Francesc Peters

Changes in Bacterioplankton Composition under Different Phytoplankton Regimens

ABSTRACT The results of empirical studies have revealed links between phytoplankton and bacterioplankton, such as the frequent correlation between chlorophyll a and bulk bacterial abundance and production. Nevertheless, little is known about possible links at the level of specific taxonomic groups. To investigate this issue, seawater microcosm experiments were performed in the northwestern Mediterranean Sea. Turbulence was used as a noninvasive means to induce phytoplankton blooms dominated by different algae. Microcosms exposed to turbulence became dominated by diatoms, while small phytoflagellates gained importance under still conditions. Denaturing gradient gel electrophoresis (DGGE) of 16S rRNA gene fragments showed that changes in phytoplankton community composition were followed by shifts in bacterioplankton community composition, both as changes in the presence or absence of distinct bacterial phylotypes and as differences in the relative abundance of ubiquitous phylotypes. Sequencing of DGGE bands showed that four Roseobacter phylotypes were present in all microcosms. The microcosms with a higher proportion of phytoflagellates were characterized by four phylotypes of the Bacteroidetes phylum: two affiliated with the family Cryomorphaceae and two with the family Flavobacteriaceae. Two other Flavobacteriaceae phylotypes were characteristic of the diatom-dominated microcosms, together with one Alphaproteobacteria phylotype (Roseobacter) and one Gammaproteobacteria phylotype (Methylophaga). Phylogenetic analyses of published Bacteroidetes 16S rRNA gene sequences confirmed that members of the Flavobacteriaceae are remarkably responsive to phytoplankton blooms, indicating these bacteria could be particularly important in the processing of organic matter during such events. Our data suggest that quantitative and qualitative differences in phytoplankton species composition may lead to pronounced differences in bacterioplankton species composition.

Helical Lévy walks: Adjusting searching statistics to resource availability in microzooplankton

The searching trajectories of different animals can be described with a broad class of flight length (lj) distributions with P(lj) = lj–μ. Theoretical studies have shown that changes in these distributions (i.e., different μ values) are key to optimizing the long-term encounter statistics under certain searcher–resource scenarios. In particular, they predict the advantage of Lévy searching (μ ≈ 2) over Brownian motion (μ ≥ 3) for low-prey-density scenarios. Here, we present experimental evidence of predicted optimal changes in the flight-time distribution of a predators walk in response to gradual density changes of its moving prey. Flight times of the dinoflagellate Oxyrrhis marina switched from an exponential to an inverse square power-law distribution when the prey (Rhodomonas sp.) decreased in abundance. Concomitantly, amplitude and frequency of the short-term helical path increased. The specific biological mechanisms involved in these searching behavioral changes are discussed. We suggest that, in a three-dimensional environment, a stronger helical component combined with a Lévy walk searching strategy enhances predators encounter rates. Our results support the idea of universality of the statistical laws in optimal searching processes despite variations in the biological details of the organisms.

The significance of the episodic nature of atmospheric deposition to Low Nutrient Low Chlorophyll regions

In the vast Low Nutrient Low-Chlorophyll (LNLC) Ocean, the vertical nutrient supply from the subsurface to the sunlit surface waters is low, and atmospheric contribution of nutrients may be one order of magnitude greater over short timescales. The short turnover time of atmospheric Fe and N supply (<1 month for nitrate) further supports deposition being an important source of nutrients in LNLC regions. Yet, the extent to which atmospheric inputs are impacting biological activity and modifying the carbon balance in oligotrophic environments has not been constrained. Here, we quantify and compare the biogeochemical impacts of atmospheric deposition in LNLC regions using both a compilation of experimental data and model outputs. A metadata-analysis of recently conducted field and laboratory bioassay experiments reveals complex responses, and the overall impact is not a simple “fertilization effect of increasing phytoplankton biomass” as observed in HNLC regions. Although phytoplankton growth may be enhanced, increases in bacterial activity and respiration result in weakening of biological carbon sequestration. The application of models using climatological or time-averaged non-synoptic deposition rates produced responses that were generally much lower than observed in the bioassay experiments. We demonstrate that experimental data and model outputs show better agreement on short timescale (days to weeks) when strong synoptic pulse of aerosols deposition, similar in magnitude to those observed in the field and introduced in bioassay experiments, is superimposed over the mean atmospheric deposition fields. These results suggest that atmospheric impacts in LNLC regions have been underestimated by models, at least at daily to weekly timescales, as they typically overlook large synoptic variations in atmospheric deposition and associated nutrient and particle inputs. Inclusion of the large synoptic variability of atmospheric input, and improved representation and parameterization of key processes that respond to atmospheric deposition, is required to better constrain impacts in ocean biogeochemical models. This is critical for understanding and prediction of current and future functioning of LNLC regions and their contribution to the global carbon cycle.

