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Ocean alkalinity enhancement (OAE) aims to transfer carbon dioxide (CO 2) from the atmosphere to the ocean by increasing the capacity of seawater to store CO 2. The potential effects of OAE-induced changes in seawater chemistry on marine biology must be assessed to understand if OAE, operated at a climate-relevant scale, would be environmentally sustainable. Here, we describe the design of the Ocean Alkalinity Enhancement Pelagic Impact Intercomparison Project (OAEPIIP) – a standardised OAE microcosm experiment with plankton communities to be conducted worldwide. OAEPIIP provides funding for participating laboratories to conduct OAE experiments in their local environments. This paper constitutes a detailed manual on the standardised methodology that shall be adopted by all OAEPIIP participants. The individual studies will provide new insights into how plankton communities respond to OAE. The synthesis of these standardised studies, without publication bias, will reveal common OAE-responses that occur across geographic and environmental gradients and are therefore particularly important to determine. The funding available to OAEPIIP and the resulting data will be shared to maximise their value and accessibility. The globally coordinated effort has potential to promote scientific consensus about the potential effects of OAE on diverse plankton communities. Such consensus, through inclusion of the global community, will provide a sounder base to facilitate political decision making as to whether OAE should be scaled up or not. [ABSTRACT FROM AUTHOR]

Understanding the impact that mineral grain size and seawater salinity have on magnesium hydroxide (Mg(OH)2) dissolution and secondary calcium carbonate (CaCO3) precipitation is critical for the success of ocean alkalinity enhancement. We tested the Mg(OH)2 dissolution kinetics in seawater using three Mg(OH)2 grain sizes (<63, 63–180 and >180 µm) and at three salinities (~36, ~28 and ~20). While Mg(OH)2 dissolution occurred quicker the smaller the grain size, salinity did not significantly impact measured rates. Our results also demonstrate that grain size can impact secondary CaCO3 precipitation, suggesting that an optimum grain size exists for ocean alkalinity enhancement (OAE) using solid Mg(OH)2. Of the three grain sizes tested, the medium grain size (63–180 µm) was optimal in terms of delaying secondary CaCO3 precipitation. We hypothesize that in the lowest grain size experiments, the higher surface area provided numerous CaCO3 precipitation nuclei, while the slower dissolution of bigger grain size maintained a higher alkalinity/pH at the surface of particles, increasing CaCO3 precipitation rates and making it observable much quicker than for the intermediate grain size. Salinity also played a role in CaCO3 precipitation where the decrease in magnesium (Mg) allowed for secondary precipitation to occur more quickly, similar in effect size to another known inhibitor, i.e., dissolved organic carbon (DOC). In summary, our results suggest that OAE efficiency as influenced by CaCO3 precipitation not only depends on seawater composition but also on the physical properties of the alkaline feedstock used. [ABSTRACT FROM AUTHOR]

Gelatinous zooplankton are increasingly recognized to play a key role in the ocean's biological carbon pump. Appendicularians, a class of pelagic tunicates, are among the most abundant gelatinous plankton in the ocean, but it is an open question how their contribution to carbon export might change in the future. Here, we conducted an experiment with large volume in situ mesocosms (~55–60 m3 and 21 m depth) to investigate how ocean acidification (OA) extreme events affect food web structure and carbon export in a natural plankton community, particularly focusing on the keystone species Oikopleura dioica, a globally abundant appendicularian. We found a profound influence of O. dioica on vertical carbon fluxes, particularly during a short but intense bloom period in the high CO2 treatment, during which carbon export was 42%–64% higher than under ambient conditions. This elevated flux was mostly driven by an almost twofold increase in O. dioica biomass under high CO2. This rapid population increase was linked to enhanced fecundity (+20%) that likely resulted from physiological benefits of low pH conditions. The resulting competitive advantage of O. dioica resulted in enhanced grazing on phytoplankton and transfer of this consumed biomass into sinking particles. Using a simple carbon flux model for O. dioica, we estimate that high CO2 doubled the carbon flux of discarded mucous houses and fecal pellets, accounting for up to 39% of total carbon export from the ecosystem during the bloom. Considering the wide geographic distribution of O. dioica, our findings suggest that appendicularians may become an increasingly important vector of carbon export with ongoing OA. [ABSTRACT FROM AUTHOR]

