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Plankton community respiration (R) is a major component of the carbon flux in aquatic ecosystems. However, current methods to measure actual respiration from oxygen consumption at relevant spatial scales are not sensitive enough in oligotrophic environments where respiration rates are very low. To overcome this drawback, more sensitive indirect enzymatic approaches are commonly used as R proxies. The in vivo electron transport system (ETSvivo) assay, which measures the reduction of (2-(4-iodophenyl)-3-(4-nitrophenyl)-5-phenyl tetrazolium chloride salt, INT) to INT-formazan in the presence of natural substrate levels, was recently proposed as an indirect reliable estimation of R for natural plankton communities. However, under in vivo conditions, formazan salts could be toxic to the cells. Here, we test the toxicity of 0.2 mM of final INT concentration, widely used for ETSvivo assays, on natural bacterial assemblages collected in coastal and oceanic waters off Gran Canaria (Canary Islands, subtropical North Atlantic), in eight independent experiments. After 0.5 h of incubation, a significant but variable decline in cell viability (14–49%) was observed in all samples inoculated with INT. Moreover, INT also inhibited leucine uptake in less than 90 min of incubation. In the light of these results, we argue that enzymatic respiratory rates obtained with the ETSvivo method need to be interpreted with caution to derive R in oceanic regions where bacteria largely contribute to community respiration. Moreover, the variable toxicity on bacterial assemblages observed in our experiments questions the use of a single R/ETSvivo relationship as a universal proxy for regional studies.

The reaction of CH ₄ with chlorine (Cl) radicals in the atmosphere is associated with an extraordinarily strong isotopic fractionation, where ¹2 CH ₄ reacts about 70 ‰ faster with Cl than ¹3 CH ₄ . Therefore, although the Cl-based sink of CH ₄ constitutes only a small contribution to its total removal rate, the uncertainty in this small sink has been identified as one of the two largest uncertainties of isotope-based CH ₄ source apportionment at the global scale. The uncertainty arises from the fact that Cl levels in the atmosphere are so low that they cannot be detected directly. One very sensitive indirect method to identify and quantify the CH ₄ + Cl reaction in the atmosphere is the detection of the extremely ¹3 C-depleted reaction product carbon monoxide (CO) from this reaction. This article reviews the concept of this approach, its successful application in the atmosphere, its challenges and opportunities for identifying and quantifying Cl-based removal of CH ₄ at the regional and global scale and its potential to detect and evaluate possible attempts to enhance CH ₄ removal from the atmosphere.

Reduction of anthropogenic CO 2 emissions alone will not sufficiently restrict global warming and enable the 1.5°C goal of the Paris agreement to be met. To effectively counteract climate change, measures to actively remove carbon dioxide from the atmosphere are required. Artificial upwelling has been proposed as one such carbon dioxide removal technique. By fueling primary productivity in the surface ocean with nutrient-rich deep water, it could potentially enhance downward fluxes of particulate organic carbon (POC) and carbon sequestration. In this study we investigated the effect of different intensities of artificial upwelling combined with two upwelling modes (recurring additions vs. one singular addition) on POC export, sinking matter stoichiometry and remineralization depth. We carried out a 39 day-long mesocosm experiment in the subtropical North Atlantic, where we fertilized oligotrophic surface waters with different amounts of deep water. The total nutrient inputs ranged from 1.6 to 11.0 μmol NO 3 – L –1 . We found that on the one hand POC export under artificial upwelling more than doubled, and the molar C:N ratios of sinking organic matter increased from values around Redfield (6.6) to ∼8–13, which is beneficial for potential carbon dioxide removal. On the other hand, sinking matter was remineralized at faster rates and showed lower sinking velocities, which led to shallower remineralization depths. Particle properties were more favorable for deep carbon export in the recurring upwelling mode, while in the singular mode the C:N increase of sinking matter was more pronounced. In both upwelling modes roughly half of the produced organic carbon was retained in the water column until the end of the experiment. This suggests that the plankton communities were still in the process of adjustment, possibly due to the different response times of producers and consumers. There is thus a need for studies with longer experimental durations to quantify the responses of fully adjusted communities. Finally, our results revealed that artificial upwelling affects a variety of sinking particle properties, and that the intensity and mode with which it is applied control the strength of the effects.

In the face of climate change there is a need to reduce atmospheric CO 2 concentrations. Artificial upwelling of nutrient-rich deep waters has been proposed as a method to enhance the biological carbon pump in oligotrophic oceanic regions in order to increase carbon sequestration. Here we examine the effect of different artificial upwelling intensities and modes (single pulse versus recurring pulses) on the dynamics of the dissolved organic matter pool (DOM). We introduced nutrient-rich deep water to large scale mesocosms (~44 m 3 ) in the oligotrophic subtropical North Atlantic and found that artificial upwelling strongly increased DOM concentrations and changed its characteristics. The magnitude of the observed changes was related to the upwelling intensity: more intense treatments led to higher accumulation of dissolved organic carbon (>70 μM of excess DOC over ambient waters for the most intense) and to comparatively stronger changes in DOM characteristics (increased proportions of chromophoric DOM (CDOM) and humic-like fluorescent DOM), suggesting a transformation of the DOM pool at the molecular level. Moreover, the single upwelling pulse resulted in higher CDOM quantities with higher molecular weight than the recurring upwelling mode. Together, our results indicate that under artificial upwelling, large DOM pools may accumulate in the surface ocean without being remineralized in the short-term. Possible reasons for this persistence could be a combination of the molecular diversification of DOM due to microbial reworking, nutrient limitation and reduced metabolic capabilities of the prokaryotic communities within the mesocosms. Our study demonstrates the importance of the DOC pool when assessing the carbon sequestration potential of artificial upwelling.