Species-specific physiological response of dinoflagellates to quantified small-scale turbulence

Turbulence has been shown to alter different aspects of the physiology of some dinoflagellates. The response appears to be species‐specific and dependent on the experimental design and setup used to generate small‐scale turbulence. We examined the variability of the response of three dinoflagellate species to the turbulence, following the same experimental design used by Berdalet (1992) on Akashiwo sanguinea (Hirasaka) Ge. Hansen et Moestrup (=Gymnodinium nelsonii G. W. Martin). In all experiments, turbulence was generated by an orbital shaker at 100 rpm, which corresponded on bulk average, to dissipation rates (ε, quantified using an acoustic Doppler velocimeter) of ≈2 cm2 · s−3. Turbulence did not appreciably affect Gymnodinium sp., a small dinoflagellate. However, Alexandrium minutum Halim and Prorocentrum triestinum J. Schiller exhibited a reduced net growth rate (33% and 28%, respectively) when shaken during the exponential growth phase. Compared to the still cultures, the shaken treatments of A. minutum and P. triestinum increased the mean cell volume (up to 1.4‐ and 2.5‐fold, respectively) and the mean DNA content (up to 1.8‐ and 5.3‐fold, respectively). Cultures affected by turbulence recovered their normal cell properties when returned to still conditions. The swimming speed of the cells exposed to agitation was half that of the unshaken ones. Overall, the response of A. minutum and P. triestinum was similar, but with lower intensity, to that observed previously on A. sanguinea. We found no clear trends related to taxonomy or morphology.

Saharan dust deposition may affect phytoplankton growth in the mediterranean sea at ecological time scales

The surface waters of the Mediterranean Sea are extremely poor in the nutrients necessary for plankton growth. At the same time, the Mediterranean Sea borders with the largest and most active desert areas in the world and the atmosphere over the basin is subject to frequent injections of mineral dust particles. We describe statistical correlations between dust deposition over the Mediterranean Sea and surface chlorophyll concentrations at ecological time scales. Aerosol deposition of Saharan origin may explain 1 to 10% (average 5%) of seasonally detrended chlorophyll variability in the low nutrient-low chlorophyll Mediterranean. Most of the statistically significant correlations are positive with main effects in spring over the Eastern and Central Mediterranean, conforming to a view of dust events fueling needed nutrients to the planktonic community. Some areas show negative effects of dust deposition on chlorophyll, coinciding with regions under a large influence of aerosols from European origin. The influence of dust deposition on chlorophyll dynamics may become larger in future scenarios of increased aridity and shallowing of the mixed layer.

Diel changes in the microstratification of the metalimnetic community in Lake Cisó

The microstratification of the metalimnetic community in Lake Cisó was followed through the diel cycle by means of a fine sampler with syringes spaced at 3 cm intervals. Populations were sharply stratified. The uppermost part of the metalimnion was occupied by the rotifer Anuraeopsis fissa. Next, layers of the ciliate Coleps hirtus and the flagellate Cryptomonas phaseolus were found. Finally, purple sulfur bacteria appeared at the bottom part of the metalimnion. Although in lower abundances, characteristic populations of ciliates were found at each depth: Strombidium inhabited zones with oxygen, while Plagiopyla and Metopus lived at depths with sulfide. Several members of the community moved vertically about 20 cm during the diel cycle. Cryptomonas performed larger vertical migrations of 40 cm. This organism concentrated at the upper metalimnion during the day, and dispersed through the lower metalimnion during the night. At night, Cryptomonas lived in an environment without oxygen and with sulfide concentrations up to 0.6 mM.

