosphatase activity is not related to phosphate availability in the deep waters. Generally, cell-specific extracellular enzymatic activities were inversely related to cell-specific prokaryotic leucine incorporation. Thus, it is apparent that the utilization of deep ocean organic matter is linked to higher cell-specific extracellular enzymatic activity and respiration and lower cell-specific PHP than in surface waters.
6.	Gasol, Josep M., et al. (author) Mesopelagic prokaryotic bulk and single-cell heterotrophic activity and community composition in the NW Africa-Canary Islands coastal-transition zone 2009 In: Progress in Oceanography. - : Elsevier BV. - 0079-6611 .- 1873-4472. ; 83, s. 189-196 Journal article (peer-reviewed)abstract Mesopelagic prokaryotic communities have often been assumed to be relatively inactive in comparison to those from epipelagic waters, and therefore unresponsive to the presence of nearby upwelled waters. We have studied the zonal (shelf-ocean), latitudinal, and depth (epipelagic-mesopelagic) variability of microbial assemblages in the NW Africa-Canary Islands coastal-transition zone (CTZ). Vertical profiles of bacterial bulk and single-cell activity through the epi- and mesopelagic waters were combined with point measurements of bacterial respiration, leucine-to-carbon conversion factors and leucine-to-thymidine incorporation ratios. The overall picture that emerges from our study is that prokaryotes in the mesopelagic zone of this area are less abundant than in the epipelagic but have comparable levels of activity. The relationship between prokaryotes and heterotrophic nanoflagellates, their main predators, remains constant throughout the water column, further contradicting the assumption that deep ocean bacterial communities are mostly inactive. Both bulk and single-cell activity showed clear differences between stations, with higher mesopelagic activities closer to the shelf or affected by upwelling features. We also tested whether differences in microbial function between stations could be related to differences in bacterial community structure, and conclude that bacterial communities are very similar at similar depths in the deep ocean, even if the stations present order-of-magnitude differences in bacterial function.
7.	Morán, Xosé Anxelu G., et al. (author) Temperature regulation of marine heterotrophic prokaryotes increases latitudinally as a breach between bottom-up and top-down controls 2017 In: Global Change Biology. - : Wiley. - 1354-1013 .- 1365-2486. ; 23:9, s. 3956-3964 Journal article (peer-reviewed)abstract Planktonic heterotrophic prokaryotes make up the largest living biomass and process most organic matter in the ocean. Determining when and where the biomass and activity of heterotrophic prokaryotes are controlled by resource availability (bottom-up), predation and viral lysis (top-down) or temperature will help in future carbon cycling predictions. We conducted an extensive survey across subtropical and tropical waters of the Atlantic, Indian and Pacific Oceans during the Malaspina 2010 Global Circumnavigation Expedition and assessed indices for these three types of controls at 109 stations (mostly from the surface to 4,000 m depth). Temperature control was approached by the apparent activation energy in eV (ranging from 0.46 to 3.41), bottom-up control by the slope of the log-log relationship between biomass and production rate (ranging from -0.12 to 1.09) and top-down control by an index that considers the relative abundances of heterotrophic nanoflagellates and viruses (ranging from 0.82 to 4.83). We conclude that temperature becomes dominant (i.e. activation energy >1.5 eV) within a narrow window of intermediate values of bottom-up (0.3-0.6) and top-down 0.8-1.2) controls. A pervasive latitudinal pattern of decreasing temperature regulation towards the Equator, regardless of the oceanic basin, suggests that the impact of global warming on marine microbes and their biogeochemical function will be more intense at higher latitudes. Our analysis predicts that 1 degrees C ocean warming will result in increased biomass of heterotrophic prokaryoplankton only in waters with <26 degrees C of mean annual surface temperature.
8.	Sarmento, Hugo, et al. (author) Phytoplankton species-specific release of dissolved free amino acids and their selective consumption by bacteria 2013 In: Limnology and Oceanography. - : Wiley. - 0024-3590 .- 1939-5590. ; 58:3, s. 1123-1135 Journal article (peer-reviewed)abstract Despite representing only a small fraction of the ocean's dissolved organic matter pool, dissolved free amino acids (DFAA) have high turnover rates and are major nitrogen and carbon sources for bacterioplankton. Both phytoplankton and bacterioplankton assimilate and release DFAA, but their consumption and production are difficult to quantify in nature due to their short residence times (min) as dissolved monomers. We segregated DFAA production by phytoplankton and bacterial consumption by measuring individual DFAA concentrations in four axenic phytoplankton cultures during the exponential growth phase, and also after 4 d incubations in the presence of a natural bacterioplankton community. The amounts and composition of the DFAA pool varied widely among phytoplankton species. The proportion of dissolved organic carbon attributed to DFAA varied among cultures. The picoeukaryotic prasinophyte, Micromonas pusilla, released higher amounts of DFAA than the other species tested (diatoms and dinoflagellate), especially alanine, which has been reported as the dominant individual DFAA in some oligotrophic environments. Community structure of heterotrophic prokaryotes responded to differences in the quality of organic matter released among microalgal species, with Roseobacter-related bacteria responding strongly to exudate composition. Our results demonstrate the specificity of DFAA extracellular release among several algal species and their preferential uptake by members of bacterial communities.