Climate change is expected to alter the intensity and frequency of upwelling in high productive coastal regions, thus impacting nutrient fluxes, primary productivity and consequently carbon cycling. However, it is unknown how these changes will impact the planktonic (phytoplankton and bacteria) community structure, which affects community respiration (CR) and hence the carbon available for sequestration or transfer to upper trophic levels. Here we present results from a 37-day mesocosm experiment where we examined the response of CR to nutrient additions by simulating upwelling events at different intensities (low, medium, high and extreme) and modes (singular and recurring additions). We also analysed the potential contribution of different plankton size classes and functional groups to CR. The trend in accumulated CR with respect to nutrient fertilisation (total nitrogen added during the experiment) was linear in the two modes. Microplankton (mostly diatoms) and nanoplankton (small flagellates) dominated under extreme upwelling intensities and high CR in both singular and recurring upwelling modes, explaining >65% of the observed variability in CR. In contrast, prokaryotic picoplankton (heterotrophic bacteria and autotrophic cyanobacteria) explained <43% of the variance in CR under the rest of the upwelling intensities and modes tested. Changes in planktonic community structure, while modulating CR variability, would regulate the metabolic balance of the ecosystem, shifting it towards net-heterotrophy when the community is dominated by small heterotrophs and to net-autotrophy when large autotrophs prevail; although depending on the mode in which nutrients are supplied to the system. This shift in the dominance of planktonic organism will hence affect not only CR but also carbon sequestration in upwelling regions

Upwelling of nutrient-rich deep waters make eastern boundary upwelling systems (EBUSs), such as the Humboldt Current system, hot spots of marine productivity. Associated settling of organic matter to depth and consecutive aerobic decomposition results in large subsurface water volumes being oxygen depleted. Under these circumstances, organic matter remineralisation can continue via denitrification, which represents a major loss pathway for bioavailable nitrogen. Additionally, anaerobic ammonium oxidation can remove significant amounts of nitrogen in these areas. Here we assess the interplay of suboxic water upwelling and nitrogen cycling in a manipulative offshore mesocosm experiment. Measured denitrification rates in incubations with water from the oxygen-depleted bottom layer of the mesocosms (via 15N label incubations) mostly ranged between 5.5 and 20 nmol N2 L−1 h−1 (interquartile range), reaching up to 80 nmol N2 L−1 h−1. However, actual in situ rates in the mesocosms, estimated via Michaelis–Menten kinetic scaling, did most likely not exceed 0.2–4.2 nmol N2 L−1 h−1 (interquartile range) due to substrate limitation. In the surrounding Pacific, measured denitrification rates were similar, although indications of substrate limitation were detected only once. In contrast, anammox (anaerobic ammonium oxidation) made only a minor contribution to the overall nitrogen loss when encountered in both the mesocosms and the Pacific Ocean. This was potentially related to organic matter C / N stoichiometry and/or process-specific oxygen and hydrogen sulfide sensitivities. Over the first 38 d of the experiment, total nitrogen loss calculated from in situ rates of denitrification and anammox was comparable to estimates from a full nitrogen budget in the mesocosms and ranged between ∼ 1 and 5.5 µmol N L−1. This represents up to ∼  20 % of the initially bioavailable inorganic and organic nitrogen standing stocks. Interestingly, this loss is comparable to the total amount of particulate organic nitrogen that was exported into the sediment traps at the bottom of the mesocosms at about 20 m depth. Altogether, this suggests that a significant portion, if not the majority of nitrogen that could be exported to depth, is already lost, i.e. converted to N2 in a relatively shallow layer of the surface ocean, provided that there are oxygen-deficient conditions like those during coastal upwelling in our study. Published data for primary productivity and nitrogen loss in all EBUSs reinforce such conclusion.