Water management practices exacerbate nitrogen retention in Mediterranean catchments

Nitrogen (N) retention sensu lato refers to all processes preventing new reactive nitrogen brought into watersheds through agricultural or industrial activities to be exported by river systems to the sea. Although such processes protect marine systems from the threat of eutrophication and anoxia, they raise other environmental issues, including the acidification of soils, the emission of ammonia and greenhouse gases, and the pollution of aquifers. Despite these implications, the factors involved in N retention are still poorly controlled, particularly in arid and semi-arid systems. The present study evaluates the N fluxes of 38 catchments in the Iberian Peninsula with contrasting climatic characteristics (temperate and Mediterranean), land uses, and water management practices. This diversity allows addressing the contribution of physical and socioecological factors in N retention, and more specifically, exploring the relation between N retention and water regulation. We hypothesise that the extreme flow regulation implemented in the Mediterranean enhances the high N retention values associated with arid and semi-arid regions. The results show that reservoirs and irrigation channels account for >50% of the variability in N retention values, and above a certain regulation threshold, N retention peaks to values >85-90%. Future climate projections forecast a decrease in rainfall and an increase in agricultural intensification and irrigation practices in many world regions, most notably in arid and semi-arid areas. Increased water demand will likely lead to greater flow regulation, and the situation in many areas may resemble that of Iberian Mediterranean catchments. High N retention and the associated environmental risks must therefore be considered and adequately addressed.

Anthropogenic versus mineral aerosols in the stimulation of microbial planktonic communities in coastal waters of the northwestern Mediterranean Sea.

The atmosphere of the northwestern (NW) Mediterranean Sea is affected by continuous inputs of anthropogenic aerosols and episodic Saharan dust events. These atmospheric inputs deliver to the surface waters high amounts of macronutrients and trace metals that can constitute their main source at certain times of the year. The effect of both anthropogenic and crustal particles over the autotrophic and heterotrophic planktonic community assembles was evaluated through three microcosm experiments carried out in the summer of 2013 and in the winter and spring of 2014 at an urban coastal location of the NW Mediterranean (Barcelona, Spain). Particles were added to seawater at a concentration of 0.8mgl-1. The results showed that (i) a greater stimulation of the whole community was observed in summer and spring than in winter; (ii) both kinds of aerosols produced an increase in the growth of phytoplankton, although the stimulation of nanoeukaryotes was significantly larger with anthropogenic aerosols; and (iii) bacterial abundance increased more with mineral dust, whereas bacterial production was more stimulated with anthropogenic inputs. Overall, the effect of atmospheric particles was dependent on their composition and solubility in seawater, as well as on the initial biogeochemical conditions present in the seawater and had the potential to change the net metabolic balance of the microbial planktonic community.

Horizontal and Vertical Distributions of Transparent Exopolymer Particles (TEP) in the NW Mediterranean Sea Are Linked to Chlorophyll a and O2 Variability

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.

Large Saharan dust storms: Implications for chlorophyll dynamics in the Mediterranean Sea

We investigate the large (LDE) and very large (VLDE) Saharan dust deposition events that occurred between 2000 and 2007 and their short-term impact on the dynamics of marine phytoplankton in the Mediterranean Sea. A total of 153 LDE were identified unevenly distributed over the years. Events were more frequent during winter, in the Eastern Mediterranean, and autumn, when they affected both the Western and the Central Mediterranean. Most of the 31 VLDE occurred during winter and autumn in the Central Mediterranean. The dynamics of chlorophyll after VLDE were studied as a proxy for phytoplankton response to atmospheric dust. A significant response of chlorophyll to dust addition was evident; this appeared to be especially true for the Western Mediterranean where a chlorophyll increases of up to 345% was recorded, whereas in the Central Mediterranean it was up to 146% and in the Eastern Mediterranean up to 121%. Chlorophyll response behavior was quite heterogeneous probably as a result of the uniqueness of each VLDE, the differences between Mediterranean areas, the community structure of phytoplankton and the interaction between bacteria and phytoplankton for new resources. An eastward decreasing trend in chlorophyll response was observed, which is in accordance with the relative importance of bacterial activity with respect to phytoplankton. The increase in mineral aerosols with increased aridity in the region together with the decrease in the depth of the mixed layer of the oceans should boost the importance of aerosols fueling marine production.