Montserrat Vidal

Phytoplankton across Tropical and Subtropical Regions of the Atlantic, Indian and Pacific Oceans

We examine the large-scale distribution patterns of the nano- and microphytoplankton collected from 145 oceanic stations, at 3 m depth, the 20% light level and the depth of the subsurface chlorophyll maximum, during the Malaspina-2010 Expedition (December 2010-July 2011), which covered 15 biogeographical provinces across the Atlantic, Indian and Pacific oceans, between 35°N and 40°S. In general, the water column was stratified, the surface layers were nutrient-poor and the nano- and microplankton (hereafter phytoplankton, for simplicity, although it included also heterotrophic protists) community was dominated by dinoflagellates, other flagellates and coccolithophores, while the contribution of diatoms was only important in zones with shallow nutriclines such as the equatorial upwelling regions. We applied a principal component analysis to the correlation matrix among the abundances (after logarithmic transform) of the 76 most frequent taxa to synthesize the information contained in the phytoplankton data set. The main trends of variability identified consisted of: 1) A contrast between the community composition of the upper and the lower parts of the euphotic zone, expressed respectively by positive or negative scores of the first principal component, which was positively correlated with taxa such as the dinoflagellates Oxytoxum minutum and Scrippsiella spp., and the coccolithophores Discosphaera tubifera and Syracosphaera pulchra (HOL and HET), and negatively correlated with taxa like Ophiaster hydroideus (coccolithophore) and several diatoms, 2) a general abundance gradient between phytoplankton-rich regions with high abundances of dinoflagellate, coccolithophore and ciliate taxa, and phytoplankton-poor regions (second principal component), 3) differences in dominant phytoplankton and ciliate taxa among the Atlantic, the Indian and the Pacific oceans (third principal component) and 4) the occurrence of a diatom-dominated assemblage (the fourth principal component assemblage), including several pennate taxa, Planktoniella sol, Hemiaulus hauckii and Pseudo-nitzschia spp., in the divergence regions. Our findings indicate that consistent assemblages of co-occurring phytoplankton taxa can be identified and that their distribution is best explained by a combination in different degrees of both environmental and historical influences.

Global distribution and vertical patterns of a prymnesiophyte-cyanobacteria obligate symbiosis

A marine symbiosis has been recently discovered between prymnesiophyte species and the unicellular diazotrophic cyanobacterium UCYN-A. At least two different UCYN-A phylotypes exist, the clade UCYN-A1 in symbiosis with an uncultured small prymnesiophyte and the clade UCYN-A2 in symbiosis with the larger Braarudosphaera bigelowii. We targeted the prymnesiophyte–UCYN-A1 symbiosis by double CARD-FISH (catalyzed reporter deposition-fluorescence in situ hybridization) and analyzed its abundance in surface samples from the MALASPINA circumnavigation expedition. Our use of a specific probe for the prymnesiophyte partner allowed us to verify that this algal species virtually always carried the UCYN-A symbiont, indicating that the association was also obligate for the host. The prymnesiophyte–UCYN-A1 symbiosis was detected in all ocean basins, displaying a patchy distribution with abundances (up to 500 cells ml−1) that could vary orders of magnitude. Additional vertical profiles taken at the NE Atlantic showed that this symbiosis occupied the upper water column and disappeared towards the Deep Chlorophyll Maximum, where the biomass of the prymnesiophyte assemblage peaked. Moreover, sequences of both prymnesiophyte partners were searched within a large 18S rDNA metabarcoding data set from the Tara-Oceans expedition around the world. This sequence-based analysis supported the patchy distribution of the UCYN-A1 host observed by CARD-FISH and highlighted an unexpected homogeneous distribution (at low relative abundance) of B. bigelowii in the open ocean. Our results demonstrate that partners are always in symbiosis in nature and show contrasted ecological patterns of the two related lineages.

Persistent natural acidification drives major distribution shifts in marine benthic ecosystems

Ocean acidification is receiving increasing attention because of its potential to affect marine ecosystems. Rare CO2 vents offer a unique opportunity to investigate the response of benthic ecosystems to acidification. However, the benthic habitats investigated so far are mainly found at very shallow water (less than or equal to 5 m depth) and therefore are not representative of the broad range of continental shelf habitats. Here, we show that a decrease from pH 8.1 to 7.9 observed in a CO2 vent system at 40 m depth leads to a dramatic shift in highly diverse and structurally complex habitats. Forests of the kelp Laminaria rodriguezii usually found at larger depths (greater than 65 m) replace the otherwise dominant habitats (i.e. coralligenous outcrops and rhodolith beds), which are mainly characterized by calcifying organisms. Only the aragonite-calcifying algae are able to survive in acidified waters, while high-magnesium-calcite organisms are almost completely absent. Although a long-term survey of the venting area would be necessary to fully understand the effects of the variability of pH and other carbonate parameters over the structure and functioning of the investigated mesophotic habitats, our results suggest that in addition of significant changes at species level, moderate ocean acidification may entail major shifts in the distribution and dominance of key benthic ecosystems at regional scale, which could have broad ecological and socio-economic implications.