Ocean alkalinity enhancement (OAE) is an emerging approach for atmospheric carbon dioxide removal (CDR). The net climatic benefit of OAE depends on how much it can increase CO 2 sequestration relative to a baseline state without OAE. This so-called "additionality" can be calculated as follows: Additionality = C OAE - Δ C baseline. So far, feasibility studies on OAE have mainly focussed on enhancing alkalinity in the oceans to stimulate CO 2 sequestration (COAE) ; however, the primary focus has not been on how such anthropogenic alkalinity would modify the natural alkalinity cycle and associated baseline CO 2 sequestration (ΔCbaseline). Here, I present incubation experiments in which materials considered for OAE (sodium hydroxide, steel slag, and olivine) are exposed to beach sand to investigate the influence of anthropogenic alkalinity on natural alkalinity sources and sinks. The experiments show that anthropogenic alkalinity can strongly reduce the generation of natural alkalinity, thereby reducing additionality. This is because the anthropogenic alkalinity increases the calcium carbonate saturation state, which reduces the dissolution of calcium carbonate from sand, a natural alkalinity source. I argue that this "additionality problem" of OAE is potentially widespread and applies to many marine systems where OAE implementation is considered – far beyond the beach scenario investigated in this study. However, the problem can potentially be mitigated by dilute dosing of anthropogenic alkalinity into the ocean environment and the avoidance of OAE in natural alkalinity cycling hotspots, such as in marine sediments. Understanding a potential slowdown of the natural alkalinity cycle through the introduction of an anthropogenic alkalinity cycle will be crucial for the assessment of OAE. [ABSTRACT FROM AUTHOR]

Ocean alkalinity enhancement (OAE) is a proposed method to counteract climate change by increasing the alkalinity of the surface ocean and thus the chemical storage capacity of seawater for atmospheric CO2. The impact of OAE on marine ecosystems, including phytoplankton communities which make up the base of the marine food web, are largely unknown. To investigate the influence of OAE on phytoplankton communities we enclosed a natural plankton community from coastal Tasmania for 22 days in nine microcosms during a spring bloom. Microcosms were split into three groups, (1) the unperturbed control, (2) the unequilibrated treatment where alkalinity was increased (+495 ± 5.2 µmol/kg) but seawater CO2 was not in equilibrium with atmospheric CO2, and (3) the equilibrated treatment where alkalinity was increased (+500 ± 3.2 µmol/kg) and seawater CO2 was in equilibrium with atmospheric CO2. Both treatments have the capacity to increase the inorganic carbon sink of seawater by 21 %. We found that simulated OAE had significant but generally moderate effects on various groups in the phytoplankton community and on heterotrophic bacteria. More pronounced effects were observed for the diatom community where silicic acid draw-down and biogenic silica build-up were reduced at increased alkalinity. Observed changes in phytoplankton communities affected the temporal trends of key biogeochemical parameters such as the organic matter carbon-to-nitrogen ratio. Interestingly, the unequilibrated treatment did not have a noticeably larger impact on the phytoplankton (and heterotrophic bacteria) community than the equilibrated treatment, even though the changes in carbonate chemistry conditions were much more severe. This was particularly evident from the occurrence and peak of the phytoplankton spring bloom during the experiment, which was not noticeably different from the control. Altogether, the inadvertent effects of increased alkalinity on the coastal phytoplankton communities appear to be justifiable, relative to the enormous climatic benefit of increasing the inorganic carbon sink of seawater by 21 %. [ABSTRACT FROM AUTHOR]

