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International audience; Time series of depth data, obtained for 6 oceanic whitetip sharks (Carcharhinus longimanus) tagged with pop-up satellite archival tags, were analyzed in detail. The aim was to improve our knowledge of the vertical behavior of this species. Individuals were tagged in the Atlantic Ocean (n=5) and Indian Ocean (n=1) between 2011 and 2012. Deployment periods for these tags varied from 100 to 178 d. The sharks spent most of their time in the mixed layer, displaying the typical behavior of an epipelagic species. However, analyses revealed complex vertical movement patterns, including marked diel changes that reflect 3 different types of behavior. Results of the generalized additive models indicate that vertical movements were strongly correlated with variations in the depth of the mixed layer. A correlation between vertical movements and sea-surface temperature (SST) was also observed. When SST was above average, oceanic whitetip sharks increased their vertical amplitude, disregarding the reduction of the mixed layer. Thermoregulation likely allows this type of vertical movement.

WOS:000426143100062; International audience; Because of the difficulty in resolving the large variability of N-2 fixation with current methods which rely on discrete sampling, the development of new methods for high resolution measurements is highly desirable. We present a new method for high-frequency measurements of aquatic N-2 fixation by continuous flow-through incubations and spectral monitoring of the acetylene (C2H2, a substrate analog for N-2) reduction to ethylene (C2H4). In this method, named Flow-through Incubation Acetylene Reduction Assays by Cavity Ring-Down Laser Absorption Spectroscopy (FARACAS), dissolved C2H2 is continuously admixed with seawater upstream of a continuous flow stirred-tank reactor (CFSR) in which C2H2 reduction takes place. Downstream of the flow-through incubator, the C2H4 gas is stripped using a bubble column contactor and circulated with a diaphragm pump into a wavelength-scanned cavity ring down laser absorption spectrometer (CRDS). Our method provides high-resolution and precise mapping of aquatic N-2 fixation, its diel cycle, and its response to environmental gradients, and can be adapted to measure other biological processes. The short-duration of the flow-through incubations without preconcentration of cells minimizes potential artifacts such as bottle containment effects while providing near real-time estimates for adaptive sampling. We expect that our new method will improve the characterization of the biogeography and kinetics of aquatic N-2 fixation rates.

This study provides extensive data on planktonic N2 fixation rates across the whole Mediterranean Sea. They show that N2 fixation occurs in Mediterranean waters during the stratification period, with a clear decreasing trend from the oligotrophic western basin (10–76 μmol m−2 d−1) to the ultra oligotrophic eastern basin (0–0.4 μmol m−2 d−1). Highest rates are measured in the less oligotrophic western basin, between the surface and 75 m-depth, where 45 to 75 % of N2 fixation are found within the picoplanktonic fraction (2 fixation in the eastern basin seems negligible, N2 fixation is able to sustain up to 35 % of new primary production during the stratified period in the western basin. These data disagree with indirect estimates of N2 fixation based on geochemical tracers and nutrient budgets, which indicates that N2 fixation increases with increasing N:P ratios and decreasing stable N isotopic signature of particulate organic nitrogen and NO3− from west to east. These results finally point out the need to assess N2 fixation at a higher temporal resolution in order to better understand the diazotrophs' dynamic under contrasted biogeochemical conditions.

The Mediterranean Sea is one of the most oligotrophic marine areas on earth where nitrogen fixation has formally believed to play an important role in carbon and nitrogen fluxes. Although this view is under debate, the diazotrophs responsible for this activity have still not been investigated in the open sea. In this study, we characterised the surface distribution and species richness of unicellular and filamentous diazotrophs across the Mediterranean Sea by combining microscopic counts with size fractionated in situ hybridization (TSA-FISH), and 16S rDNA and nifH genes phylogenies. These genetic analyses were possible owing to the development of a new PCR protocol adapted to scarce microorganisms that can detect as few as 1 cell ml−1 in cultures. Low concentrations of diazotrophic cyanobacteria were detected and this community was dominated at 99.9% by picoplankton hybridized to the Nitro821 probe, specific for unicellular diazotrophic cyanobacteria (UCYN). Among filamentous cyanobacteria only 0.02 filament ml−1 of Richelia were detected in the eastern basin, while small (0.7–1.5 μm) and large (2.5–3.2 μm) Nitro821-targeted cells were recovered at all stations with a mean concentration of 3.5 cell ml−1. The affiliation of the small Nitro821-targeted cells to UCYN-A was confirmed by 16S and nifH phylogenies in the western Mediterranean Sea. In the central and the eastern Mediterranean Sea no 16S rDNA and nifH sequence from UCYN was obtained as cells concentration were close to, or below PCR detection limit. Bradyrhizobium sequences dominated nifH clone libraries from picoplanktonic size fractions. A few sequences of γ-proteobacteria were also detected in the central Mediterranean Sea. While low phosphate and iron concentrations could explain the absence of Trichodesmium sp., the factors that prevent the development of UCYN-B and C remain unknown. We also propose that the dominating picoplankters probably developed specific strategies, such as associations with protists or particles, and/or photosynthetic activity, to acquire carbon for sustaining diazotrophy.

