Description: Nitrogen fixation by filamentous cyanobacteria supplies significant amounts of new nitrogen (N) to the Baltic Sea. This balances N loss processes such as denitrification and anammox and forms an important N source supporting primary and secondary production in N-limited post-spring bloom plankton communities. Laboratory studies suggest that filamentous diazotrophic cyanobacteria growth and N2-fixation rates are sensitive to ocean acidification with potential implications for new N supply to the Baltic Sea. In this study, our aim was to assess the effect of ocean acidification on diazotroph growth and activity as well as the contribution of diazotrophically-fixed N to N supply in a natural plankton assemblage. We enclosed a natural plankton community in a summer season in the Baltic Sea near the entrance to the Gulf of Finland in six large-scale mesocosms (volume ~ 55 m3) and manipulated fCO2 over a range relevant for projected ocean acidification by the end of this century (average treatment fCO2: 365–1231 μatm). The direct response of diazotroph growth and activity was followed in the mesocosms over a 47 day study period during N-limited growth in the summer plankton community. Diazotrophic filamentous cyanobacteria abundance throughout the study period and N2-fixation rates (determined only until day 21 due to subsequent use of contaminated commercial 15N-N2 gas stocks) remained low. Thus estimated new N inputs from diazotrophy were too low to relieve N limitation and stimulate a summer phytoplankton bloom. Instead regeneration of organic N sources likely sustained growth in the plankton community. We could not detect significant CO2-related differences in inorganic or organic N pools sizes, or particulate matter N : P stoichiometry. Additionally, no significant effect of elevated CO2 on diazotroph activity was observed. Therefore, ocean acidification had no observable impact on N cycling or biogeochemistry in this N-limited, post-spring bloom plankton assemblage in the Baltic Sea.

Description: This doctoral dissertation presents the results from two independent mesocosm studies on naturally present summer plankton communities in the Baltic Sea. The aim was to investigate the impact of ocean acidification (increased CO2 concentration and decreased seawater pH) as well as the combination of ocean acidification and ocean warming (increased seawater temperature) on the abundance and activity of diazotrophic organisms and on N-limited plankton communities.

Description: 30.12.2018 8° 30‘S 081° 00‘W ca. 120 Seemeilen westlich der Küste Perus MSM80 CUSCO Zweiter Wochenbericht für die Zeit vom 23.12. bis 30.12.2018

Description: In June 2012 we deployed six 19 m long KOSMOS units for 8 weeks in the Tvärminne Storfjärden (59° 51.5’ N, 23° 15.5’ E), an open archipelago area on the south-west corner of Finland, in order to study the influence of ocean acidification on the succession of a plankton community under in-situ conditions in the Baltic Sea. At the beginning of the study (18th of June), we mounted an underwater camera on a diving torch and slowly lowered this setup to a depth of approximately 18.5 m. The video during camera descent shows the post-spring bloom plankton community that we studied within the KOSMOS system. In contrast to mesocosm studies in other regions, the plankton community was less lively: there was a comparatively high number of organic matter aggregations and low abundance of zooplankton. The conical mesocosm sediment trap can be seen at the end of the video in 19 m water depth. The results of this mesocosm study are presented in Biogeosciences in a Special Issue 'Effects of rising CO2 on a Baltic Sea plankton community: ecological and biogeochemical impacts', currently in Biogeosciences Discussion. The video can be downloaded on OceanRep but is also available on the KOSMOS channel of the streaming platform YouTube.

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: 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: Upwelling of nutrient-rich waters into the sunlit surface layer of the ocean supports high primary productivity in eastern boundary upwelling systems (EBUSs). However, subsurface waters contain not only macronutrients (N, P, Si) but also micronutrients, organic matter and seed microbial communities that may modify the response to macronutrient inputs via upwelling. These additional factors are often neglected when investigating upwelling impacts on surface ocean productivity. Here, we investigated how different components of upwelled water (macronutrients, organic nutrients and seed communities) drive the response of surface plankton communities to upwelling in the Peruvian coastal zone. Results from our short-term (10 d) study show that the most influential drivers in upwelled deep water are (1) the ratio of inorganic nutrients (NOx : PO) and (2) the microbial community present that can seed heterogeneity in phytoplankton succession and modify the stoichiometry of residual inorganic nutrients after phytoplankton blooms. Hence, this study suggests that phytoplankton succession after upwelling is modified by factors other than the physical supply of inorganic nutrients. This would likely affect trophic transfer and overall productivity in these highly fertile marine ecosystems.

Description: Ocean acidification (OA) will affect marine biotas from the organism to the ecosystem level. Yet, the consequences for the biological carbon pump and thereby the oceanic sink for atmospheric CO2 are still unclear. Here we show that OA considerably alters the C/N ratio of organic-matter export (C/Nexport), a key factor determining efficiency of the biological pump. By synthesizing sediment-trap data from in situ mesocosm studies in different marine biomes, we find distinct but highly variable impacts of OA on C/Nexport, reaching up to a 20% increase/decrease under partial pressure of CO2 (pCO2) conditions projected for 2100. These changes are driven by pCO2 effects on a variety of plankton taxa and corresponding shifts in food-web structure. Notably, our findings suggest a pivotal role of heterotrophic processes in controlling the response of C/Nexport to OA, thus contradicting the paradigm of primary producers as the principal driver of biogeochemical responses to ocean change.

Description: Marine nitrogen (N2) fixation supports significant primary productivity in the global ocean. However, in one of the most productive regions of the world ocean, the northern Humboldt Upwelling System (HUS), the magnitude and spatial distribution of this process remains poorly characterized. This study presents a spatially resolved dataset of N2 fixation rates across six coastal transects of the northern HUS off Peru (8°S – 16°S) during austral summer. N2 fixation rates were detected throughout the waters column including within the OMZ between 12°S and 16°S. N2 fixation rates were highest where the subsurface Oxygen Minimum Zone (OMZ, O2 〈20 µmol L-1) was most intense and estimated nitrogen (N) loss was highest. There, rates were measured throughout the water column. Hence the vertical and spatial distribution of rates indicates colocation of N2 fixation with N loss in the coastal productive waters of the northern HUS. Despite high phosphate and total dissolvable iron (TdFe) concentrations throughout the study area, N2 fixation was still generally low (1.19 ± 3.81 nmol L-1 d-1) and its distribution could not be directly explained by these two factors. Our results suggest that the distribution was likely influenced by a complex interplay of environmental factors including phytoplankton biomass and organic matter availability, and potentially iron, or other trace metal (co)-limitation of both N2 fixation and primary production. In general, our results support previous conclusions that N2 fixation in the northern HUS plays a minor role as a source of new N and to replenish the regional N loss. Key Points: A north-to-south pattern in N2 fixation rates was observed implying increased N turnover between 12°S and 16°S where N loss was pronounced Highest N2 fixation rates were measured in coastal productive waters above and within the OMZ, showing no clear relationship with Fe or P The magnitude of N2 fixation was low compared to predictions, estimated to account for ∼0.3% of primary production and 〈2% of local N loss