Description: In the autumn of 2014, nine large mesocosms were deployed in the oligotrophic subtropical North-Atlantic coastal waters off Gran Canaria (Spain). Their deployment was designed to address the acidification effects of CO2 levels from 400 to 1,400 mu atm, on a plankton community experiencing upwelling of nutrient-rich deep water. Among other parameters, chlorophyll a (chl-a), potential respiration (Phi), and biomass in terms of particulate protein (B) were measured in the microplankton community (0.7-50.0 mu m) during an oligotrophic phase (Phase I), a phytoplankton-bloom phase (Phase II), and a post-bloom phase (Phase III). Here, we explore the use of the Phi/chl-a ratio in monitoring shifts in the microplankton community composition and its metabolism. Phi/chl-a values below 2.5 mu L O-2 h(-1) (mu g chl-a)(-1) indicated a community dominated by photoautotrophs. When Phi/chl-a ranged higher, between 2.5 and 7.0 mu L O-2 h(-1) (pg chl-a)(-1) , it indicated a mixed community of phytoplankton, microzooplankton and heterotrophic prokaryotes. When Phi/chl-a rose above 7.0 mu L O-2 h(-1) (mu g chl-a)(-1), it indicated a community where microzooplankton proliferated (〉10.0 mu L O-2 h(-1) (mu g chl-a)(-1)), because heterotrophic dinoflagellates bloomed. The first derivative of B, as a function of time (dB/dt), indicates the rate of protein build-up when positive and the rate of protein loss, when negative. It revealed that the maximum increase in particulate protein (biomass) occurred between 1 and 2 days before the chl-a peak. A day after this peak, the trough revealed the maximum net biomass loss. This analysis did not detect significant changes in particulate protein, neither in Phase I nor in Phase III. Integral analysis of Phi/chl-a and B, over the duration of each phase, for each mesocosm, reflected a positive relationship between 4) and pCO(2) during Phase II [alpha = 230.10-5 mu L O-2 h(-1) L-1 (patm CO2)(-1) (phase-day)(-1), R-2 = 0.30] and between chl-a and pCO(2) during Phase III [alpha = 100.10(-5) Ag chl-a L-1 (mu atmCO(2))(-1) (phase-day)(-1), R-2 = 0.84]. At the end of Phase II, a harmful algal species (HAS), Vicicitus globosus, bloomed in the high pCO(2) mesocosms. In these mesocosms, microzooplankton did not proliferate, and chl-a retention time in the water column increased. In these V globosus-disrupted communities, the (Phi/chl-a ratio [4.1 +/- 1.5 /mu L O-2 h(-1) (mu g chl-a)(-1)] was more similar to the Phi/chl-a ratio in a mixed plankton community than to a photoautotroph-dominated one.

Description: Artificial upwelling has been discussed as a nature-based solution to fertilize currently unproductive areas of the ocean to enhance food web productivity and atmospheric CO2 sequestration. The efficacy of this approach may be closely tied to the nutrient stoichiometry of the upwelled water, as Si-rich water upwelling should benefit the growth of diatoms, who are key players for primary production, carbon export and food web efficiency. With a mesocosm experiment in subtropical waters, we assessed the physiological and functional responses of an oligotrophic phytoplankton community to artificial upwelling under varying Si:N ratios (0.07-1.33). Deep water fertilization led to strongly enhanced primary productivity rates and net autotrophy across Si scenarios. At the community level, Si-rich upwelling temporarily increased primary production and consistently enhanced diatom growth, producing up to 10-fold higher abundances compared to Si-deficient upwelling. At the organism level, contrasting effects were observed. On the one hand, silicification and size of diatom cells remained unaffected by Si:N, which is surprising given the direct dependency of these traits on Si. On the other hand, diatom Chlorophyll a density and carbon density were strongly reduced and particulate matter C:N was elevated under Si-rich upwelling. This suggests a reduced nutritional value for higher trophic levels under high Si:N ratios. Despite these strong qualitative changes under high Si, diatom cells appeared healthy and showed high photosynthetic efficiency. Our findings reveal great physiological plasticity and adaptability in phytoplankton under artificial upwelling, with Si-dependent trade-offs between primary producer quantity and quality.

