Beatriz Mouriño-Carballido

Resource supply overrides temperature as a controlling factor of marine phytoplankton growth.

The universal temperature dependence of metabolic rates has been used to predict how ocean biology will respond to ocean warming. Determining the temperature sensitivity of phytoplankton metabolism and growth is of special importance because this group of organisms is responsible for nearly half of global primary production, sustains most marine food webs, and contributes to regulate the exchange of CO2 between the ocean and the atmosphere. Phytoplankton growth rates increase with temperature under optimal growth conditions in the laboratory, but it is unclear whether the same degree of temperature dependence exists in nature, where resources are often limiting. Here we use concurrent measurements of phytoplankton biomass and carbon fixation rates in polar, temperate and tropical regions to determine the role of temperature and resource supply in controlling the large-scale variability of in situ metabolic rates. We identify a biogeographic pattern in phytoplankton metabolic rates, which increase from the oligotrophic subtropical gyres to temperate regions and then coastal waters. Variability in phytoplankton growth is driven by changes in resource supply and appears to be independent of seawater temperature. The lack of temperature sensitivity of realized phytoplankton growth is consistent with the limited applicability of Arrhenius enzymatic kinetics when substrate concentrations are low. Our results suggest that, due to widespread resource limitation in the ocean, the direct effect of sea surface warming upon phytoplankton growth and productivity may be smaller than anticipated.

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.

High sensitivity of ultra-oligotrophic marine ecosystems to atmospheric nitrogen deposition

Using a model of plankton and organic-matter cycling we demonstrate that variable stoichiometric ratios can lead to a more than 5-fold higher sensitivity of simulated carbon export to atmospheric N deposition in the ultra-oligotrophic eastern part of the North Atlantic subtropical gyre compared to the westerly oligotrophic region near Bermuda, often used as a reference site for subtropical regions. Stronger nutrient limitation in the ultra-oligotrophic east causes higher phytoplankton C:N ratios and lower carbon assimilation efficiency of zooplankton in the model, which results in a higher export efficiency of carbon to the deep ocean compared to the less nutrient-limited western site. Our results indicate that previous estimates of oceanic carbon uptake associated with atmospheric nitrogen deposition may not be fully robust and that spatial variability in nutrient stress and ecological stoichiometry could significantly affect the biogeochemical impact of increasing atmospheric deposition of anthropogenic nitrogen.

Nutrient limitation suppresses the temperature dependence of phytoplankton metabolic rates

Climate warming has the potential to alter ecosystem function through temperature-dependent changes in individual metabolic rates. The temperature sensitivity of phytoplankton metabolism is especially relevant, since these microorganisms sustain marine food webs and are major drivers of biogeochemical cycling. Phytoplankton metabolic rates increase with temperature when nutrients are abundant, but it is unknown if the same pattern applies under nutrient-limited growth conditions, which prevail over most of the ocean. Here we use continuous cultures of three cosmopolitan and biogeochemically relevant species (Synechococcus sp., Skeletonema costatum and Emiliania huxleyi) to determine the temperature dependence (activation energy, Ea) of metabolism under different degrees of nitrogen (N) limitation. We show that both CO2 fixation and respiration rates increase with N supply but are largely insensitive to temperature. Ea of photosynthesis (0.11 ± 0.06 eV, mean ± SE) and respiration (0.04 ± 0.17 eV) under N-limited growth is significantly smaller than Ea of growth rate under nutrient-replete conditions (0.77 ± 0.06 eV). The reduced temperature dependence of metabolic rates under nutrient limitation can be explained in terms of enzyme kinetics, because both maximum reaction rates and half-saturation constants increase with temperature. Our results suggest that the direct, stimulating effect of rising temperatures upon phytoplankton metabolic rates will be circumscribed to ecosystems with high-nutrient availability.