9.	Alonso-Saez, Laura, et al. (author) Leucine-to-carbon empirical conversion factor experiments: does bacterial community structure have an influence? 2010 In: Environmental Microbiology. - : Wiley. - 1462-2912 .- 1462-2920. ; 12:11, s. 2988-2997 Journal article (peer-reviewed)abstract The suitability of applying empirical conversion factors (eCFs) to determine bacterial biomass production remains unclear because seawater cultures are usually overtaken by phylotypes that are not abundant in situ. While eCFs vary across environments, it has not been tested whether differences in eCFs are driven by changes in bacterial community composition or by in situ environmental conditions. We carried out seawater cultures throughout a year to analyse the correlation between eCFs and bacterial community structure, analysed by catalysed reporter deposition fluorescence in situ hybridization. Gammaproteobacteria usually dominated seawater cultures, but their abundance exhibited a wide range (25–73% of cell counts) and significantly increased with inorganic nutrient enrichment. Flavobacteria were less abundant but increased up to 40% of cells counts in winter seawater cultures, when in situ chlorophyll a was high. The correlations between eCFs and the abundance of the main broad phylogenetic groups (Gamma-, Alphaproteobacteria and Flavobacteria) were significant, albeit weak, while more specific groups (Alteromonadaceae and Rhodobacteraceae) were not significantly correlated. Our results show that the frequent development of the fast-growing group Alteromonadaceae in seawater cultures does not strongly drive the observed variations in eCFs. Rather, the results imply that environmental conditions and the growth of specific phylotypes interact to determine eCFs.
10.	Aparicio, Fran L., et al. (author) Eutrophication and acidification : Do they induce changes in the dissolved organic matter dynamics in the coastal Mediterranean Sea? 2016 In: Science of the Total Environment. - : Elsevier BV. - 0048-9697 .- 1879-1026. ; 563, s. 179-189 Journal article (peer-reviewed)abstract Two mesocosms experiments were conducted in winter 2010 and summer 2011 to examine how increased pCO(2) and/or nutrient concentrations potentially perturbate dissolved organic matter dynamics in natural microbial assemblages. The fluorescence signals of protein-and humic-like compounds were used as a proxy for labile and non-labile material, respectively, while the evolution of bacterial populations, chlorophyll a (Chl a) and dissolved organic carbon (DOC) concentrations were used as a proxy for biological activity. For both seasons, the presence of elevated pCO(2) did not cause any significant change in the DOC dynamics (p-value < 0.05). The conditions that showed the greatest changes in prokaryote abundances and Chl a content were those amended with nutrients, regardless of the change in pH. The temporal evolution of fluorophores and optical indices revealed that the degree of humification of the organic molecules and their molecular weight changed significantly in the nutrient-amended treatment. The generation of protein-like compounds was paired to increases in the prokaryote abundance, being higher in the nutrient-amended tanks than in the control. Different patterns in the magnitude and direction of the generation of humic-like molecules suggested that these changes depended on initial microbial populations and the availability of extra nutrient inputs. Based on our results, it is expected that in the future projected coastal scenarios the eutrophication processes will favor the transformations of labile and recalcitrant carbon regardless of changes in pCO(2). (c) 2016 The Authors. Published by Elsevier B.V.
11.	Baltar, Federico, et al. (author) Bacterial Versus Archaeal Origin of Extracellular Enzymatic Activity in the Northeast Atlantic Deep Waters 2013 In: Microbial Ecology. - Springer-Verlag : Springer Science and Business Media LLC. - 0095-3628 .- 1432-184X. ; 65:2, s. 277-288 Journal article (peer-reviewed)abstract We determined the total and dissolved extracellularenzymatic activity (EEA) of α-glucosidase and β-glucosidase(AGase and BGase), alkaline phosphatase (APase) and leucineaminopeptidase (LAPase) activities in the epi-, meso- andbathypelagic waters of the subtropical Northeast Atlantic.EEA was also determined in treatments in which bacterialEEAwas inhibited by erythromycin. Additionally, EEA decayexperiments were performed with surface and deep waters todetermine EEA lifetimes in both water masses. The proportionof dissolved to total EEA (66–89 %, 44–88 %, 57–82 % and86–100 % for AGase, BGase, APase and LAPase, respectively)was generally higher than the cell-associated (i.e.,particulate) EEA. The percentage of dissolved to total EEAwas inversely proportional to the percentage of erythromycininhibitedto total EEA. Since erythromycin-inhibited plusdissolved EEA equaled total EEA, this tentatively suggeststhat cell-associated EEA in the open oceanic water column isalmost exclusively of bacterial origin. The decay constants ofdissolved EEAwere in the range of 0.002–0.048 h−1 dependingon the type of extracellular enzyme, temperature and depthin the water column. Although dissolved EEA can have differentorigins, the major contribution of Bacteria to cellassociatedEEA and the long life-time of dissolved EEAsuggest that Bacteria—and not mesophilic Archaea—areessentially the main producers of EEA in the open subtropicalNortheast Atlantic down to bathypelagic layers.