Description: Ocean alkalinity enhancement (OAE) has been proposed as a carbon dioxide removal technology (CDR) allowing for long term storage of carbon dioxide in the ocean. By changing the carbonate speciation in seawater, OAE may potentially alter marine ecosystems with implications for the biological carbon pump. Using mesocosmsthe subtropical North Atlantic, we provide first empirical insights into impacts of carbonate-based OAE on the vertical flux and attenuation of sinking particles in an oligotrophic plankton community. We enhanced total alkalinity (TA) in increments of 300 μmol kg-1, reaching up to ΔTA = 2400 µmol kg-1 compared to ambient TA. We applied a pCO2-equilibrated OAE approach, i.e. dissolved inorganic carbon (DIC) was raised simultaneously with TA to maintain seawater pCO2 in equilibrium with the atmosphere, thereby keeping perturbations of seawater carbonate chemistry moderate. The vertical flux of major elements including carbon, nitrogen, phosphorus and silicon, as well as their stoichiometric ratios (e.g. carbon-to-nitrogen) remained unaffected over 29 days of OAE. The particle properties controlling the flux attenuationinking velocities and remineralization rates also remained unaffected by OAE. However, we observed abiotic mineral precipitation at high OAE levels (ΔTA = 1800 μmol kg-1 and higher) that resulted in a substantial increase in PIC formation. The associated consumption of alkalinity reduces the efficiency of CO2 removal and emphasizes the importance of maintaining OAE within a carefully defined operating range. Our findings suggest that carbon export by oligotrophic plankton communities is insensitive to OAE perturbations using a CO2 pre-equilibrated approach. The integrity of ecosystem services is a prerequisite for large-scale application and should be further tested across a variety of nutrient-regimes and for less idealized OAE approaches.

Description: Anthropogenic CO2 emissions cause a drop in seawater pH and shift the inorganic carbon speciation. Collectively, the term ocean acidification (OA) summarizes these changes. Few studies have examined OA effects on predatory plankton, e.g. Hydrozoa and fish larvae as well as their interaction in complex natural communities. Because Hydrozoa can seriously compete with and prey on other higher-level predators such as fish, changes in their abundances may have significant consequences for marine food webs and ecosystem services. To investigate the interaction between Hydrozoa and fish larvae influenced by OA, we enclosed a natural plankton community in Raunefjord, Norway, for 53 days in eight ≈ 58 m³ pelagic mesocosms. CO2 levels in four mesocosms were increased to ≈ 2000 µatm pCO2, whereas the other four served as untreated controls. We studied OA-induced changes at the top of the food web by following ≈2000 larvae of Atlantic herring (Clupea harengus) hatched inside each mesocosm during the first week of the experiment, and a Hydrozoa population that had already established inside the mesocosms. Under OA, we detected 20% higher abundance of hydromedusae staged jellyfish, but 25% lower biomass. At the same time, survival rates of Atlantic herring larvae were higher under OA (control pCO2: 0.1%, high pCO2: 1.7%) in the final phase of the study. These results indicate that a decrease in predation pressure shortly after hatch likely shaped higher herring larvae survival, when hydromedusae abundance was lower in the OA treatment compared to control conditions. We conclude that indirect food-web mediated OA effects drove the observed changes in the Hydrozoa – fish relationship, based on significant changes in the phyto-, micro-, and mesoplankton community under high pCO2. Ultimately, the observed immediate consequences of these changes for fish larvae survival and the balance of the Hydrozoa – fish larvae predator – prey relationship has important implications for the functioning of oceanic food webs.

