ALBERT

Description: Diatoms are silicifying phytoplankton contributing about one quarter to primary 79 production on Earth. Ocean acidification (OA) could alter the competitiveness of diatoms 80 relative to other taxa and/or lead to shifts among diatom species. In spring 2016, we set 81 up a plankton community experiment at the coast of Gran Canaria (Canary Islands, 82 Spain) to investigate the response of subtropical diatom assemblages to elevated 83 84 seawater pCO2.

Description: Gravitational sinking of photosynthetically fixed particulate organic carbon (POC) constitutes a key component of the biological carbon pump. The fraction of POC leaving the surface ocean depends on POC sinking velocity (SV) and remineralization rate (Cremin), both of which depend on plankton community structure. However, the key drivers in plankton communities controlling SV and Cremin are poorly constrained. In fall 2014, we conducted a 6 weeks mesocosm experiment in the subtropical NE Atlantic Ocean to study the influence of plankton community structure on SV and Cremin. Oligotrophic conditions prevailed for the first 3 weeks, until nutrient‐rich deep water injected into all mesocosms stimulated diatom blooms. SV declined steadily over the course of the experiment due to decreasing CaCO3 ballast and – according to an optical proxy proposed herein – due to increasing aggregate porosity mostly during an aggregation event after the diatom bloom. Furthermore, SV was positively correlated with the contribution of picophytoplankton to the total phytoplankton biomass. Cremin was highest during a Synechococcus bloom under oligotrophic conditions and in some mesocosms during the diatom bloom after the deep‐water addition while it was particularly low during harmful algal blooms. The temporal changes were considerably larger in Cremin (max. 15‐fold) than in SV (max. 3‐fold). Accordingly, estimated POC transfer efficiency to 1000 m was mainly dependent on how the plankton community structure affected Cremin. Our approach revealed key players and interactions in the plankton food web influencing POC export efficiency thereby improving our mechanistic understanding of the biological carbon pump.

Description: Coccolithophores are a group of unicellular phytoplankton species whose ability to calcify has a profound influence on biogeochemical element cycling. Calcification rates are controlled by a large variety of biotic and abiotic factors. Among these factors, carbonate chemistry has gained considerable attention during the last years as coccolithophores have been identified to be particularly sensitive to ocean acidification. Despite intense research in this area, a general concept harmonizing the numerous and sometimes (seemingly) contradictory responses of coccolithophores to changing carbonate chemistry is still lacking to date. Here, we present the "substrate-inhibitor concept" which describes the dependence of calcification rates on carbonate chemistry speciation. It is based on observations that calcification rate scales positively with bicarbonate (HCO3-), the primary substrate for calcification, and carbon dioxide (CO2), which can limit cell growth, whereas it is inhibited by protons (H+). This concept was implemented in a model equation, tested against experimental data, and then applied to understand and reconcile the diverging responses of coccolithophorid calcification rates to ocean acidification obtained in culture experiments. Furthermore, we (i) discuss how other important calcification-influencing factors (e.g. temperature and light) could be implemented in our concept and (ii) embed it in Hutchinson's niche theory, thereby providing a framework for how carbonate chemistry-induced changes in calcification rates could be linked with changing coccolithophore abundance in the oceans. Our results suggest that the projected increase of H+ in the near future (next couple of thousand years), paralleled by only a minor increase of inorganic carbon substrate, could impede calcification rates if coccolithophores are unable to fully adapt. However, if calcium carbonate (CaCO3) sediment dissolution and terrestrial weathering begin to increase the oceans' HCO3- and decrease its H+ concentrations in the far future (10 -100 kyears), coccolithophores could find themselves in carbonate chemistry conditions which may be more favorable for calcification than they were before the Anthropocene.

Description: Ocean acidification is considered as a crucial stressor for marine communities. In this study, we tested the effects of the IPCC RPC6.0 end-of-century acidification scenario on a natural plankton community in the Gullmar Fjord, Sweden, during a long-term mesocosm experiment from a spring bloom to a mid-summer situation. The focus of this study was on microzooplankton and its interactions with phytoplankton and mesozooplankton. The microzooplankton community was dominated by ciliates, especially small Strombidium sp., with the exception of the last days when heterotrophic dinoflagellates increased in abundance. We did not observe any effects of high CO2 on the community composition and diversity of microzooplankton. While ciliate abundance, biomass and growth rate were not affected by elevated CO2, we observed a positive effect of elevated CO2 on dinoflagellate abundances. Additionally, growth rates of dinoflagellates were significantly higher in the high CO2 treatments. Given the higher Chlorophyll a content measured under high CO2, our results point at mainly indirect effects of CO2 on microzooplankton caused by changes in phytoplankton standing stocks, in this case most likely an increase in small-sized phytoplankton of 〈8 µm. Overall, the results from the present study covering the most important part of the growing season indicate that coastal microzooplankton communities are rather robust towards realistic acidification scenarios.

