Ocean alkalinity enhancement (OAE) through mineral dissolution is a promising marine carbon dioxide removal strategy because it increases the buffering capacity of seawater and thereby enhances passive storage of atmospheric CO2. However, the ecological consequences of OAE for zooplankton, particularly gelatinous species, remain poorly understood. Here, we assessed the response of a key gelatinous zooplankton species to OAE in a 53-day mesocosm experiment in a temperate Norwegian fjord. Oikopleura dioica is a globally distributed zooplankton member, known for its high secondary production capacity and key role in vertical carbon flux. O. dioica continuously produces mucous feeding structures ('houses'), which efficiently retain submicron particles. Once discarded, these houses can sink rapidly and contribute to vertical carbon exports. To test the impacts of OAE on O. dioica abundances and their house production capacity, we exposed natural plankton communities to non-CO2-equilibrated OAE scenarios spanning a ΔTA range from 0-600 μmol kg-1, using silicate-based (olivine) and calcium-based (slaked lime) minerals. Population dynamics of O. dioica were monitored alongside the plankton community, and targeted bottle incubations were used to quantify house production and feeding rates. We show that O. dioica abundances varied by an order of magnitude within and across treatments. No interaction between O. dioica abundance and alkalinity levels or mineral types could be detected. Instead, O. dioica abundance variations were primarily explained by prey availability (picoplankton). Additionally, house production and feeding rate experiments showed that O. dioica were unaffected by OAE across all treatments. These findings indicate that O. dioica, as a key gelatinous zooplankton member, is physiologically resilient to OAE within the tested range. Future studies should incorporate gelatinous zooplankton into OAE assessments and investigate higher alkalinity perturbations to evaluate potential ecosystem impacts and larvacean-mediated changes in carbon export during under OAE deployments above ΔTA 600 μmol kg-1.

The Humboldt upwelling system (HUS) is known for its extraordinary productivity due to wind-driven upwelling of nutrient-rich deep water, resulting in the highest fish catches per unit area worldwide. However, contrary to other Eastern boundary upwelling systems, upwelling intensity is highest in winter, while primary productivity reaches its peak during the summer months. Our current understanding of the counterintuitive relationship between upwelling intensity and productivity is insufficient to predict the consequences of climate change on this ecosystem. To elucidate the drivers of the upwelling-productivity relationship in the HUS, we tested the hypothesis that low light availability limits primary productivity in winter despite strong upwelling intensity supplying plenty of nutrients into the surface layer, while light availability in the shallower mixed layer in summer enables an effective use of the upwelled nutrients. To disentangle the interactive effects of light and nutrients on primary production and carbon cycling, we enclosed natural plankton communities off the coast of Callao (Peru) during a 35-day mesocosm experiment by recreating summer-time high light and winter-time low light conditions under different upwelling intensities (0 %, 15 %, 30 %, 45 % and 60 %). Primary productivity and phytoplankton biomass scaled with both nitrate and light availability. Comparing the same upwelling intensity at different light levels, our data confirmed the hypothesis that light limitation due to deepening of the mixed layer is a key driver for the out-of-phase observations in primary productivity in the Humboldt upwelling system. Under light limiting conditions phytoplankton had low POC:Chla ratios indicating photoacclimation and low POC:PON ratios indicating light limitation of nitrate uptake which leads to increased food quality for grazers in winter. Our study indicates that small seasonal changes in phytoplankton biomass (estimated using Chla) might hide larger changes in primary productivity (carbon uptake), and highlights the importance of combining satellite studies with in situ observations and experimental studies to predict the fate of upwelling systems in a changing ocean. Increased stratification caused by global warming in upwelling systems such as the HUS would lead to less phytoplankton biomass with higher POC:Chla and POC:PON ratios. This phytoplankton community would have lower food quality for grazers and might lead to a decline in the transfer to higher trophic levels, but at the same time might lead to increased CO2 drawdown in an otherwise CO2 emitting ecosystem. Understanding the unique relationship between upwelling intensity and productivity in the HUS contributes to predicting the reaction of this valuable ecosystem for fisheries to the impacts of climate change.

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 mesocosms in the 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 ratios), remained unaffected over 29 d of OAE. The particle properties controlling the flux attenuation, including sinking 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 particulate inorganic carbon (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.