Importance of salt fingering for new nitrogen supply in the oligotrophic ocean.

The input of new nitrogen into the euphotic zone constrains the export of organic carbon to the deep ocean and thereby the biologically mediated long-term CO2 exchange between the ocean and atmosphere. In low-latitude open-ocean regions, turbulence-driven nitrate diffusion from the oceans interior and biological fixation of atmospheric N2 are the main sources of new nitrogen for phytoplankton productivity. With measurements across the tropical and subtropical Atlantic, Pacific and Indian oceans, we show that nitrate diffusion (171±190 μmol m−2 d−1) dominates over N2 fixation (9.0±9.4 μmol m−2 d−1) at the time of sampling. Nitrate diffusion mediated by salt fingers is responsible for ca. 20% of the new nitrogen supply in several provinces of the Atlantic and Indian Oceans. Our results indicate that salt finger diffusion should be considered in present and future ocean nitrogen budgets, as it could supply globally 0.23–1.00 Tmol N yr−1 to the euphotic zone.

Factors controlling seasonal variability of benthic ammonium release and oxygen uptake in Alfacs Bay (Ebro Delta, NW Mediterranean)

The seasonal variability of sediment–water ammonium flux andoxygen uptake was studied in an estuarine bay (Alfacs Bay, Ebro Delta, NWMediterranean) influenced by temporal freshwater discharges. Three stationswith different organic loading were sampled. The relationships of benthicfluxes to bottom water (temperature, dissolved oxygen, ammonium, nitrateplus nitrite) and to sediment (porosity, chlorophyll a derivative pigments,organic carbon and nitrogen) variables were examined. Oxygen uptake rangedfrom 0.3 to 2.5 mmol m−2 h−1 and ammoniumrelease ranged from 6 to 230 µmol m−2 h−1.The lowest value was recorded at the station furthest from the freshwaterinputs, and the highest was at the littoral station nearest the freshwaterdischarge channels (for oxygen uptake) and at the deep station at the saltwedge front (for ammonium flux). Water temperature and the concentration ofchlorophyll a derivative pigments on the surface sediment were revealed asthe main variables to be taken into account to explain the variabilityfound. Changes in fluxes reflecting temperature changes were found at thestation furthest from the freshwater inputs, while at the other, fluxvariability was found to be related to the cycle of functioning offreshwater discharge channels. The different patterns of variability arediscussed in relation to the dynamics of the estuary and to the mainfeatures of benthic nitrogen cycling.

Factors controlling spatial variability in ammonium release within an estuarine bay (Alfacs Bay, Ebro Delta, NW Mediterranean)

Sediment-water ammonium fluxes, oxygen uptake and sediment characteristics were studied in an estuarine bay influenced by temporal freshwater discharges. Sediment at seven stations representing a gradient imposed by freshwater inputs was sampled for sediment-water ammonium and oxygen fluxes, chlorophyll a derivative pigments, organic content, porosity and elemental composition (Fe, Mn, Si, Al). Oxygen uptake decreased along the gradient and correlated with total chlorophyll a derivatives indicating the close coupling between aerobic metabolism and short-time sedimentation events. Ammonium release showed a discontinuous pattern of decrease along the gradient and only correlated with the Fe:Mn atomic ratio. Correlation between the structural properties of the sediment (Si:Al atomic ratio, porosity and organic content) and ammonium release was also found (excluding data from the station with the highest ammonium flux). The extent of the influence of metabolism and sediment structure on ammonium release is discussed.

