Beschreibung: Diazotrophic dinitrogen (N2) fixation contributes ~76% to "new" nitrogen inputs to the sunlit open ocean, but environmental factors determining N2 fixation rates are not well constrained. Excess phosphate (phosphate-nitrate/16 〉 0) and iron availability control N2 fixation rates in the eastern tropical North Atlantic (ETNA), but it remains an open question how excess phosphate is generated within or supplied to the phosphate-depleted sunlit layer. Our observations in the ETNA region (8°N-15°N, 19°W-23°W) suggest that Prochlorococcus and Synechococcus, the two ubiquitous non-diazotrophic cyanobacteria with cellular N:P ratios higher than the Redfield ratio, create an environment of excess phosphate, which cannot be explained by diapycnal mixing, atmospheric, and riverine inputs. Thus, our results unveil a new biogeochemical niche construction mechanism by non-diazotrophic cyanobacteria for their diazotrophic phylum group members (N2 fixers). Our observations may help to understand the prevalence of diazotrophy in low-phosphate, oligotrophic regions.

Beschreibung: Ocean acidification is expected to influence plankton community structure and biogeochemical element cycles. To date, the response of plankton communities to elevated CO2 has been studied primarily during nutrient-stimulated blooms. In this CO2 manipulation study, we used large-volume (~ 55 m3) pelagic in situ mesocosms to enclose a natural summer, post-spring-bloom plankton assemblage in the Baltic Sea to investigate the response of organic matter pools to ocean acidification. The carbonate system in the six mesocosms was manipulated to yield average fCO2 ranging between 365 and ~ 1230 μatm with no adjustment of naturally available nutrient concentrations. Plankton community development and key biogeochemical element pools were subsequently followed in this nitrogen-limited ecosystem over a period of 7 weeks. We observed higher sustained chlorophyll a and particulate matter concentrations (~ 25 % higher) and lower inorganic phosphate concentrations in the water column in the highest fCO2 treatment (1231 μatm) during the final 2 weeks of the study period (Phase III), when there was low net change in particulate and dissolved matter pools. Size-fractionated phytoplankton pigment analyses indicated that these differences were driven by picophytoplankton (〈 2 μm) and were already established early in the experiment during an initial warm and more productive period with overall elevated chlorophyll a and particulate matter concentrations. However, the influence of picophytoplankton on bulk organic matter pools was masked by high biomass of larger plankton until Phase III, when the contribution of the small size fraction (〈 2 μm) increased to up to 90 % of chlorophyll a. In this phase, a CO2-driven increase in water column particulate carbon did not lead to enhanced sinking material flux but was instead reflected in increased dissolved organic carbon concentrations. Hence ocean acidification may induce changes in organic matter partitioning in the upper water column during the low-nitrogen summer period in the Baltic Sea.

Beschreibung: Ocean acidification is caused by increasing amounts of carbon dioxide dissolving in the oceans leading to lower seawater pH. We studied the effects of lowered pH on the calanoid copepod Eurytemora affinis during a mesocosm experiment conducted in a coastal area of the Baltic Sea. We measured copepod reproductive success as a function of pH, chlorophyll a concentration, diatom and dinoflagellate biomass, carbon to nitrogen (C : N) ratio of suspended particulate organic matter, as well as copepod fatty acid composition. The laboratory-based experiment was repeated four times during four consecutive weeks, with water and copepods sampled from pelagic mesocosms enriched with different CO2 concentrations. In addition, oxygen radical absorbance capacity (ORAC) of animals from the mesocosms was measured weekly to test whether the copepod's defence against oxidative stress was affected by pH. We found no effect of pH on offspring production. Phytoplankton biomass, as indicated by chlorophyll a concentration, had a strong positive effect. The concentration of polyunsaturated fatty acids in the females were reflected in the eggs and had a positive effect on offspring production, whereas monounsaturated fatty acids of the females were reflected in their eggs but had no significant effect. ORAC was not affected by pH. From these experiments we conclude that E. affinis seems robust against direct exposure to ocean acidification on a physiological level, for the variables covered in the study. E. affinis may not have faced acute pH stress in the treatments as the species naturally face large pH fluctuations.

