ALBERT

Description: Enrichment of the oceans with CO2 may be beneficial for some marine phytoplankton, including harmful algae. Numerous laboratory experiments provided valuable insights into the effects of elevated pCO(2) on the growth and physiology of harmful algal species, including the production of phycotoxins. Experiments close to natural conditions are the next step to improve predictions, as they consider the complex interplay between biotic and abiotic factors that can confound the direct effects of ocean acidification. We therefore investigated the effect of ocean acidification on the occurrence and abundance of phycotoxins in bulk plankton samples during a long-term mesocosm experiment in the Gullmar Fjord, Sweden, an area frequently experiencing harmful algal blooms. During the experimental period, a total of seven phycotoxin-producing harmful algal genera were identified in the fjord, and in accordance, six toxin classes were detected. However, within the mesocosms, only domoic acid and the corresponding producer Pseudo-nitzschia spp. was observed. Despite high variation within treatments, significantly higher particulate domoic acid contents were measured in the mesocosms with elevated pCO(2). Higher particulate domoic acid contents were additionally associated with macronutrient limitation. The risks associated with potentially higher phycotoxin levels in the future ocean warrants attention and should be considered in prospective monitoring strategies for coastal marine waters.

Description: The extracellular concentration of H2O2 in surface aquatic environments is controlled by a balance between photochemical production and the microbial synthesis of catalase and peroxidase enzymes to remove H2O2 from solution. In any kind of incubation experiment, the formation rates and equilibrium concentrations of reactive oxygen species (ROSs) such as H2O2 may be sensitive to both the experiment design, particularly to the regulation of incident light, and the abundance of different microbial groups, as both cellular H2O2 production and catalase–peroxidase enzyme production rates differ between species. Whilst there are extensive measurements of photochemical H2O2 formation rates and the distribution of H2O2 in the marine environment, it is poorly constrained how different microbial groups affect extracellular H2O2 concentrations, how comparable extracellular H2O2 concentrations within large-scale incubation experiments are to those observed in the surface-mixed layer, and to what extent a mismatch with environmentally relevant concentrations of ROS in incubations could influence biological processes differently to what would be observed in nature. Here we show that both experiment design and bacterial abundance consistently exert control on extracellular H2O2 concentrations across a range of incubation experiments in diverse marine environments. During four large-scale (〉1000 L) mesocosm experiments (in Gran Canaria, the Mediterranean, Patagonia and Svalbard) most experimental factors appeared to exert only minor, or no, direct effect on H2O2 concentrations. For example, in three of four experiments where pH was manipulated to 0.4–0.5 below ambient pH, no significant change was evident in extracellular H2O2 concentrations relative to controls. An influence was sometimes inferred from zooplankton density, but not consistently between different incubation experiments, and no change in H2O2 was evident in controlled experiments using different densities of the copepod Calanus finmarchicus grazing on the diatom Skeletonema costatum (〈1 % change in [H2O2] comparing copepod densities from 1 to 10 L−1). Instead, the changes in H2O2 concentration contrasting high- and low-zooplankton incubations appeared to arise from the resulting changes in bacterial activity. The correlation between bacterial abundance and extracellular H2O2 was stronger in some incubations than others (R2 range 0.09 to 0.55), yet high bacterial densities were consistently associated with low H2O2. Nonetheless, the main control on H2O2 concentrations during incubation experiments relative to those in ambient, unenclosed waters was the regulation of incident light. In an open (lidless) mesocosm experiment in Gran Canaria, H2O2 was persistently elevated (2–6-fold) above ambient concentrations; whereas using closed high-density polyethylene mesocosms in Crete, Svalbard and Patagonia H2O2 within incubations was always reduced (median 10 %–90 %) relative to ambient waters.

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: Morphological changes in coccoliths, tiny calcite platelets covering the outer surface of coccolithophores, can be induced by 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 environmental 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 of those of the fossilized ancestors. To investigate this, we exposed four living coccolithophore species (Emiliania huxleyi, Gephyrocapsa oceanica, Coccolithus pelagicus subsp. braarudii, and Pleurochrysis carterae) that have been evolutionarily distinct for hundreds of thousands to millions of years, to a range of 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 to test if there is a consistent morphological response of the four species to changes in any of the tested abiotic environmental factors. If this was the case, then this could suggest that coccolith morphology can serve as a paleo-proxy for that specific factor because this response is conserved across species that have been evolutionary distinct over geological timescales. However, 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 paleo-proxy for these environmental drivers. 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 of adverse carbonate chemistry conditions.