12.	Baltar, Federico, et al. (author) Evidence of prokaryotic metabolism on suspended particulate organic matter in the dark waters of the (sub)tropical North Atlantic 2009 In: Limnology and Oceanography. - 0024-3590 .- 1939-5590. ; 54:1, s. 182-193 Journal article (peer-reviewed)abstract The distribution of prokaryotic abundance (PA), respiratory activity (ETS), heterotrophic production (PHP), and suspended particulate (POM) and dissolved (DOM) organic matter was determined in the meso- and bathypelagic waters of the (sub) tropical North Atlantic. PA decreased by one order of magnitude from the lower euphotic zone to the bathypelagic waters, while ETS decreased by two and PHP by three orders of magnitude. On a section following the Mid-Atlantic Ridge from 35 degrees N to 5 degrees N, ETS below 1000-m depth increased southwards up to three-fold. This latitudinal gradient in the deep waters was paralleled by a six-fold increase in Particulate Organic Carbon (POC), whereas no trend was apparent in the DOM distribution. Significant correlations between POM and ETS were obtained in the water masses between 1000-m and 3000-m depth, the Antarctic Intermediate Water and the North East Atlantic Deep Water. A strong imbalance in the dark ocean was found between prokaryotic carbon demand (estimated through two different approaches) and the carbon sinking flux derived from sediment-trap records corrected with Th-230. The imbalance was greater when deeper in the water column, suggesting that the suspended carbon pool must account for most of the carbon deficit. Our results, together with other recent findings discussed in this paper, indicate that microbial life in the dark ocean is likely more dependent on slowly sinking or buoyant, laterally advected suspended particles than hitherto assumed.
13.	Baltar, Federico, et al. (author) Marine bacterial community structure resilience to changes in protist predation under phytoplankton bloom conditions 2016 In: The ISME Journal. - : Springer Science and Business Media LLC. - 1751-7362 .- 1751-7370. ; 10:3, s. 568-581 Journal article (peer-reviewed)abstract To test whether protist grazing selectively affects the composition of aquatic bacterial communities, we combined high-throughput sequencing to determine bacterial community composition with analyses of grazing rates, protist and bacterial abundances and bacterial cell sizes and physiological states in a mesocosm experiment in which nutrients were added to stimulate a phytoplankton bloom. A large variability was observed in the abundances of bacteria (from 0.7 to 2.4 x 10(6) cells per ml), heterotrophic nanoflagellates (from 0.063 to 2.7 x 10(4) cells per ml) and ciliates (from 100 to 3000 cells per l) during the experiment (similar to 3-, 45- and 30-fold, respectively), as well as in bulk grazing rates (from 1 to 13 x 10(6) bacteria per ml per day) and bacterial production (from 3 to 379 mu g per Cl per day) (1 and 2 orders of magnitude, respectively). However, these strong changes in predation pressure did not induce comparable responses in bacterial community composition, indicating that bacterial community structure was resilient to changes in protist predation pressure. Overall, our results indicate that peaks in protist predation (at least those associated with phytoplankton blooms) do not necessarily trigger substantial changes in the composition of coastal marine bacterioplankton communities.
14.	Baltar, Federico, 1982-, et al. (author) Mesoscale eddies: hot-spots for prokaryotic diversity and function in the ocean 2010 In: The ISME Journal. - : Springer Science and Business Media LLC. - 1751-7362 .- 1751-7370. ; 4, s. 975-988 Journal article (peer-reviewed)abstract To investigate the effects of mesoscale eddies on prokaryotic assemblage structure and activity, we sampled two cyclonic eddies (CEs) and two anticyclonic eddies (AEs) in the permanent eddy-field downstream the Canary Islands. The eddy stations were compared with two far-field (FF) stations located also in the Canary Current, but outside the influence of the eddy field. The distribution of prokaryotic abundance (PA), bulk prokaryotic heterotrophic activity (PHA), various indicators of single-cell activity (such as nucleic acid content, proportion of live cells, and fraction of cells actively incorporating leucine), as well as bacterial and archaeal community structure were determined from the surface to 2000 m depth. In the upper epipelagic layer (0–200 m), the effect of eddies on the prokaryotic community was more apparent, as indicated by the higher PA, PHA, fraction of living cells, and percentage of active cells incorporating leucine within eddies than at FF stations. Prokaryotic community composition differed also between eddy and FF stations in the epipelagic layer. In the mesopelagic layer (200–1000 m), there were also significant differences in PA and PHA between eddy and FF stations, although in general, there were no clear differences in community composition or single-cell activity. The effects on prokaryotic activity and community structure were stronger in AE than CE, decreasing with depth in both types of eddies. Overall, both types of eddies show distinct community compositions (as compared with FF in the epipelagic), and represent oceanic ‘hotspots’ of prokaryotic activity (in the epi- and mesopelagic realms).