Ocean alkalinity enhancement (OAE) is a proposed method for removing carbon dioxide (CO2) from the atmosphere by the accelerated weathering of (ultra-) basic minerals to increase alkalinity - the chemical capacity of seawater to store CO2. During the weathering of OAE-relevant minerals relatively large amounts of trace metals will be released and may perturb pelagic ecosystems. Nickel (Ni) is of particular concern as it is abundant in olivine, one of the most widely considered minerals for OAE. However, so far there is limited knowledge about the impact of Ni on marine biota including phytoplankton. To fill this knowledge gap, this study tested the growth and photo-physiological response of 11 marine phytoplankton species to a wide range of dissolved Ni concentrations (from 0 nmol/L to 50,000 nmol/L). We found that the phytoplankton species were not very sensitive to Ni concentrations under the culturing conditions established in our experiments, but the responses were species-specific. The growth rates of 6 of the 11 tested species showed small but significant responses to changing Ni concentrations. Photosynthetic performance, assessed by measuring the maximum quantum yield (Fv/Fm) and the functional absorption cross-section (σPSII) of photosystem II, was also only mildly sensitive to changing Ni in 3 out of 11 species and 4 out of 11 species, respectively. The limited effect of Ni may be partly due to the provision of nitrate as the nitrogen source for growth, as previous studies suggest higher sensitivities when urea is the nitrogen source. Furthermore, limited influence may be due to the relatively high concentrations of organic ligands in the growth media in our experiments. These ligands reduced bioavailable Ni (i.e., "free Ni2+") concentrations by binding the majority of the dissolved Ni. Our data suggest that dissolved Ni does not have a strong effect on phytoplankton under our experimental conditions, but we emphasize that a deeper understanding of nitrogen sources, ligand concentrations and phytoplankton composition is needed when assessing the influence of Ni release associated with OAE. We discuss if applications of OAE with Ni-rich minerals may be safer in regions with high organic ligand concentrations and low concentrations of urea as such boundary conditions may lead to less impact of Ni on phytoplankton communities. [ABSTRACT FROM AUTHOR]

Eastern boundary upwelling systems (EBUS) are among the most productive marine ecosystems on Earth. The high productivity in surface waters is facilitated by upwelling of nutrient-rich deep waters, with high light availability enabling fast phytoplankton growth and nutrient utilization. However, there are numerous biotic and abiotic factors modifying productivity and biogeochemical processes. Determining these factors is important because EBUS are considered hotspots of climate change, and reliable predictions on their future functioning requires understanding of the mechanisms driving biogeochemical cycles therein. In this study, we used in situ mesocosms to obtain mechanistic understanding of processes controlling productivity, organic matter export, and particulate matter stoichiometry in the coastal Peruvian upwelling system. Therefore, eight mesocosm units with a volume of ~50m³ were deployed for 50 days ~6km off Callao during austral summer 2017, coinciding with a coastal El Niño event. To compare how upwelling of different water bodies influences plankton succession patterns, we collected two subsurface waters at different locations in the regional oxygen minimum zone (OMZ) and injected these into four replicate mesocosms, respectively (mixing ratio ≈ 1.5:1 mesocosm: OMZ water). The differences in nutrient concentrations between the collected water bodies were relatively small, and therefore we do not consider treatment differences in the present paper. The phytoplankton communities were initially dominated by diatoms but shifted towards a pronounced dominance of the mixotrophic harmful dinoflagellate (Akashiwo sanguinea) when inorganic nitrogen was exhausted in surface layers. The community shift resulted in a major short-term increase in productivity during A. sanguinea growth which left a pronounced imprint on organic matter C:N:P stoichiometry. However, C, N, and P export fluxes were not affected by this ecological regime shift because A. sanguinea persisted in the water column and did not sink out during the experiment. Accordingly, ongoing export fluxes during the study were maintained mainly by a remaining "background" plankton community. Overall, biogeochemical pools and fluxes were surprisingly constant in between the ecological regime shifts. We explain this constancy by light limitation through self-shading by phytoplankton and inorganic nitrogen limitation which constrained phytoplankton growth. Thus, gain and loss processes seemed to be relatively well balanced and there was little opportunity for blooms, which represents an event where the system becomes unbalanced. The mesocosm study revealed key links between ecological and biogeochemical processes for one of the economically most important regions in the oceans. [ABSTRACT FROM AUTHOR]

Morphological changes in coccoliths, tiny calcite platelets covering the outer surface of coccolithophores, can be the result of physiological responses to environmental changes. Coccoliths recovered from sedimentary successions may therefore provide information on paleo-environmental conditions prevailing at the time when the coccolithophores were alive. To calibrate the biomineralization responses of ancient coccolithophore to climatic changes studies often compared the biological responses of living coccolithophore species with paleo-data from calcareous nannofossils. However, there is uncertainty whether the morphological responses of living coccolithophores are representative for those of the fossilized ancestors. To investigate this, we cultured four living coccolithophore species (Emiliania huxleyi, Gephyrocapsa oceanica, Coccolithus pelagicus subsp. braarudii, and Pleurochrysis carterae) that have been evolutionarily distinct for millions of years, exposed them to changing environmental conditions (i.e. changing light intensity, Mg / Ca ratio, nutrient availability, temperature and carbonate chemistry) and evaluated their responses in coccolith morphology (i.e. size, length, width, malformation). The motivation for this study was that if the species show the same morphological response to changes in any of the tested abiotic environmental factors, then there is a reason to assume that this response is conserved over geological timescales and that coccolith morphology can serve as a paleo-proxy for that specific factor. In contrast with this concept, we found that the four species responded differently to changing light intensity, Mg / Ca ratio, nutrient availability and temperature in terms of coccolith morphology. The lack of a common response reveals the difficulties in using coccolith morphology as a proxy for paleo-environmental conditions. However, a common response was observed under changing seawater carbonate chemistry (i.e. rising CO2) which consistently induced malformations. This commonality provides some confidence that malformations found in the sedimentary record could be indicative for high CO2 levels. [ABSTRACT FROM AUTHOR]