Surface distribution of dissolved trace metals in the oligotrophic ocean and their influence on phytoplankton biomass and productivity

The distribution of bioactive trace metals has the potential to enhance or limit primary productivity and carbon export in some regions of the world ocean. To study these connections, the concentrations of Cd, Co, Cu, Fe, Mo, Ni, and V were determined for 110 surface water samples collected during the Malaspina 2010 Circumnavigation Expedition (MCE). Total dissolved Cd, Co, Cu, Fe, Mo, Ni, and V concentrations averaged 19.0 ± 5.4 pM, 21.4 ± 12 pM, 0.91 ± 0.4 nM, 0.66 ± 0.3 nM, 88.8 ± 12 nM, 1.72 ± 0.4 nM, and 23.4 ± 4.4 nM, respectively, with the lowest values detected in the Central Pacific and increased values at the extremes of all transects near coastal zones. Trace metal concentrations measured in surface waters of the Atlantic Ocean during the MCE were compared to previously published data for the same region. The comparison revealed little temporal changes in the distribution of Cd, Co, Cu, Fe, and Ni over the last 30 years. We utilized a multivariable linear regression model to describe potential relationships between primary productivity and the hydrological, biological, trace nutrient and macronutrient data collected during the MCE. Our statistical analysis shows that primary productivity in the Indian Ocean is best described by chlorophyll a, NO3, Ni, temperature, SiO4, and Cd. In the Atlantic Ocean, primary productivity is correlated with chlorophyll a, NO3, PO4, mixed layer depth, Co, Fe, Cd, Cu, V, and Mo. The variables salinity, temperature, SiO4, NO3, PO4, Fe, Cd, and V were found to best predict primary productivity in the Pacific Ocean. These results suggest that some of the lesser studied trace elements (e.g., Ni, V, Mo, and Cd) may play a more important role in regulating oceanic primary productivity than previously thought and point to the need for future experiments to verify their potential biological functions.

Polyunsaturated aldehydes from large phytoplankton of the Atlantic Ocean surface (42°n to 33°s).

Polyunsaturated aldehydes (PUAs) are organic compounds mainly produced by diatoms, after cell wounding. These compounds are increasingly reported as teratogenic for species of grazers and deleterious for phytoplanktonic species, but there is still scarce information regarding concentration ranges and the composition of PUAs in the open ocean. In this study, we analyzed the spatial distribution and the type of aldehydes produced by the large-sized (>10 μm) phytoplankton in the Atlantic Ocean surface. Analyses were conducted on PUAs released after mechanical disruption of the phytoplankton cells, referred to here as potential PUAs (pPUAs). Results show the ubiquitous presence of pPUA in the open ocean, including upwelling areas, as well as oligotrophic gyres. Total pPUA concentrations ranged from zero to 4.18 pmol from cells in 1 L. Identified PUAs were heptadienal, octadienal and decadienal, with heptadienal being the most common (79% of total stations). PUA amount and composition across the Atlantic Ocean was mainly related to the nitrogen:phosphorus ratio, suggesting nutrient-driven mechanisms of PUA production. Extending the range of trophic conditions considered by adding data reported for productive coastal waters, we found a pattern of PUA variation in relation to trophic status.

Close and delayed benthic-pelagic coupling in coastal ecosystems : the role of physical constraints

We aimed to analyse the temporal scales of the variability of benthic ammonium flux using data from an estuarine bay (Alfacs Bay, N. W. Mediterranean). Several nitrogen (N) pools in the sediment, their reactivities and their associated fluxes were estimated. Decomposition of labile buried N (4.5 mol N m-2) was found to cause an ammonium flux of 0.1 mmol N m-2 d -1, referred to as background flux. The fluxes measured from bell-jar incubations were usually higher, between 2 and 6 mmol N m-2 d -1, which reflects mineralization of recent sedimentation. A typical sedimentation event was found to account for 25 – 75 mmol m-2 of freshly settled N, which should bring on an ammonium flux of about 1.7–5.0 mmol N m-2 d -1, referred to as fast flux. The concordance between measured and computed benthic fluxes is associated with the coupling of benthic fluxes to production and sedimentation. Close benthic–pelagic coupling was observed in winter and early spring, while a delayed flux response to sedimentation, with transient variations of pore water ammonium profiles, showing surface peaks and decreasing concentrations with depth, was found in autumn. Structures, such as viscous layers, which develop over the sediment–water interface, were found to be essential to the regulation of benthic processes and to lead to transient variations of pore water nutrient concentrations and associated fluxes. The temporal scales of the benthic flux response to sedimentation were discussed in terms of the physical structures involved in decomposition (the bulk sediment, the viscous layers or the fresh settled layer), the processes controlling kinetics and diffusion laws. Several scenarios for the benthic- pelagic coupling in Alfacs Bay, in which local (estuarine circulation) and climatic components combine to yield the variability observed, were examined.