The eastern North Atlantic region receives large Saharan dust deposition inputs, providing nutrients and trace metals to the surface waters. We assessed the effects of intense dust deposition on phytoplankton and bacteria cell abundances, metabolic activity, and community structure, along a surface productivity gradient in the Mauritanian-Senegalese upwelling system. Dust concentrations above 4 mg L -1 were added to triplicate microcosms in four bioassay experiments, each lasting three days, increasing nitrate, phosphate and, to a lesser extent, silicate seawater concentrations. Even though dust deposition enhanced both heterotrophic and photosynthetic activity, bacterial production responded faster and stronger than primary production, especially as oligotrophic conditions increased. Bacterial production rates in oligotrophic waters almost tripled one day after the enrichment. However, such favorable response could not be observed on the total organic carbon production until a lag phase of 2 days and whilst under moderate eutrophic conditions. Dust enrichment benefited the presence of certain planktonic groups over others according to their nutrient requirements. Indicator species analysis revealed that our dust-treated microcosms were consistently characterized by Raphid-pennate diatoms, as well as by Hyphomonas genus of Alphaproteobacteria and several species of Alteromonas Gammaproteobacteria. Yet, changes in microbial community structure and composition were primarily shaped by the starting conditions of each experiment. These findings indicate that increasing dust deposition events and the weakening of the Mauritanian-Senegalese upwelling system under climate change may result in a more heterotrophic system, particularly in oligotrophic waters, reducing its potential to function as an atmospheric carbon sink.

The subtropical oceans are home to one of the largest ecosystems on Earth, contributing to nearly one third of global oceanic primary production. Ocean warming leads to enhanced stratification in the oligotrophic ocean but also intensification in cross-shore wind gradients and thus in eddy kinetic energy across eastern boundary regions of the subtropical gyres. Phytoplankton thriving in a future warmer oligotrophic subtropical ocean with enhanced CO2 levels could therefore be patchily fertilized by increased mesoscale and submesoscale variability inducing nutrient pumping into the surface ocean. Under this premise, we have tested the response of three size classes (0.2-2, 2-20 and >20 m) of subtropical phytoplankton communities in terms of primary production, chlorophyll and cell biomass, to increasing CO2 concentrations and nutrient fertilization during an in situ mesocosm experiment in oligotrophic waters off of the island of Gran Canaria. We found no significant CO2-related effect on primary production and biomass under oligotrophic conditions (phase I). In contrast, primary production, chlorophyll and biomass displayed a significant and pronounced increase under elevated CO2 conditions in all groups after nutrient fertilization, both during the bloom (phase II) and post-bloom (phase III) conditions. Although the relative increase of primary production in picophytoplankton (250%) was 2.5 higher than in microphytoplankton (100%) after nutrient fertilization, comparing the high and low CO2 treatments, microphytoplankton dominated in terms of biomass, contributing >57% to the total. These results contrast with similar studies conducted in temperate and cold waters, where consistently small phytoplankton benefitted after nutrient additions at high CO2, pointing to different CO2–sensitivities across plankton communities and ecosystem types in the ocean.

The enzymatic electron transport system (ETS) assay is frequently used as a proxy of respiratory activity in planktonic communities. It is thought to estimate the maximum overall activity of the enzymes associated with the respiratory ETS systems in both eukaryotic and prokaryotic organisms. Thus, in order to derive actual respiration rates (R) from ETS it is necessary to determine empirical R/ETS conversion algorithms. In this study we explore the temporal development of R and ETS activity in natural plankton communities (from bacteria to large phytoplankton) enclosed in mesocosms, treated with different CO2 concentrations. The experiment lasted 30 days, during which abrupt changes in community structure and biomass occurred through a sharp transition from oligotrophy (phase I) to highly eutrophic conditions (phase II) after nutrient-induced fertilization (day 18). R and ETS did not show any response to CO2 under oligotrophic conditions, but R increased significantly more in the two high CO2 mesocosms after fertilization, coinciding with a sharp rise in large phytoplankton (mostly diatoms). R and ETS were significantly correlated only during the eutrophic phase. The R/ETS ranged more than 3 fold in magnitude during the experiment, with phase-averaged values significantly higher under oligotrophic conditions (0.7-1.1) than after nutrient fertilization (0.5-0.7). We did not find any significant relationship between R/ETS and community structure or biomass, although R correlated significantly with total biomass after fertilization in the four mesocosms. Multiple stepwise regression models show that large phytoplankton explains most of the variance in R during phases I (86%) and II (53%) and of ETS (86%) during phase II, while picophytoplankton contributes up to 73% to explain the variance in the ETS model during phase I. Our results suggest that R/ETS may be too variable in the ocean as to apply constant values to different communities living under contrasting environmental conditions. Controlled experiments with natural communities, like the present one, would help to constrain the range of variability of the R/ETS ratio, and to understand the factors driving it.

Conectamos investigación y práctica docente con el análisis del diseño de una secuencia que integra la contextualización, indagación, modelización y la evaluación de su efectividad desde las ideas del alumnado que moviliza. La metodología cualitativo-interpretativa utilizada permite analizar la práctica docente y los resultados muestran que la contextualización en un problema cercano (muerte de peces y algas en la playa) genera en los estudiantes diversidad de explicaciones sobre las posibles causas y coherencia en qué pruebas necesitarían para demostrarlas o refutarlas. La búsqueda de pruebas para detectar la auténtica causa de la muerte, en este caso por deshidratación, ofrece conocimiento descriptivo que condiciona sus explicaciones, necesarias para que el modelo de simulación diseñado de ósmosis tenga sentido para el alumnado y les sea de utilidad para transferir a otros contextos.