Studies investigating the effect of increasing CO2 levels on the phosphorus cycle in natural waters are lacking although phosphorus often controls phytoplankton development in many aquatic systems. The aim of our study was to analyse effects of elevated CO2 levels on phosphorus pool sizes and uptake. The phosphorus dynamic was followed in a CO2-manipulation mesocosm experiment in the Storfjärden (western Gulf of Finland, Baltic Sea) in summer 2012 and was also studied in the surrounding fjord water. In all mesocosms as well as in surface waters of Storfjärden, dissolved organic phosphorus (DOP) concentrations of 0.26±0.03 and 0.23±0.04 μmol L-1, respectively, formed the main fraction of the total P-pool (TP), whereas phosphate (PO4) constituted the lowest fraction with mean concentration of 0.15±0.02 in the mesocosms and 0.17±0.07 μmol L-1 in the fjord. Transformation of PO4 into DOP appeared to be the main pathway of PO4 turnover. About 82% of PO4 was converted into DOP whereby only 18% of PO4 was transformed into particulate phosphorus (PP). PO4 uptake rates measured in the mesocosms ranged between 0.6 and 3.9 nmol L-1 h-1. About 86% of them was realized by the size fraction < 3 μm. Adenosine triphosphate (ATP) uptake revealed that additional P was supplied from organic compounds accounting for 25-27%of P provided by PO4 only. CO2 additions did not cause significant changes in phosphorus (P) pool sizes, DOP composition, and uptake of PO4 and ATP when the whole study period was taken into account. However, significant short-term effects were observed for PO4 and PP pool sizes in CO2 treatments > 1000 μatm during periods when phytoplankton biomass increased. In addition, we found significant relationships (e.g., between PP and Chl a/ in the untreated mesocosms which were not observed under high fCO2 conditions. Consequently, it can be hypothesized that the relationship between PP formation and phytoplankton growth changed with CO2 elevation. It can be deduced from the results, that visible effects of CO2 on P pools are coupled to phytoplankton growth when the transformation of PO4 into POP was stimulated. The transformation of PO4 into DOP on the other hand does not seem to be affected. Additionally, there were some indications that cellular mechanisms of P regulation might be modified under CO2 elevation changing the relationship between cellular constituents. [ABSTRACT FROM AUTHOR]

Gelatinous zooplankton are increasingly recognized to play a key role in the ocean's biological carbon pump. Appendicularians, a class of pelagic tunicates, are among the most abundant gelatinous plankton in the ocean, but it is an open question how their contribution to carbon export might change in the future. Here, we conducted an experiment with large volume in situ mesocosms (~55-60 m 3 and 21 m depth) to investigate how ocean acidification (OA) extreme events affect food web structure and carbon export in a natural plankton community, particularly focusing on the keystone species Oikopleura dioica, a globally abundant appendicularian. We found a profound influence of O. dioica on vertical carbon fluxes, particularly during a short but intense bloom period in the high CO 2 treatment, during which carbon export was 42%-64% higher than under ambient conditions. This elevated flux was mostly driven by an almost twofold increase in O. dioica biomass under high CO 2 . This rapid population increase was linked to enhanced fecundity (+20%) that likely resulted from physiological benefits of low pH conditions. The resulting competitive advantage of O. dioica resulted in enhanced grazing on phytoplankton and transfer of this consumed biomass into sinking particles. Using a simple carbon flux model for O. dioica, we estimate that high CO 2 doubled the carbon flux of discarded mucous houses and fecal pellets, accounting for up to 39% of total carbon export from the ecosystem during the bloom. Considering the wide geographic distribution of O. dioica, our findings suggest that appendicularians may become an increasingly important vector of carbon export with ongoing OA., (© 2023 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)