Ocean alkalinity enhancement (OAE) is considered for the long-term removal of gigatonnes of carbon dioxide (CO2) from the atmosphere to achieve our climate goals. Little is known, however, about the ecosystem-level changes in biogeochemical functioning that may result from the chemical sequestration of CO2 in seawater and how stable the sequestration is. We studied these two aspects in natural plankton communities under carbonate-based, CO2-equilibrated OAE of up to a doubling of ambient alkalinity (+2400 µeq kg−1, Ωaragonite∼10) in the nutrient-poor North Atlantic. During our month-long mesocosm experiment, the majority of biogeochemical pools, including inorganic nutrients, particulate organic carbon and phosphorus, and biogenic silica, remained unaltered across all OAE levels. Noticeable exceptions were a minor decrease in particulate organic nitrogen and an increase in the carbon-to-nitrogen ratio (C:N) of particulate organic matter in response to OAE. Thus, in our nitrogen-limited system, nitrogen turnover processes appear more susceptible than those of other elements, which could lead to decreased food quality and increased organic carbon storage. However, alkalinity and chemical CO2 sequestration were not stable at all levels of OAE. Two weeks after alkalinity addition, we measured a loss of added alkalinity and of the initially stored CO2 in the mesocosm where alkalinity was highest. The loss rate in this mesocosm accelerated over time and amounted to ∼10 % of stored CO2 within 4 weeks after alkalinity enhancement. Additional tests showed that such secondary precipitation can be initiated by particles acting as precipitation nuclei and that this process can occur even at lower levels of OAE. In conclusion, in scenarios like our study with carbonate-based OAE, where the carbon is already sequestered, the risk of major and sustained impacts on biogeochemical functioning may be low in the nutrient-poor ocean. However, the durability of carbon sequestration using OAE could be constrained by alkalinity loss in supersaturated waters with precipitation nuclei present. Our study provides an evaluation of the ecosystem impacts of an idealised OAE deployment for monitoring, reporting, and verification in an oligotrophic system. Whether biogeochemical functioning is resilient to more technically simple and economically viable approaches that induce stronger water chemistry perturbations remains to be seen.

Abstract. Ocean alkalinity enhancement (OAE) stands as a promising carbon dioxide removal technology. Yet, this solution to climate change entails shifts in water chemistry with unknown consequences for marine fish that are critical to ecosystem health and food security. With a laboratory and mesocosm experiment, we show that early life stages of fish can be resistant to OAE. We examined metabolic rate, swimming behavior, growth and survival in Atlantic herring (Clupea harengus) and other temperate coastal fish species. Neither direct physiological nor indirect food web-mediated impacts of OAE were apparent. This was despite non-CO2-equilibrated OAE (ΔTA = +600 µmol kg-1) that induces strong perturbations (ΔpH = +0.7, pCO2 = 75 µatm) compared to alternative deployment scenarios. Whilst our results give cause for optimism regarding the large-scale application of OAE, other life history stages (embryos) and habitats (open ocean) may prove more vulnerable. Still, our study across ecological scales (organism to community) and exposure times (short- to long-term) suggests that some fish populations, including key fisheries species, may be resilient to the carbonate chemistry changes under OAE.

Enhancing ocean productivity by artificial upwelling is evaluated as a nature-based solution for food security and climate change mitigation. Fish production is intended through diatom-based plankton food webs as these are assumed to be short and efficient. However, our findings from mesocosm experiments on artificial upwelling in the oligotrophic ocean disagree with this classical food web model. Here, diatoms did not reduce trophic length and instead impaired the transfer of primary production to crustacean grazers and small pelagic fish. The diatom-driven decrease in trophic efficiency was likely mediated by changes in nutritional value for the copepod grazers. Whilst diatoms benefitted the availability of essential fatty acids, they also caused unfavorable elemental compositions via high carbon-to-nitrogen ratios (i.e. low protein content) to which the grazers were unable to adapt. This nutritional imbalance for grazers was most pronounced in systems optimized for CO2 uptake through carbon-to-nitrogen ratios well beyond Redfield. A simultaneous enhancement of fisheries production and carbon sequestration via artificial upwelling may thus be difficult to achieve given their opposing stoichiometric constraints. Our study suggest that food quality can be more critical than quantity to maximize food web productivity during shorter-term fertilization of the oligotrophic ocean.