Description: In this study, we report from a long-term in situ mesocosm experiment, where we investigated the response of natural plankton communities in temperate waters (Gullmarfjord, Sweden) to elevated CO2 concentrations and OA as expected for the end of the century (~760 µatm pCO2). Based on a plankton-imaging approach, we examined size structure, community composition and food web characteristics of the whole plankton assemblage, ranging from picoplankton to mesozooplankton, during an entire winter-to-summer succession. The plankton imaging system revealed pronounced temporal changes in the size structure of the copepod community over the course of the plankton bloom. These observed shift towards smaller individuals resulted in an overall decrease of copepod biomass by 25%, despite increasing numerical abundances. Furthermore, we observed distinct effects of elevated CO2 on biomass and size structure of the entire plankton community. Notably, the biomass of copepods, dominated by Pseudocalanus acuspes, displayed a tendency towards elevated biomass by up to 30-40% under simulated ocean acidification. This effect was significant for certain copepod size classes and was most likely driven by CO2-stimulated responses of primary producers and a complex interplay of trophic interactions that allowed this CO2 effect to propagate up the food web.

Description: Climate change challenges phytoplankton communities but evolutionary adaptation could mitigate potential impacts. Here, we tested whether adaptation to a stressor under laboratory conditions leads to equivalent fitness gains in a more natural environment. We found that fitness advantages that had evolved under laboratory conditions were masked by pleiotropic effects in natural plankton communities. Moreover, new genotypes with highly variable competitive abilities evolved on timescales significantly shorter than climate change.

Description: Thermal reaction norms for growth rates of six Emiliania huxleyi isolates originating from the central Atlantic (Azores, Portugal) and five isolates from the coastal North Atlantic (Bergen, Norway) were assessed. We used the template mode of variation model to decompose variations in growth rates into modes of biological interest: vertical shift, horizontal shift, and generalist-specialist variation. In line with the actual habitat conditions, isolates from Bergen (Bergen population) grew well at lower temperatures, and isolates from the Azores (Azores population) performed better at higher temperatures. The optimum growth temperature of the Azores population was significantly higher than that of the Bergen population. Neutral genetic differentiation was found between populations by microsatellite analysis. These findings indicate that E. huxleyi populations are adapted to local temperature regimes. Next to between-population variation, we also found variation within populations. Genotype-by-environment interactions resulted in the most pronounced phenotypic differences when isolates were exposed to temperatures outside the range they naturally encounter. Variation in thermal reaction norms between and within populations emphasizes the importance of using more than one isolate when studying the consequences of global change on marine phytoplankton. Phenotypic plasticity and standing genetic variation will be important in determining the potential of natural E. huxleyi populations to cope with global climate change.

Description: Although coccolithophore physiological responses to CO2-induced changes in seawater carbonate chemistry have been widely studied in the past, there is limited knowledge on the variability of physiological responses between populations from different areas. In the present study, we investigated the specific responses of growth, particulate organic (POC) and inorganic carbon (PIC) production rates of three populations of the coccolithophore Emiliania huxleyi from three regions in the North Atlantic Ocean (Azores: six strains, Canary Islands: five strains, and Norwegian coast near Bergen: six strains) to a CO2 partial pressure (pCO2) range from 120 to 2630µatm. Physiological rates of each population and individual strain increased with rising pCO2 levels, reached a maximum and declined thereafter. Optimal pCO2 for growth, POC production rates, and tolerance to low pH (i.e., high proton concentration) was significantly higher in an E. huxleyi population isolated from the Norwegian coast than in those isolated near the Azores and Canary Islands. This may be due to the large environmental variability including large pCO2 and pH fluctuations in coastal waters off Bergen compared to the rather stable oceanic conditions at the other two sites. Maximum growth and POC production rates of the Azores and Bergen populations were similar and significantly higher than that of the Canary Islands population. This pattern could be driven by temperature–CO2 interactions where the chosen incubation temperature (16°C) was slightly below what strains isolated near the Canary Islands normally experience. Our results indicate adaptation of E. huxleyi to their local environmental conditions and the existence of distinct E. huxleyi populations. Within each population, different growth, POC, and PIC production rates at different pCO2 levels indicated strain-specific phenotypic plasticity. Accounting for this variability is important to understand how or whether E. huxleyi might adapt to rising CO2 levels.

Description: Coccolithophores -single-celled calcifying phytoplankton- are an important group of marine primary producers and the dominant builders of calcium carbonate globally. Coccolithophores form extensive blooms and increase the density and sinking speed of organic matter via calcium carbonate ballasting. Thereby, they play a key role in the marine carbon cycle. Coccolithophore physiological responses to experimental ocean acidification have ranged from moderate stimulation to substantial decline in growth and calcification rates, combined with enhanced malformation of their calcite platelets. Here we report on a mesocosm experiment conducted in a Norwegian fjord in which we exposed a natural plankton community to a wide range of CO2-induced ocean acidification, to test whether these physiological responses affect the ecological success of coccolithophore populations. Under high-CO2 treatments, Emiliania huxleyi, the most abundant and productive coccolithophore species, declined in population size during the pre-bloom period and lost the ability to form blooms. As a result, particle sinking velocities declined by up to 30% and sedimented organic matter was reduced by up to 25% relative to controls. There were also strong reductions in seawater concentrations of the climate-active compound dimethylsulfide in CO2-enriched mesocosms. We conclude that ocean acidification can lower calcifying phytoplankton productivity, potentially creating a positive feedback to the climate system.