Description: Coccolithophores are an important group of marine phytoplankton which cover themselves with the coccosphere – a shell composed of numerous calcium carbonate (CaCO3) platelets. Despite more than a century of coccolithophore research, it remains speculative why coccolithophores calcify. Resolving this question is essential to assess the competitive fitness of coccolithophores in the future ocean where changes in calcification are expected. Here, we used the Emiliania huxleyi – Emiliania huxleyi Virus 86 host-virus model system to test the hypothesis that the coccosphere serves as a physical barrier reducing viral infection. Therefore, we removed the coccosphere from living E. huxleyi cells and compared the infection progress relative to calcified cells in a series of 6 experiments under different growth conditions. We found that the coccosphere does not constitute an effective physical barrier against viral penetration, since non-growing calcified cells were susceptible to viral infection and lysis (growth stopped by light limitation). However, we also found that protection against the virus may depend on the daily growth cycle. E. huxleyi reached higher peak abundances when decalcified cells were allowed to rebuild their coccosphere before entering cell division phase and being exposed to the virus, thereby suggesting that rates of viral infection could be reduced by the coccosphere during the critical phase in the cell cycle. However, the benefit of this potential protection is arguably of limited ecological significance since the concentrations of both, calcified and decalcified E. huxleyi approached similar values until the end of the bloom. We conclude that the coccosphere provides at best a limited protection against infection with the EhV86.

Description: Eastern boundary upwelling systems (EBUS) are among the most productive marine ecosystems on Earth. The production of organic material is fueled by upwelling of nutrient-rich deep waters and high incident light at the sea surface. However, biotic and abiotic factors can modify surface production and related biogeochemical processes. Determining these factors is important because EBUS are considered hotspots of climate change, and reliable predictions of their future functioning requires understanding of the mechanisms driving the biogeochemical cycles therein. In this field experiment, we used in situ mesocosms as tools to improve our mechanistic understanding of processes controlling organic matter cycling in the coastal Peruvian upwelling system. Eight mesocosms, each with a volume of ∼55 m3, were deployed for 50 d ∼6 km off Callao (12∘ S) during austral summer 2017, coinciding with a coastal El Niño phase. After mesocosm deployment, we collected subsurface waters at two different locations in the regional oxygen minimum zone (OMZ) and injected these into four mesocosms (mixing ratio ≈1.5 : 1 mesocosm: OMZ water). The focus of this paper is on temporal developments of organic matter production, export, and stoichiometry in the individual mesocosms. The mesocosm phytoplankton communities were initially dominated by diatoms but shifted towards a pronounced dominance of the mixotrophic dinoflagellate (Akashiwo sanguinea) when inorganic nitrogen was exhausted in surface layers. The community shift coincided with a short-term increase in production during the A. sanguinea bloom, which left a pronounced imprint on organic matter C : N : P stoichiometry. However, C, N, and P export fluxes did not increase because A. sanguinea persisted in the water column and did not sink out during the experiment. Accordingly, export fluxes during the study were decoupled from surface production and sustained by the remaining plankton community. Overall, biogeochemical pools and fluxes were surprisingly constant for most of the experiment. We explain this constancy by light limitation through self-shading by phytoplankton and by inorganic nitrogen limitation which constrained phytoplankton growth. Thus, gain and loss processes remained balanced and there were few opportunities for blooms, which represents an event where the system becomes unbalanced. Overall, our mesocosm study revealed some key links between ecological and biogeochemical processes for one of the most economically important regions in the oceans.

Description: Phytoplankton growth rate is a key variable controlling species succession and ecosystem structure throughout the surface ocean. Carbonate chemistry conditions are known to influence phytoplankton growth rates but there is no conceptual framework allowing us to compare growth rate responses across taxa. Here we analyse the literature to show that phytoplankton growth rates follow an optimum curve response pattern whenever the tested species is exposed to a sufficiently large gradient in proton (H+) concentrations. Based on previous findings with coccolithophores and diatoms, we argue that this ‘universal reaction norm’ is shaped by the stimulating influence of increasing inorganic carbon substrate (left side of the optimum) and the inhibiting influence of increase H+ (right side of the optimum). We envisage that exploration of carbonate chemistry-dependent optimum curves as a default experimental approach will boost our mechanistic understanding of phytoplankton responses to ocean acidification, like temperature curves have already boosted our mechanistic understanding to global warming.

Description: The Paris Agreement to limit global warming to well below 2 °C requires ambitious emission reduction and the balancing of remaining emissions through carbon sinks, i.e. the deployment of carbon dioxide removal (CDR). While ambitious climate protection scenarios until now consider primarily land-based CDR methods, there is growing concern about their potential to deliver sufficient CDR, and marine CDR options receive more and more interest. Based on idealized theoretical studies, Ocean Alkalinity Enhancement (OAE) appears as a promising marine CDR method. However, the knowledge base is insufficient for a robust assessment of its practical feasibility, of its side effects, social and governance aspects as well as monitoring, reporting and verification issues. A number of research efforts aim to improve this in a timely manner. We provide an overview on the current situation of developing OAE as marine CDR method, and describe the history that has led to the creation of the OAE research Best Practices Guide.