Vivaspin6 (R) ultrafiltration units with molecular weight "cut-off" membranes of 5, 10, 30, 50, and 100 kDa were used together to examine the size distribution of newly formed iron (Fe) colloids in natural seawater samples and in the presence of several different Fe chelators with varying Fe binding strength. Artificial Fe chelators, such as TAC, and 2 kDG, when added at equimolar levels to Fe, supported the formation of a continuum of Fe-ligand colloids between 5 and 100 kDa. More than 90% of the added Fe-55 in these solutions occurred in Fe aggregates/particles larger than 100 kDa. The strong siderophore DFO held the majority of the added Fe-55 in the "truly" soluble fraction = 10 kDa showed similar "cut-off" properties on natural seawater samples collected in the water column off the Peruvian coast. Fe solubility determined with these membranes was approximately six times greater than Fe solubility determined with the 5 kDa membrane and the 0.02 mu m syringe filters. This suggests that a seamless size continuum of organic chelators (

Iron (Fe) is a limiting nutrient for phytoplankton productivity in many different oceanic regions. A critical aspect underlying iron limitation is its low solubility in seawater as this controls the distribution and transport of iron through the ocean. Processes which enhance the solubility of iron in seawater, either through redox reactions or organic complexation, are central to understanding the biogeochemical cycling of iron. In this work we combined iron solubility measurements with parallel factor (PARAFAC) data analysis of Coloured Dissolved Organic Matter (CDOM) fluorescence along a meridional transect through the Atlantic (PS ANT XXVI-4) to examine the hypothesis that marine humic fluorescence is a potential proxy for iron solubility in the surface ocean. PARAFAC analysis revealed 4 components (C1-4), two humic like substances (C2&4) and two protein-like (C1&3). Overall none of the 4 components were significantly correlated with iron solubility, though humic-like components were weakly correlated with iron solubility in iron replete waters. Our analysis suggests that the ligands responsible for maintaining iron in solution in the euphotic zone are sourced from both remineralisation processes and specific ligands produced in response to iron stress and are not easily related to bulk CDOM properties. The humic fluorescence signal was sharply attenuated in surface waters presumably most likely due to photo bleaching, though there was only a weak correlation with the transient photo product H2O2, suggesting longer lifetimes in the photic zone for the fluorescent components identified here. Fil: Heller, M. I.. Geomar-Helmholtz Centre for Ocean Research Kiel; Alemania Fil: Gaiero, Diego Marcelo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Córdoba. Centro de Investigaciones en Ciencias de la Tierra. Universidad Nacional de Córdoba. Facultad de Ciencias Exactas Físicas y Naturales. Centro de Investigaciones en Ciencias de la Tierra; Argentina Fil: Croot, P. L.. Geomar-Helmholtz Centre for Ocean Research Kiel; Alemania

It is now well established that Iron (Fe) is a limiting element in many regions of the open ocean. Our current understanding of the key processes which control iron distribution in the open ocean have been largely based on thermodynamic measurements performed under the assumption of equilibrium conditions. Using this equilibrium approach, researchers have been able to detect and quantify organic complexing ligands in seawater and examine their role in increasing the overall solubility of iron. Our current knowledge about iron bioavailability to phytoplankton and bacteria is also based heavily on carefully controlled laboratory studies where it is assumed the chemical species are in equilibrium in line with the free ion association model and/or its successor the biotic ligand model. Similarly most field work on iron biogeochemistry generally consists of a single profile which is in essence a "snap-shot" in time of the system under investigation. However it is well known that the surface ocean is an extremely dynamic environment and it is unlikely if thermodynamic equilibrium between all the iron species present is ever truly achieved. In sunlit waters this is mostly due to the daily passage of the sun across the sky leading to photoredox processes which alter Fe speciation by cycling between redox states and between inorganic and organic species. Episodic deposition events, dry and wet, are also important perturbations to iron cycling as they bring in new iron to the system and alter the equilibrium between iron species and phases. Here we utilize new field data collected in the open ocean on the complexation kinetics of iron in the surface ocean to identify the important role of weak iron binding ligands (i.e., those that cannot maintain iron in solution indefinitely at seawater pH: alpha(FeL) < alpha(Fe')) in allowing transient increases in iron solubility in response to iron deposition events. Experiments with the thermal O-2(-) source SOTS-1 also indicate the short term impact of this species on iron solubility also with relevance to the euphotic zone. This data highlights the roles of kinetics, redox, and weaker iron binding ligands in the biogeochemical cycling of iron in the ocean.