Beschreibung: Anthropogenic carbon dioxide (CO2) emissions are reducing the pH in the world's oceans. The plankton community is a key component driving biogeochemical fluxes, and the effect of increased CO2 on plankton is critical for understanding the ramifications of ocean acidification on global carbon fluxes. We determined the plankton community composition and measured primary production, respiration rates and carbon export (defined here as carbon sinking out of a shallow, coastal area) during an ocean acidification experiment. Mesocosms ( ∼  55 m3) were set up in the Baltic Sea with a gradient of CO2 levels initially ranging from ambient ( ∼  240 µatm), used as control, to high CO2 (up to  ∼  1330 µatm). The phytoplankton community was dominated by dinoflagellates, diatoms, cyanobacteria and chlorophytes, and the zooplankton community by protozoans, heterotrophic dinoflagellates and cladocerans. The plankton community composition was relatively homogenous between treatments. Community respiration rates were lower at high CO2 levels. The carbon-normalized respiration was approximately 40 % lower in the high-CO2 environment compared with the controls during the latter phase of the experiment. We did not, however, detect any effect of increased CO2 on primary production. This could be due to measurement uncertainty, as the measured total particular carbon (TPC) and combined results presented in this special issue suggest that the reduced respiration rate translated into higher net carbon fixation. The percent carbon derived from microscopy counts (both phyto- and zooplankton), of the measured total particular carbon (TPC), decreased from  ∼  26 % at t0 to  ∼  8 % at t31, probably driven by a shift towards smaller plankton (〈 4 µm) not enumerated by microscopy. Our results suggest that reduced respiration leads to increased net carbon fixation at high CO2. However, the increased primary production did not translate into increased carbon export, and consequently did not work as a negative feedback mechanism for increasing atmospheric CO2 concentration.

Beschreibung: Ocean acidification is challenging phenotypic plasticity of individuals and populations. Calanoid copepods (zooplankton) are shown to be fairly plastic against altered pH conditions, and laboratory studies indicate that transgenerational effects are one mechanism behind this plasticity. We studied phenotypic plasticity of the copepod Acartia sp. in the course of a pelagic, large-volume mesocosm study that was conducted to investigate ecosystem and biogeochemical responses to ocean acidification. We measured copepod egg production rate, egg-hatching success, adult female size and adult female antioxidant capacity (ORAC) as a function of acidification (fCO2  ∼  365–1231 µatm) and as a function of quantity and quality of their diet. We used an egg transplant experiment to reveal whether transgenerational effects can alleviate the possible negative effects of ocean acidification on offspring development. We found significant negative effects of ocean acidification on adult female size. In addition, we found signs of a possible threshold at high fCO2, above which adaptive maternal effects cannot alleviate the negative effects of acidification on egg-hatching and nauplii development. We did not find support for the hypothesis that insufficient food quantity (total particulate carbon 〈 55 µm) or quality (C : N) weakens the transgenerational effects. However, females with high-ORAC-produced eggs with high hatching success. Overall, these results indicate that Acartia sp. could be affected by projected near-future CO2 levels

Beschreibung: About a quarter of anthropogenic CO2 emissions are currently taken up by the oceans, decreasing seawater pH. We performed a mesocosm experiment in the Baltic Sea in order to investigate the consequences of increasing CO2 levels on pelagic carbon fluxes. A gradient of different CO2 scenarios, ranging from ambient ( ∼  370 µatm) to high ( ∼  1200 µatm), were set up in mesocosm bags ( ∼  55 m3). We determined standing stocks and temporal changes of total particulate carbon (TPC), dissolved organic carbon (DOC), dissolved inorganic carbon (DIC), and particulate organic carbon (POC) of specific plankton groups. We also measured carbon flux via CO2 exchange with the atmosphere and sedimentation (export), and biological rate measurements of primary production, bacterial production, and total respiration. The experiment lasted for 44 days and was divided into three different phases (I: t0–t16; II: t17–t30; III: t31–t43). Pools of TPC, DOC, and DIC were approximately 420, 7200, and 25 200 mmol C m−2 at the start of the experiment, and the initial CO2 additions increased the DIC pool by  ∼  7 % in the highest CO2 treatment. Overall, there was a decrease in TPC and increase of DOC over the course of the experiment. The decrease in TPC was lower, and increase in DOC higher, in treatments with added CO2. During phase I the estimated gross primary production (GPP) was  ∼  100 mmol C m−2 day−1, from which 75–95 % was respired,  ∼  1 % ended up in the TPC (including export), and 5–25 % was added to the DOC pool. During phase II, the respiration loss increased to  ∼  100 % of GPP at the ambient CO2 concentration, whereas respiration was lower (85–95 % of GPP) in the highest CO2 treatment. Bacterial production was  ∼  30 % lower, on average, at the highest CO2 concentration than in the controls during phases II and III. This resulted in a higher accumulation of DOC and lower reduction in the TPC pool in the elevated CO2 treatments at the end of phase II extending throughout phase III. The “extra” organic carbon at high CO2 remained fixed in an increasing biomass of small-sized plankton and in the DOC pool, and did not transfer into large, sinking aggregates. Our results revealed a clear effect of increasing CO2 on the carbon budget and mineralization, in particular under nutrient limited conditions. Lower carbon loss processes (respiration and bacterial remineralization) at elevated CO2 levels resulted in higher TPC and DOC pools than ambient CO2 concentration. These results highlight the importance of addressing not only net changes in carbon standing stocks but also carbon fluxes and budgets to better disentangle the effects of ocean acidification.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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.

Beschreibung: 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