IntroductionArtificial 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 upwelling should benefit the growth of diatoms, who are key players for primary production, carbon export and food web efficiency.MethodsWith 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).ResultsDeep water fertilization led to strongly enhanced primary productivity rates and net autotrophy across Si scenarios. At the community level, Si-rich upwelling50 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.DiscussionThis 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.

Ocean alkalinity enhancement (OAE) is currently discussed as a potential negative emission technology to sequester atmospheric carbon dioxide in seawater. Yet, its potential risks or cobenefits for marine ecosystems are still mostly unknown, thus hampering its evaluation for large-scale application. Here, we assessed the impacts OAE may have on plankton communities, focusing on phytoplankton and microzooplankton. In a mesocosm study in the oligotrophic subtropical North Atlantic, we investigated the response of a natural plankton community to CO2-equilibrated OAE across a gradient from ambient alkalinity (2400 μmol kg-1) to double (4800 μmol kg-1). Abundance and biomass of phytoplankton and microzooplankton were insensitive to OAE across all size classes (pico, nano and micro), nutritional modes (autotrophic, mixotrophic and heterotrophic) and taxonomic groups (cyanobacteria, diatoms, haptophytes, dinoflagellates, and ciliates). Consequently, plankton communities under OAE maintained their natural chlorophyll a levels, size structure, taxonomic composition and biodiversity. These findings suggest a high tolerance of phytoplankton and microzooplankton to CO2-equilibrated OAE in the oligotrophic ocean. However, alternative application schemes involving more drastic perturbations in water chemistry and nutrient-rich ecosystems require further investigation. Nevertheless, our study on idealized OAE will help develop an environmentally safe operating space for this climate change mitigation solution.

Ocean alkalinity enhancement (OAE) stands as a promising carbon dioxide removal technology. Yet, this solution to climate change entails shifts in environmental drivers with unknown consequences for marine fish that are critical to ecosystem health and food security. Fish and their supporting food webs may be stressed by the novel carbonate chemistry or the nutrients contained in the deployed minerals. With a mesocosm experiment on natural plankton communities, we studied early life stages of fish under alkalinity (+600 µmol kg−1) and silicate (+75 µmol L−1) addition. Larvae and young juveniles of temperate coastal species, including Atlantic herring (Clupea harengus) and cod (Gadus morhua), were exposed to direct physiological and indirect food-web-mediated effects of OAE for 49 d. Neither in the shorter nor in the longer term did we find an impairment of fish growth and survival. Alkalization even led to an increase in fish biomass. This resistance to OAE was despite using non-CO2-equilibrated deployment that induces more severe perturbations in carbonate chemistry (ΔpH =+0.7, pCO2=75 µatm) compared to alternative scenarios. Overall, our community-level study suggests that some fish populations, including key fisheries' species, may be resilient to the water chemistry changes under OAE. Whilst these results give cause for optimism regarding the large-scale application of OAE, other life history stages (embryos) and habitats (open ocean) may prove more vulnerable.

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 CO 2 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.

Ocean artificial upwelling has been suggested to boost primary production and increase harvestable resources such as fish. Yet, for this ecosystem-based approach to work, an effective energy transfer up the food web is required. Here, we studied the trophic role of microzooplankton under artificial upwelling via biomass and community composition as well as grazing rates on phytoplankton. Using mesocosms in the oligotrophic ocean, we supplied nutrient-rich deep water at varying intensities (low to high) and addition modes (a Singular large pulse or smaller Recurring pulses). Deep-water fertilization created a diatom-dominated bloom that scaled with the amount of inorganic nutrients added, but also Synechococcus-like cells, picoeukaryotes and nanophytoplankton increased in abundance with added nutrients. After 30 days, towards the end of the experiment, coccolithophores bloomed under recurring upwelling of high intensity. Across all upwelling scenarios, the microzooplankton community was dominated by ciliates, dinoflagellates (mixo- and heterotrophic) and radiolarians. Under the highest upwelling intensity, the average grazing rates of Synechococcus-like cells, picoeukaryotes and nanophytoplankton by microzooplankton were 0.35 d-1 ± 0.18 (SD), 0.09 d-1 ± 0.12 (SD), and 0.11 d-1 ± 0.13 (SD), respectively. There was little temporal variation in grazing of nanophytoplankton but grazing of Synechococcus-like cells and picoeukaryotes were more variable. There were positive correlations between abundance of these groups and grazing rates, suggesting a response in the microzooplankton community to prey availability. The average phytoplankton to microzooplankton ratio (biovolume) increased with added deep water, and this increase was highest in the Singular treatment, reaching ~30 (m3 m-3), whereas the phytoplankton to total zooplankton biomass ratio (weight) increased from ~1 under low upwelling to ~6 (g g-1) in the highest upwelling but without a difference between the Singular and the Recurring mode. Several smaller, recurring upwelling events increased the importance of microzooplankton compared with one large pulse of deep water. Our results demonstrate that microzooplankton would be an important component for trophic transfer if artificial upwelling would be carried out at scale in the oligotrophic ocean.