The performance of 36 models (22 ocean color models and 14 biogeochemical ocean circulation models (BOGCMs)) that estimate depth-integrated marine net primary productivity (NPP) was assessed by comparing their output to in situ 14C data at the Bermuda Atlantic Time series Study (BATS) and the Hawaii Ocean Time series (HOT) over nearly two decades. Specifically, skill was assessed based on the models' ability to estimate the observed mean, variability, and trends of NPP. At both sites, more than 90% of the models underestimated mean NPP, with the average bias of the BOGCMs being nearly twice that of the ocean color models. However, the difference in overall skill between the best BOGCM and the best ocean color model at each site was not significant. Between 1989 and 2007, in situ NPP at BATS and HOT increased by an average of nearly 2% per year and was positively correlated to the North Pacific Gyre Oscillation index. The majority of ocean color models produced in situ NPP trends that were closer to the observed trends when chlorophyll-a was derived from high-performance liquid chromatography (HPLC), rather than fluorometric or SeaWiFS data. However, this was a function of time such that average trend magnitude was more accurately estimated over longer time periods. Among BOGCMs, only two individual models successfully produced an increasing NPP trend (one model at each site). We caution against the use of models to assess multiannual changes in NPP over short time periods. Ocean color model estimates of NPP trends could improve if more high quality HPLC chlorophyll-a time series were available. © 2010 by the American Geophysical Union.

Unicellular diazotrophic cyanobacteria (UCYN2-Fix lineage) were detected using whole-cell hybridization of specific Nitro821 oligonucleotide probe at the coastal and oligotrophic station SOMLIT off Marseilles (France). This station was sampled monthly, for a year and a half (June 2006-November 2007). The UCYN2-Fix community was dominated at 99.9% by picoplankters (0.7-1.5 mu m) mainly as free living in the 0.2-3-mu m size fraction. They were present all the year long with a mean density of 4.6 cell mL(-1), except in summer 2006, when concentrations reached 1.9 x 10(3) cell mL(-1) and 5.3 x 10(3) cell mL(-1) in June and July, respectively. During this bloom, picoplanktonic UCYN2-Fix represented 10.7% of the picocyanobacterial community. The larger size fractions (3-10 mu m and > 10 mu m) were also dominated (98.9%) by picoplanktonic UCYN2-Fix cells, associated with inert particles, or dinoflagellates. However, hardly any nanoplanktonic UCYN2-Fix were detected (0-0.12 cell mL(-1)), and Trichodesmium sp. was observed only once in summer at low concentration (0.03 trichome mL(-1)). We hypothesize that a combination of abiotic parameters, such as elevated temperature, absence of nitrate, presence of phosphate, and an exceptionally high urban pollution event, explain the large bloom of potentially diazotrophic picocyanobacteria. Further studies are needed to confirm the identity of these small cells and their role in nitrogen cycle and marine productivity, especially since some effects of climate change (e. g., increased surface warming and upper-water-column stratification) may increase their importance in a near future.

Scientists and resource managers often use methods and tools that assume ecosystem components respond linearly to environmental drivers and human stressors. However, a growing body of literature demonstrates that many relationships are-non-linear, where small changes in a driver prompt a disproportionately large ecological response. We aim to provide a comprehensive assessment of the relationships between drivers and ecosystem components to identify where and when non-linearities are likely to occur. We focused our analyses on one of the best-studied marine systems, pelagic ecosystems, which allowed us to apply robust statistical techniques on a large pool of previously published studies. In this synthesis, we (1) conduct a wide literature review on single driver-response relationships in pelagic systems, (2) use statistical models to identify the degree of non-linearity in these relationships, and (3) assess whether general patterns exist in the strengths and shapes of non-linear relationships across drivers. Overall we found that non-linearities are common in pelagic ecosystems, comprising at least 52% of all driver-response relation- ships. This is likely an underestimate, as papers with higher quality data and analytical approaches reported non-linear relationships at a higher frequency (on average 11% more). Consequently, in the absence of evidence for a linear relationship, it is safer to assume a relationship is non-linear. Strong non-linearities can lead to greater ecological and socioeconomic consequences if they are unknown (and/or unanticipated), but if known they may provide clear thresholds to inform management targets. In pelagic systems, strongly non-linear relationships are often driven by climate and trophodynamic variables but are also associated with local stressors, such as overfishing and pollution, that can be more easily controlled by managers. Even when marine resource managers cannot influence ecosystem change, they can use information about threshold responses to guide how other stressors are managed and to adapt to new ocean conditions. As methods to detect and reduce uncertainty around threshold values improve, managers will be able to better understand and account for ubiquitous non-linear relationships.