Biological N2 Fixation in the Upwelling Region off NW Iberia: Magnitude, Relevance, and Players

The classical paradigm about marine N2 fixation establishes that this process is mainly constrained to nitrogen-poor tropical and subtropical regions, and sustained by the colonial cyanobacterium Trichodesmium spp. and diatom-diazotroph symbiosis. However, the application of molecular techniques allowed determining a high phylogenic diversity and a wide distribution of marine diazotrophs, which extends the range of ocean environments where biological N2 fixation may be relevant. Between February 2014 and December 2015, we carried out 10 one-day samplings in the upwelling system off NW Iberia in order to: 1) investigate the seasonal variability in the magnitude of N2 fixation, 2) determine its biogeochemical role as a mechanism of new nitrogen supply, and 3) quantify the main diazotrophs in the region under contrasting hydrographic regimes. Our results indicate that the magnitude of N2 fixation in this region was relatively low (0.001±0.002 – 0.095±0.024 µmol N m-3 d-1), comparable to the lower-end of rates described for the subtropical NE Atlantic. Maximum rates were observed at surface during both upwelling and relaxation conditions. The comparison with nitrate diffusive fluxes revealed the minor role of N2 fixation (2 fixation activity detected in the region. Quantitative PCR targeting the nifH gene revealed the highest abundances of two sublineages of Candidatus Atelocyanobacterium thalassa or UCYN-A (UCYN-A1 and UCYN-A2) mainly at surface waters during upwelling and relaxation conditions, and of Gammaproteobacteria γ-24774A11 at deep waters during downwelling. Maximum abundance for the three groups were up to 6.7 × 102, 1.5 × 103 and 2.4 × 104 nifH copies L-1, respectively. Our findings demonstrate measurable N2 fixation activity and presence of diazotrophs throughout the year in a nitrogen-rich temperate region.

Marine Primary Productivity Is Driven by a Selection Effect

The number of species of autotrophic communities can increase ecosystem productivity through species complementarity or through a selection effect which occurs when the biomass of the community approaches the monoculture biomass of the most productive species. Here we explore the effect of resource supply on marine primary productivity under the premise that the high local species richness of phytoplankton communities increases resource use through transient selection of productive species. Using concurrent measurements of phytoplankton community structure, nitrate fluxes into the euphotic zone and productivity from a temperate coastal ecosystem, we find that observed productivities are best described by a population growth model in which the dominant species of the community approach their maximum growth rates. We interpret these results as evidence of species selection in communities containing a vast taxonomic repertory. The prevalence of selection effect was supported by open ocean data that show an increase in community dominance across a gradient of nutrient availability. These results highlight the way marine phytoplankton optimize resources and sustain world food stocks. We suggest that the maintenance of phytoplankton species richness is essential to sustain marine primary productivity since it guarantees the occurrence of highly productive species.

Modulation of the Semidiurnal Cycle of Turbulent Dissipation by Wind‐Driven Upwelling in a Coastal Embayment

With two 25‐hour series of turbulent microstructure and currents observations carried out in August 2013, during spring (CHAOS 1) and neap tides (CHAOS 2), we investigated the semidiurnal cycle of turbulent dissipation in an embayment affected by coastal upwelling (Ria de Vigo, NW Iberia). At the time of sampling, the bay hosted a net, wind‐driven bi‐directional positive exchange flow and thermal stratification. Turbulent kinetic energy (TKE) dissipation (e) at the interface between upwelled and surface waters was enhanced by two orders of magnitude during the ebbs (∼ 10−6 W kg−1) with respect to the floods (∼ 10−8 W kg−1). This pattern was caused by the constructive interference of the shear associated with the upwelling and tidal currents. The vertical structure of the tidal currents was consistent with a deformation of tidal ellipses by stratification, which was tightly coupled to the intensity of upwelling. This two‐pronged interaction resulted in a modulation of the semidiurnal cycle of turbulent dissipation by coastal upwelling. Thus, as a result of the upwelling relaxation conditions experienced during CHAOS 1, depth‐integrated interior TKE dissipation rates were higher, by a factor of∼ 2, compared to CHAOS 2. By using a simple model, we determined that observed variations in turbulent mixing had a limited influence on the tidal variations of stratification, which were dominated by straining and advection. The mixing mechanism described here is potentially relevant for the ecology of upwelling bays, as it can stimulate the transport of nutrients from deep‐upwelled waters to the sun‐lit surface layers where primary production takes place.