<p>In light of the climate crisis and the necessity to meet the Paris Agreement goal of staying well below the 2°C temperature increase whilst respecting other UN sustainable development goals, an array of technologies to absorb and store atmospheric , named Carbon Dioxide Removal (CDR) technologies, are being proposed, developed and researched. One such technologies is Ocean Alkalinization, or Ocean Alkalinity Enhancement (OAE), which stems from the natural process of rock weathering. Several alkalinity sources and deployment options have been proposed, each associated with differing biological drivers. Among these, a carbonate-based, dissolved, equilibrated addition stands as the most optimistic deployment scenario, both from a carbon sequestration verification and an ecosystem impact stand point.</p><p>We decided to test this implementation in the first community-level mesocosm experiment to be done in the field. Pelagic mesocosms were deployed in Taliarte (Gran Canaria), enclosing 8 m³ of oligotrophic coastal waters with their associated natural plankton community. Nine OAE addition scenarios were simulated, increasing alkalinity (TA) in steps of 300 µeq/L from ambient levels up to its doubling, using a mixture of sodium carbonate and bicarbonate. Particular focus was placed on the impacts of enhanced TA on the ecosystem service (ES) of food production. This was addressed via zooplankton properties pertaining to ecosystem stability, food quantity and nutritional quality. Zooplankton diversity, functional composition, biomass, CN stoichiometry, population size structure, secondary production, trophic length, and a number of fatty acid nutritional indexes and trophic markers were monitored throughout the 33-day experiment.</p><p>Here, only 4 out of the over 30 different stability, food quantity and quality proxies were significantly affected by enhanced TA.<em> </em>Out of these, two were interpreted as negative impacts: a shorter-term halving in small copepod production, coinciding with a halving in copepod nauplii biomass, with a doubling in TA. These responses could be partly explained by the halving in large microplankton, an assumed preferred food source for copepods, detected right after treatment. However, none of these were sustained until the end of the experiment, thus suggesting no longer-term consequences. All in all, this study provides evidence for a low impact risk of enhanced TA on zooplankton, and ultimately the ES of food production. These findings set a promising stage towards the safe implementation of CO2-equilibrated OAE in oligotrophic coastal waters.</p>

Artificial upwelling brings nutrient-rich deep water to the sun-lit surface to boost fisheries or carbon sequestration. Deep water sources under consideration range widely in inorganic silicon (Si) relative to nitrogen (N). Yet, little is known about how such differences in nutrient composition may influence the effectiveness of the fertilization. Si is essential primarily for diatoms that may increase food web and export efficiency via their large size and ballasting mineral shells, respectively. With a month-long mesocosm study in the subtropical North Atlantic, we tested the biological response to artificial upwelling with varying Si:N ratios (0.07-1.33). Community biomass increased 10-fold across all mesocosms, indicating that basic bloom dynamics were upheld despite the wide range in nutrient composition. Key properties of these blooms, however, were influenced by Si. Photosynthetic capacity and nutrient-use efficiency doubled from Si-poor to Si-rich upwelling, leading to C:N ratios as high as 17, well beyond Redfield. Si-rich upwelling also resulted in 6-fold higher diatom abundance and mineralized Si and a corresponding shift from smaller towards larger phytoplankton. The pronounced change in both plankton quantity (biomass) and quality (C:N ratio, size and mineral ballast) for trophic transfer and export underlines the pivotal role of Si in shaping the response of oligotrophic regions to upwelled nutrients. Our findings indicate a benefit of active Si management during artificial upwelling with the potential to optimize fisheries production and CO2 removal.