Description: To keep global warming below 1.5 °C, technologies that remove carbon from the atmosphere will be needed. Ocean artificial upwelling of nutrient-rich water stimulates primary productivity and could enhance the biological carbon pump for natural CO2 removal. Its potential may depend on the Si availability in the upwelled water, which regulates the abundance of diatoms that are key carbon exporters. In a mesocosm experiment, we tested the effect of nutrient composition (Si relative to N) in artificially upwelled waters on export quantity and quality in a subtropical oligotrophic environment. Upwelling led to a doubling of exported particulate matter and increased C:N ratios to well beyond Redfield (9.5 to 11.1). High Si availability stimulated this carbon over-consumption further, resulting in a temporary ~5-fold increase in POC export and ~30% increase in C:N ratios compared to Si-scarce upwelling. Whilst the biogenic Si ballast of the export flux increased more than 3.5-fold over the Si:N gradient, these heavier particles did not sink faster. On the contrary, sinking velocity decreased considerably under high Si:N, most likely due to reduced particle size. Respiration rates remained similar across all treatments indicating that biogenic Si did not protect particles against microbial degradation. Si availability thus influenced key processes of the biological carbon pump in counteracting ways by increasing the export magnitude and associated C:N ratios but decreasing the efficiency of carbon transfer to depth. These opposing effects need to be considered when evaluating the potential of artificial upwelling as negative emission technology.

Description: The oceans’ uptake of anthropogenic carbon dioxide (CO2) decreases seawater pH and alters the inorganic carbon speciation – summarized in the term ocean acidification (OA). Already today, coastal regions experience episodic pH events during which surface layer pH drops below values projected for the surface ocean at the end of the century. Future OA is expected to further enhance the intensity of these coastal extreme pH events. To evaluate the influence of such episodic OA events in coastal regions, we deployed eight pelagic mesocosms for 53 days in Raunefjord, Norway, and enclosed 56–61 m3 of local seawater containing a natural plankton community under nutrient limited post-bloom conditions. Four mesocosms were enriched with CO2 to simulate extreme pCO2 levels of 1978 – 2069 μatm while the other four served as untreated controls. Here, we present results from multivariate analyses on OA-induced changes in the phyto-, micro-, and mesozooplankton community structure. Pronounced differences in the plankton community emerged early in the experiment, and were amplified by enhanced top-down control throughout the study period. The plankton groups responding most profoundly to high CO2 conditions were cyanobacteria (negative), chlorophyceae (negative), auto- and heterotrophic microzooplankton (negative), and a variety of mesozooplanktonic taxa, including copepoda (mixed), appendicularia (positive), hydrozoa (positive), fish larvae (positive), and gastropoda (negative). The restructuring of the community coincided with significant changes in the concentration and elemental stoichiometry of particulate organic matter. Results imply that extreme CO2 events can lead to a substantial reorganization of the planktonic food web, affecting multiple trophic levels from phytoplankton to primary and secondary consumers.

Description: Reduction of anthropogenic CO2 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 NO3– 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

Description: Diatoms account for up to 40% of marine primary production(1,2) and require silicic acid to grow and build their opal shell(3). On the physiological and ecological level, diatoms are thought to be resistant to, or even benefit from, ocean acidification(4-6). Yet, global-scale responses and implications for biogeochemical cycles in the future ocean remain largely unknown. Here we conducted five in situ mesocosm experiments with natural plankton communities in different biomes and find that ocean acidification increases the elemental ratio of silicon (Si) to nitrogen (N) of sinking biogenic matter by 17 +/- 6 per cent under p(CO2) conditions projected for the year 2100. This shift in Si:N seems to be caused by slower chemical dissolution of silica at decreasing seawater pH. We test this finding with global sediment trap data, which confirm a widespread influence of pH on Si:N in the oceanic water column. Earth system model simulations show that a future pH-driven decrease in silica dissolution of sinking material reduces the availability of silicic acid in the surface ocean, triggering a global decline of diatoms by 13-26 per cent due to ocean acidification by the year 2200. This outcome contrasts sharply with the conclusions of previous experimental studies, thereby illustrating how our current understanding of biological impacts of ocean change can be considerably altered at the global scale through unexpected feedback mechanisms in the Earth system.