15.	Baltar, Federico, et al. (author) Mesoscale variability modulates seasonal changes in the trophic structure of nano- and picoplankton communities across the NW Africa-Canary Islands transition zone 2009 In: Progress in Oceanography. - : Elsevier BV. - 0079-6611 .- 1873-4472. ; 83:1-4, s. 180-188 Journal article (peer-reviewed)abstract The variability of picoplankton and nanoplankton autotrophic (A) and heterotrophic (H) communities was studied along a zonal gradient extending from the NW African shelf to 500 km offshore in two contrasting seasons of the year: early spring (spring) and summer (summer). Plankton abundance was significantly higher in summer than in spring. In particular, heterotrophic prokaryotes (HP) and Prochlorococcus (Proc) were an order of magnitude more abundant in summer, presumably due to a higher loading of dissolved organic matter and higher temperatures. The average ratio of A to H biomass was lower during the summer. Over the African shelf, picoplankton was lowest during summer, while both the autotrophic and heterotrophic nanoflagellates (ANF and HNF) showed the highest abundances. In contrast, in spring, the highest abundance of Picoeukaryotes (PE) and Synechococcus (Syn) was found over or close to the shelf. The offshore sampling sections intersected a complex area of strong mesoscale variability, which affected the plankton distribution. In summer, the entrainment of an upwelling filament around a cyclonic eddy provoked the increase of HP and Syn abundances by about one order of magnitude over the surrounding waters, while PE were more abundant over the core of the eddy (probably due to nutrient pumping). In spring, HP and Syn were more abundant at the boundaries of an anticyclonic eddy and in the filament (where PE also increased). Proc abundance increased up to one order of magnitude in the core of the eddy and in the eddy-oceanic waters front. ANF and HNF showed the highest abundances in the filament and the eddy. in summary, although seasonality affects the background variability in microplankton communities, the mesoscale variability found in the Canary Islands transition zone strongly modulates the patterns of distribution, abundances and changes in community structure, altering the A to H ratio and concomitantly playing a key role modifying the carbon pathways within the food web in the region.
16.	Baltar, Federico, et al. (author) Microbial functioning and community structure variability in the mesopelagic and epipelagic waters of the subtropical Northeast Atlantic Ocean. 2012 In: Applied and Environmental Microbiology. - 0099-2240 .- 1098-5336. ; 78:9, s. 3309-3316 Journal article (peer-reviewed)abstract We analyzed the regional distribution of bulk heterotrophic prokaryotic activity (leucine incorporation) and selected single-cell parameters (cell viability and nucleic acid content) as parameters for microbial functioning, as well as bacterial and archaeal community structure in the epipelagic (0-200 m) and mesopelagic (200-1000 m) subtropical Northeast Atlantic Ocean. We selectively sampled three contrasting regions covering a wide range of surface productivity and oceanographic properties within the same basin: (i) the eddy field south of the Canary Islands, (ii) the open-ocean Subtropical Gyre and (iii) the upwelling filament off Cape Blanc. In the epipelagic waters, a high regional variation in hydrographic parameters and bacterial community structure was detected accompanied, however, by a low variability in microbial functioning. In contrast, mesopelagic microbial functioning was highly variable between the studied regions despite the homogeneous abiotic conditions found therein. More microbial functioning parameters indicated differences among the three regions within the mesopelagic (i.e., viability of cells, nucleic acid content, cell-specific heterotrophic activity, nanoflagellate abundance, prokaryotic to nanoflagellate abundance ratio) than in the epipelagic (i.e., bulk activity, nucleic acid content and nanoflagellate abundance) waters. Our results show that the mesopelagic realm in the NE Atlantic is, in terms of microbial activity, more heterogeneous than its epipelagic counterpart, probably linked to mesoscale hydrographical variations.
17.	Baltar, Federico, et al. (author) Prokaryotic carbon utilization in the dark ocean: : growth efficiency, leucine-to-carbon conversion factors, and their relation 2010 In: Aquatic Microbial Ecology. - : Inter-Research Science Center. - 0948-3055 .- 1616-1564. ; 60:3, s. 227-232 Journal article (peer-reviewed)abstract Experiments were conducted in the mesopelagic subtropical northeast Atlantic Ocean to determine the range of variability in the prokaryotic leucine-to-carbon conversion factor (CF), and prokaryotic growth efficiency (PGE). The way prokaryotic heterotrophic production (PHP) is calcu- lated directly influences PGE (variations of PGE between 1 and 31% were found for a single sample). The empirically obtained deep-water CFs showed a 7-fold variability (0.13 to 0.85 kg C mol–1 Leu), but were always lower than the theoretical CF of 1.55 kg C mol–1 Leu assuming no isotope dilution. Empirically determined CFs were highly correlated to PGE, suggesting that both parameters are rep- resentations of the same basic metabolic processes. Overall, the PGEs obtained in this study suggest that mesopelagic prokaryotic assemblages can sometimes be as important in carbon processing as their epipelagic counterparts.
18.	Baltar, Federico, et al. (author) Response of rare, common and abundant bacterioplankton to anthropogenic perturbations in a Mediterranean coastal site 2015 In: FEMS Microbiology Ecology. - : Oxford University Press (OUP). - 0168-6496 .- 1574-6941. ; 91:6 Journal article (peer-reviewed)abstract Bacterioplankton communities are made up of a small set of abundant taxa and a large number of low-abundant organisms (i.e. 'rare biosphere'). Despite the critical role played by bacteria in marine ecosystems, it remains unknown how this large diversity of organisms are affected by human-induced perturbations, or what controls the responsiveness of rare compared to abundant bacteria. We studied the response of a Mediterranean bacterioplankton community to two anthropogenic perturbations (i.e. nutrient enrichment and/or acidification) in two mesocosm experiments (in winter and summer). Nutrient enrichment increased the relative abundance of some operational taxonomic units (OTUs), e.g. Polaribacter, Tenacibaculum, Rhodobacteraceae and caused a relative decrease in others (e.g. Croceibacter). Interestingly, a synergistic effect of acidification and nutrient enrichment was observed on specific OTUs (e.g. SAR86). We analyzed the OTUs that became abundant at the end of the experiments and whether they belonged to the rare (<0.1% of relative abundance), the common (0.1-1.0% of relative abundance) or the abundant (>1% relative abundance) fractions. Most of the abundant OTUs at the end of the experiments were abundant, or at least common, in the original community of both experiments, suggesting that ecosystem alterations do not necessarily call for rare members to grow.
19.	Baltar, Federico, et al. (author) Significance of non-sinking particulate organic carbon and dark CO2 fixation to heterotrophic carbon demand in the mesopelagic Atlantic 2010 In: Geophysical Research Letters. - 0094-8276 .- 1944-8007. ; 37, s. L09602- Journal article (peer-reviewed)abstract It is generally assumed that sinking particulate organic carbon (POC) constitutes the main source of organic carbon supply to the deep ocean's food webs. However, a major discrepancy between the rates of sinking POC supply (collected with sediment traps) and the prokaryotic organic carbon demand (the total amount of carbon required to sustain the heterotrophic metabolism of the prokaryotes; i.e., production plus respiration, PCD) of deep-water communities has been consistently reported for the dark realm of the global ocean. While the amount of sinking POC flux declines exponentially with depth, the concentration of suspended, buoyant non-sinking POC (nsPOC; obtained with oceanographic bottles) exhibits only small variations with depth in the (sub) tropical Northeast Atlantic. Based on available data for the North Atlantic we show here that the sinking POC flux would contribute only 4-12% of the PCD in the mesopelagic realm (depending on the primary production rate in surface waters). The amount of nsPOC potentially available to heterotrophic prokaryotes in the mesopelagic realm can be partly replenished by dark dissolved inorganic carbon fixation contributing between 12% to 72% to the PCD daily. Taken together, there is evidence that the mesopelagic microheterotrophic biota is more dependent on the nsPOC pool than on the sinking POC supply. Hence, the enigmatic major mismatch between the organic carbon demand of the deep-water heterotrophic microbiota and the POC supply rates might be substantially smaller by including the potentially available nsPOC and its autochthonous production in oceanic carbon cycling models. Citation: Baltar, F., J. Aristegui, E. Sintes, J. M. Gasol, T. Reinthaler, and G. J. Herndl (2010), Significance of non-sinking particulate organic carbon and dark CO2 fixation to heterotrophic carbon demand in the mesopelagic northeast Atlantic.
20.	Baltar, Federico, et al. (author) Strong coast-ocean and surface-depth gradients in prokaryotic assemblage structure and activity in a coastal transition zone region 2007 In: Aquatic Microbial Ecology. - : Inter-Research Science Center. - 0948-3055 .- 1616-1564. ; 50:1, s. 63-74 Journal article (peer-reviewed)abstract The distribution of marine Crenarchaeota Group 1, marine Euryarchaeota Group II and some major groups of Bacteria (SAR 11, Roseobacter, Gammaproteobacteria and Bacteroidetes) was investigated in the North Atlantic water column (surface to 2000 m depth) along a transect from the coastal waters of the NW African upwelling to the offshore waters of the Canary Coastal Transition Zone (CTZ). Catalyzed reporter deposition-fluorescence in situ hybridization (CARD-FISH) was used to describe the prokaryotic assemblages. Bulk picoplankton abundance and leucine incorporation were determined. Pronounced changes in prokaryotic assemblage composition were observed from the coast to the open ocean and at the deep chlorophyll maximum (DCM) with decreasing bulk heterotrophic activity. All bacterial groups decreased in absolute abundances from the coast to the open ocean; both archaeal groups increased towards the open ocean. Prokaryotic abundance and activity decreased 2 and 3 orders of magnitude, respectively, from the surface to 2000 m. Prokaryotic growth rates were high in the mesopelagic zone (similar to 0.13 d(-)1), compared to other reports from the central North Atlantic. SARI 1 in total picoplankton abundance decreased from 42 % in the DCM to 4 % at 2000 m, while marine Crenarchaeota Group I increased from 1 % in the DCM to 39 % in the oxygen minimum layer. A clear influence of the different intermediate water masses was observed on the bulk heterotrophic picoplankton activity, with lower leucine incorporation rates corresponding to layers where patches of Antarctic Intermediate Water were detected. Coast-ocean and surface-depth gradients in bulk prokaryotic abundance and production and assemblage composition were comparable to changes observed in basin-scale studies, pinpointing the CTZs as regions of strong variability in microbial diversity and metabolism.