Description: As one of Earth's most productive marine ecosystems, the Peruvian upwelling system transports large amounts of biogenic matter from the surface to the deep ocean. Whilst particle sinking velocity is a key factor controlling the biological pump, thereby affecting carbon sequestration and O2-depletion, it has not yet been measured in this system. During a 50 d mesocosm experiment in the surface waters off the coast of Peru, we assessed particle sinking velocities and their biogeochemical and physical drivers. We further characterized the general properties of exported particles under different phytoplankton communities and nutritional states. Average sinking velocities varied between size classes and ranged from 12.8 ± 0.7 m d−1 (particles 40–100 µm) to 19.4 ± 0.7 m d−1 (particles 100–250 µm) and 34.2 ± 1.5 m d−1 (particles 250–1000 µm) (± 95 % CI). Despite a distinct plankton succession from diatoms to dinoflagellates with concomitant 5-fold drop in opal ballasting, substantial changes in sinking velocity were not observed. This illustrates the complexity of counteracting factors driving the settling behaviour of marine particles. In contrast, we found higher sinking velocities with increasing particle size and roundness and decreasing porosity. Size had by far the strongest influence among these physical particle properties, despite a high amount of unexplained variability. Our study provides a detailed analysis of the drivers of particle sinking velocity in the Peruvian upwelling system, which allows modellers to optimize local particle flux parameterization. This will help to better project oxygen concentrations and carbon sequestration in a region that is subject to substantial climate-driven changes.

Description: The Humboldt Current Upwelling System (HCS) is the most productive eastern boundary upwelling system (EBUS) in terms of fishery yield on the planet. EBUSs are considered hotspots of climate change with predicted expansion of mesopelagic oxygen minimum zones (OMZs) and related changes in the frequency and intensity of upwelling of nutrient-rich, low-oxygen deep water. To increase our mechanistic understanding of how upwelling impacts plankton communities and trophic links, we investigated mesozooplankton community succession and gut fluorescence, fatty acid and elemental compositions (C, N, O, P), and stable isotope (δ13C, δ15N) ratios of dominant mesozooplankton and microzooplankton representatives in a mesocosm setup off Callao (Peru) after simulated upwelling with OMZ water from two different locations and different N:P signatures (moderate and extreme treatments). An oxycline between 5 and 15 m with hypoxic conditions (〈50 µmol L−1) below ∼10 m persisted in the mesocosms throughout the experiment. No treatment effects were determined for the measured parameters, but differences in nutrient concentrations established through OMZ water additions were only minor. Copepods and polychaete larvae dominated in terms of abundance and biomass. Development and reproduction of the dominant copepod genera Paracalanus sp., Hemicyclops sp., Acartia sp., and Oncaea sp. were hindered as evident from accumulation of adult copepodids but largely missing nauplii. Failed hatching of nauplii in the hypoxic bottom layer of the mesocosms and poor nutritional condition of copepods suggested from very low gut fluorescence and fatty acid compositions most likely explain the retarded copepod development. Correlation analysis revealed no particular trophic relations between dominant copepods and phytoplankton groups. Possibly, particulate organic matter with a relatively high C:N ratio was a major diet of copepods. C:N ratios of copepods and polychaetes ranged 4.8–5.8 and 4.2–4.3, respectively. δ15N was comparatively high (∼13 ‰–17 ‰), potentially because the injected OMZ source water was enriched in δ15N as a result of anoxic conditions. Elemental ratios of dinoflagellates deviated strongly from the Redfield ratio. We conclude that opportunistic feeding of copepods may have played an important role in the pelagic food web. Overall, projected changes in the frequency and intensity of upwelling hypoxic waters may make a huge difference for copepod reproduction and may be further enhanced by varying N:P ratios of upwelled OMZ water masses.