21.	Gasol, Josep M., et al. (author) Towards a better understanding of microbial carbon flux in the sea 2008 In: Aquatic Microbial Ecology. - : Inter-Research Science Center. - 0948-3055 .- 1616-1564. ; 53:1, s. 21-38 Journal article (peer-reviewed)abstract We now have a relatively good idea of how bulk microbial processes shape the cycling of organic matter and nutrients in the sea. The advent of the molecular biology era in microbial ecology has resulted in advanced knowledge about the diversity of marine microorganisms, suggesting that we might have reached a high level of understanding of carbon fluxes in the oceans. However, it is becoming increasingly clear that there are large gaps in the understanding of the role of bacteria in regulating carbon fluxes. These gaps may result from methodological as well as conceptual limitations. For example, should bacterial production be measured in the light? Can bacterial production conversion factors be predicted, and how are they affected by loss of tracers through respiration? Is it true that respiration is relatively constant compared to production? How can accurate measures of bacterial growth efficiency be obtained? In this paper, we discuss whether such questions could (or should) be addressed. Ongoing genome analyses are rapidly widening our understanding of possible metabolic pathways and cellular adaptations used by marine bacteria in their quest for resources and struggle for survival (e.g. utilization of light, acquisition of nutrients, predator avoidance, etc.). Further, analyses of the identity of bacteria using molecular markers (e.g. subgroups of Bacteria and Archaea) combined with activity tracers might bring knowledge to a higher level. Since bacterial growth (and thereby consumption of DOC and inorganic nutrients) is likely regulated differently in different bacteria, it will be critical to learn about the life strategies of the key bacterial species to achieve a comprehensive understanding of bacterial regulation of C fluxes. Finally, some processes known to occur in the microbial food web are hardly ever characterized and are not represented in current food web models. We discuss these issues and offer specific comments and advice for future research agendas.
22.	Gómez-Consarnau, Laura, et al. (author) Structuring of bacterioplankton communities by specific dissolved organic carbon compounds 2012 In: Environmental Microbiology. - : Wiley. - 1462-2912 .- 1462-2920. ; 14:9, s. 2361-2378 Journal article (peer-reviewed)abstract The main role of microorganisms in the cycling of the bulk dissolved organic carbon pool in the ocean is well established. Nevertheless, it remains unclear if particular bacteria preferentially utilize specific carbon compounds and whether such compounds have the potential to shape bacterial community composition. Enrichment experiments in the Mediterranean Sea, Baltic Sea and the North Sea (Skagerrak) showed that different low-molecular-weight organic compounds, with a proven importance for the growth of marine bacteria (e.g. amino acids, glucose, dimethylsulphoniopropionate, acetate or pyruvate), in most cases differentially stimulated bacterial growth. Denaturing gradient gel electrophoresis fingerprints and 16S rRNA gene sequencing revealed that some bacterial phylotypes that became abundant were highly specific to enrichment with specific carbon compounds (e.g. Acinetobacter sp. B1-A3 with acetate or Psychromonas sp. B3-U1 with glucose). In contrast, other phylotypes increased in relative abundance in response to enrichment with several, or all, of the investigated carbon compounds (e.g. Neptuniibacter sp. M2-A4 with acetate, pyruvate and dimethylsulphoniopropionate, and Thalassobacter sp. M3-A3 with pyruvate and amino acids). Furthermore, different carbon compounds triggered the development of unique combinations of dominant phylotypes in several of the experiments. These results suggest that bacteria differ substantially in their abilities to utilize specific carbon compounds, with some bacteria being specialists and others having a more generalist strategy. Thus, changes in the supply or composition of the dissolved organic carbon pool can act as selective forces structuring bacterioplankton communities.
23.	Lekunberri, Itziar, et al. (author) The phylogenetic and ecological context of cultured and whole genome-sequenced planktonic bacteria from the coastal NW Mediterranean Sea 2014 In: Systematic and Applied Microbiology. - : Elsevier BV. - 0723-2020 .- 1618-0984. ; 37:3, s. 216-228 Journal article (peer-reviewed)abstract Microbial isolates are useful models for physiological and ecological studies and can also be used to reassemble genomes from metagenomic analyses. However, the phylogenetic diversity that can be found among cultured marine bacteria may vary significantly depending on the isolation. Therefore, this study describes a set of 136 bacterial isolates obtained by traditional isolation techniques from the Blanes Bay Microbial Observatory, of which seven strains have had the whole genome sequenced. The complete set was compared to a series of environmental sequences obtained by culture-independent techniques (60 DGGE sequences and 303 clone library sequences) previously obtained by molecular methods. In this way, each isolate was placed in both its "ecological" (time of year, nutrient limitation, chlorophyll and temperature values) context or setting, and its "phylogenetic" landscape (i.e. similar organisms that were found by culture-independent techniques, when they were relevant, and when they appeared). Nearly all isolates belonged to the Gammaproteobacteria, Alphaproteobacteria, or the Bacteroidetes (70, 40 and 20 isolates, respectively). Rarefaction analyses showed similar diversity patterns for sequences from isolates and molecular approaches, except for Alphaproteobacteria where cultivation retrieved a higher diversity per unit effort. Approximately 30% of the environmental clones and isolates formed microdiversity clusters constrained at 99% 16S rRNA gene sequence identity, but the pattern was different in Bacteroidetes (less microdiversity) than in the other main groups. Seventeen cases (12.5%) of nearly complete (98-100%) rRNA sequence identity between isolates and environmental sequences were found: nine in the Alphaproteobacteria, five in the Gammaproteobacteria, and three in the Bacteroidetes, indicating that cultivation could be used to obtain at least some organisms representative of the various taxa detected by molecular methods. Collectively, these results illustrated the largely unexplored potential of culturing on standard media for complementing the study of microbial diversity by culture-independent techniques and for obtaining phylogenetically distinct model organisms from natural seawater.
24.	Malits, Andrea, et al. (author) Viral-Mediated Microbe Mortality Modulated by Ocean Acidification and Eutrophication : Consequences for the Carbon Fluxes Through the Microbial Food Web 2021 In: Frontiers in Microbiology. - : Frontiers Media S.A.. - 1664-302X. ; 12 Journal article (peer-reviewed)abstract Anthropogenic carbon emissions are causing changes in seawater carbonate chemistry including a decline in the pH of the oceans. While its aftermath for calcifying microbes has been widely studied, the effect of ocean acidification (OA) on marine viruses and their microbial hosts is controversial, and even more in combination with another anthropogenic stressor, i.e., human-induced nutrient loads. In this study, two mesocosm acidification experiments with Mediterranean waters from different seasons revealed distinct effects of OA on viruses and viral-mediated prokaryotic mortality depending on the trophic state and the successional stage of the plankton community. In the winter bloom situation, low fluorescence viruses, the most abundant virus-like particle (VLP) subpopulation comprising mostly bacteriophages, were negatively affected by lowered pH with nutrient addition, while the bacterial host abundance was stimulated. High fluorescence viruses, containing cyanophages, were stimulated by OA regardless of the nutrient conditions, while cyanobacteria of the genus Synechococcus were negatively affected by OA. Moreover, the abundance of very high fluorescence viruses infecting small haptophytes tended to be lower under acidification while their putative hosts' abundance was enhanced, suggesting a direct and negative effect of OA on viral-host interactions. In the oligotrophic summer situation, we found a stimulating effect of OA on total viral abundance and the viral populations, suggesting a cascading effect of the elevated pCO(2) stimulating autotrophic and heterotrophic production. In winter, viral lysis accounted for 30 +/- 16% of the loss of bacterial standing stock per day (VMMBSS) under increased pCO(2) compared to 53 +/- 35% in the control treatments, without effects of nutrient additions while in summer, OA had no significant effects on VMMBSS (35 +/- 20% and 38 +/- 5% per day in the OA and control treatments, respectively). We found that phage production and resulting organic carbon release rates significantly reduced under OA in the nutrient replete winter situation, but it was also observed that high nutrient loads lowered the negative effect of OA on viral lysis, suggesting an antagonistic interplay between these two major global ocean stressors in the Anthropocene. In summer, however, viral-mediated carbon release rates were lower and not affected by lowered pH. Eutrophication consistently stimulated viral production regardless of the season or initial conditions. Given the relevant role of viruses for marine carbon cycling and the biological carbon pump, these two anthropogenic stressors may modulate carbon fluxes through their effect on viruses at the base of the pelagic food web in a future global change scenario.
25.	Pinhassi, Jarone, et al. (author) Functional responses of key marine bacteria to environmental change - toward genetic counselling for coastal waters 2022 In: Frontiers in Microbiology. - : Frontiers Media S.A.. - 1664-302X. ; 13 Journal article (peer-reviewed)abstract Coastal ecosystems deteriorate globally due to human-induced stress factors, like nutrient loading and pollution. Bacteria are critical to marine ecosystems, e.g., by regulating nutrient cycles, synthesizing vitamins, or degrading pollutants, thereby providing essential ecosystem services ultimately affecting economic activities. Yet, until now bacteria are overlooked both as mediators and indicators of ecosystem health, mainly due to methodological limitations in assessing bacterial ecosystem functions. However, these limitations are largely overcome by the advances in molecular biology and bioinformatics methods for characterizing the genetics that underlie functional traits of key bacterial populations - "key" in providing important ecosystem services, being abundant, or by possessing high metabolic rates. It is therefore timely to analyze and define the functional responses of bacteria to human-induced effects on coastal ecosystem health. We posit that categorizing the responses of key marine bacterial populations to changes in environmental conditions through modern microbial oceanography methods will allow establishing the nascent field of genetic counselling for our coastal waters. This requires systematic field studies of linkages between functional traits of key bacterial populations and their ecosystem functions in coastal seas, complemented with systematic experimental analyses of the responses to different stressors. Research and training in environmental management along with dissemination of results and dialogue with societal actors are equally important to ensure the role of bacteria is understood as fundamentally important for coastal ecosystems. Using the responses of microorganisms as a tool to develop genetic counselling for coastal ecosystems can ultimately allow for integrating bacteria as indicators of environmental change.
26.	Pinhassi, Jarone, et al. (author) In Memoriam Tom Berman 2013 In: Aquatic Microbial Ecology. - : Inter-Research Science Center. - 0948-3055 .- 1616-1564. ; 70:3, s. 187-187 Journal article (pop. science, debate, etc.)
27.	Vikström, Kevin, 1988- (author) Importance of bacterial maintenance respiration and baseline respiration for development of coastal hypoxia 2018 Doctoral thesis (other academic/artistic)abstract Reduced oxygen concentrations and increasing hypoxic zones havebecome more common in the sea due to climate change andeutrophication. The main cause of oxygen loss in oxygenatedenvironments is respiration. Respiration rates can be estimated usingoptode methodologies which utilize dynamic luminescence quenching toestimate the oxygen concentration declines in dark incubations. Apublished optode methodology was improved by using optodes withtitanium housing instead of plastic housing plausibly trapping oxygen.Drift was highly reduced by the titanium casings leading to a higherprecision and lower detection limit of 0.97 mmol O2 m-3 d-1. 28% ofmeasurements were shown to have non-linear oxygen concentrationdeclines. The rate of oxygen change was derived with a 2nd degreepolynomial at 1 hour from the incubation start. The majority of non-lineardeclines were concave and due to carbon substrate limitation. Analyzingnon-linear trends linearly, a common practice, leads to anunderestimation of respiration by up to 64%.Bacterial maintenance respiration (Rm) was studied using anecophysiological model unverified in natural environments. The modelwas applicable at high productivities but a quadratic model wasdemonstrated to give a better fit. Rm was found to represent a significantpart in the sub-arctic estuary contributing to 58% of the annual specificbacterial respiration. Therefore, Rm may be more important in nature thanpreviously recognized. The ecophysiological model is driven solely by thebacterial specific growth rate (μ) where the relative influence of Rm iselevated as μ decreases. As a consequence, I hypothesize that a reductionin nutrients may not decrease the oxygen consumption but rather shiftbacterial growth based respiration to Rm as μ approaches zero.Baseline respiration (Rbl), defined as ecosystem respiration disconnectedfrom contemporary primary produced carbon, was also studied. Rbl wasshown to be largely supplied by allochthonous carbon in a coastalecosystem and had a contribution of 50% to the annual planktoncommunity respiration in the sub-arctic estuary studied. I claim that Rbland Rm are crucial to include for understanding and managingdevelopment of aquatic hypoxia in an effective and economic manner.
28.	Vila-Costa, Maria, et al. (author) Microbial consumption of organophosphate esters in seawater under phosphorus limited conditions 2019 In: Scientific Reports. - : Nature Publishing Group. - 2045-2322. ; 9 Journal article (peer-reviewed)abstract The anthropogenic perturbation of the phosphorus (P) marine biogeochemical cycle due to synthetic organophosphorus compounds remains unexplored. The objective of this work was to investigate the microbial degradation of organophosphate triesters (OPEs), widely used as plasticizers and flame retardants, in seawater and their effects on the physiology and composition of microbial communities. Experiments were performed in July 2014 using surface seawater from the Blanes Bay Microbial Observatory (NW Mediterranean) to which OPEs were added at environmentally relevant concentrations. The concentrations of OPEs in the dissolved-phase generally decreased after 24 hours of incubation at in situ conditions. The fitted first order reaction constants were significantly different than zero for the trihaloalkyl phosphate, tris(2-chloroethyl) phosphate and trialyl phosphate tricresyl phosphate. In general, OPEs triggered an increase of the percentage of actively respiring bacteria, total bacterial activity, and the number of low-nucleic acid bacteria, and a decrease in the percentage of membrane-compromised bacteria. Members of some bacterial groups, in particular Flavobacteria, increased their specific activity, indicating that seawater contains bacteria with the potential to degrade OPEs. In aged seawater that was presumably depleted of labile dissolved organic carbon and inorganic P, alkaline phosphatase activities significantly decreased when OPEs were added, indicating a relief on P stress, consistent with the role of OPEs as potential P sources.

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