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  • 1
    Publication Date: 2024-03-25
    Description: Enhancing ocean productivity by artificial upwelling is evaluated as a nature-based solution for food security and climate change mitigation. Fish production is intended through diatom-based plankton food webs as these are assumed to be short and efficient. However, our findings from mesocosm experiments on artificial upwelling in the oligotrophic ocean disagree with this classical food web model. Here, diatoms did not reduce trophic length and instead impaired the transfer of primary production to crustacean grazers and small pelagic fish. The diatom-driven decrease in trophic efficiency was likely mediated by changes in nutritional value for the copepod grazers. Whilst diatoms benefitted the availability of essential fatty acids, they also caused unfavorable elemental compositions via high carbon-to-nitrogen ratios (i.e. low protein content) to which the grazers were unable to adapt. This nutritional imbalance for grazers was most pronounced in systems optimized for CO2 uptake through carbon-to-nitrogen ratios well beyond Redfield. A simultaneous enhancement of fisheries production and carbon sequestration via artificial upwelling may thus be difficult to achieve given their opposing stoichiometric constraints. Our study suggest that food quality can be more critical than quantity to maximize food web productivity during shorter-term fertilization of the oligotrophic ocean.
    Repository Name: EPIC Alfred Wegener Institut
    Type: Article , isiRev
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  • 2
    Publication Date: 2021-04-23
    Description: The oceans absorb about a quarter of the annually produced anthropogenic atmospheric carbon dioxide (CO2), resulting in a decrease in surface water pH, a process termed ocean acidification (OA). Surprisingly little is known about how OA affects the physiology of heterotrophic bacteria or the coupling of heterotrophic bacteria to phytoplankton when nutrients are limited. Previous experiments were, for the most part, undertaken during productive phases or following nutrient additions designed to stimulate algal blooms. Therefore, we performed an in situ large-volume mesocosm ( ∼  55 m3) experiment in the Baltic Sea by simulating different fugacities of CO2 (fCO2) extending from present to future conditions. The study was conducted in July–August after the nominal spring bloom, in order to maintain low-nutrient conditions throughout the experiment. This resulted in phytoplankton communities dominated by small-sized functional groups (picophytoplankton). There was no consistent fCO2-induced effect on bacterial protein production (BPP), cell-specific BPP (csBPP) or biovolumes (BVs) of either free-living (FL) or particle-associated (PA) heterotrophic bacteria, when considered as individual components (univariate analyses). Permutational Multivariate Analysis of Variance (PERMANOVA) revealed a significant effect of the fCO2 treatment on entire assemblages of dissolved and particulate nutrients, metabolic parameters and the bacteria–phytoplankton community. However, distance-based linear modelling only identified fCO2 as a factor explaining the variability observed amongst the microbial community composition, but not for explaining variability within the metabolic parameters. This suggests that fCO2 impacts on microbial metabolic parameters occurred indirectly through varying physicochemical parameters and microbial species composition. Cluster analyses examining the co-occurrence of different functional groups of bacteria and phytoplankton further revealed a separation of the four fCO2-treated mesocosms from both control mesocosms, indicating that complex trophic interactions might be altered in a future acidified ocean. Possible consequences for nutrient cycling and carbon export are still largely unknown, in particular in a nutrient-limited ocean.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 3
    Publication Date: 2021-04-23
    Description: 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.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 4
    Publication Date: 2023-02-08
    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.
    Type: Article , PeerReviewed
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  • 5
  • 6
    Publication Date: 2023-12-21
    Description: Ocean alkalinity enhancement (OAE) has been proposed as a carbon dioxide removal technology (CDR) allowing for long term storage of carbon dioxide in the ocean. By changing the carbonate speciation in seawater, OAE may potentially alter marine ecosystems with implications for the biological carbon pump. Using mesocosmsthe subtropical North Atlantic, we provide first empirical insights into impacts of carbonate-based OAE on the vertical flux and attenuation of sinking particles in an oligotrophic plankton community. We enhanced total alkalinity (TA) in increments of 300 μmol kg-1, reaching up to ΔTA = 2400 µmol kg-1 compared to ambient TA. We applied a pCO2-equilibrated OAE approach, i.e. dissolved inorganic carbon (DIC) was raised simultaneously with TA to maintain seawater pCO2 in equilibrium with the atmosphere, thereby keeping perturbations of seawater carbonate chemistry moderate. The vertical flux of major elements including carbon, nitrogen, phosphorus and silicon, as well as their stoichiometric ratios (e.g. carbon-to-nitrogen) remained unaffected over 29 days of OAE. The particle properties controlling the flux attenuationinking velocities and remineralization rates also remained unaffected by OAE. However, we observed abiotic mineral precipitation at high OAE levels (ΔTA = 1800 μmol kg-1 and higher) that resulted in a substantial increase in PIC formation. The associated consumption of alkalinity reduces the efficiency of CO2 removal and emphasizes the importance of maintaining OAE within a carefully defined operating range. Our findings suggest that carbon export by oligotrophic plankton communities is insensitive to OAE perturbations using a CO2 pre-equilibrated approach. The integrity of ecosystem services is a prerequisite for large-scale application and should be further tested across a variety of nutrient-regimes and for less idealized OAE approaches.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 7
    Publication Date: 2024-02-07
    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.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 8
    Publication Date: 2024-02-07
    Description: Ocean artificial upwelling has been suggested to boost primary production and increase harvestable resources such as fish. Yet, for this ecosystem-based approach to work, an effective energy transfer up the food web is required. Here, we studied the trophic role of microzooplankton under artificial upwelling via biomass and community composition as well as grazing rates on phytoplankton. Using mesocosms in the oligotrophic ocean, we supplied nutrient-rich deep water at varying intensities (low to high) and addition modes (a Singular large pulse or smaller Recurring pulses). Deep-water fertilization created a diatom-dominated bloom that scaled with the amount of inorganic nutrients added, but also Synechococcus -like cells, picoeukaryotes and nanophytoplankton increased in abundance with added nutrients. After 30 days, towards the end of the experiment, coccolithophores bloomed under recurring upwelling of high intensity. Across all upwelling scenarios, the microzooplankton community was dominated by ciliates, dinoflagellates (mixo- and heterotrophic) and radiolarians. Under the highest upwelling intensity, the average grazing rates of Synechococcus -like cells, picoeukaryotes and nanophytoplankton by microzooplankton were 0.35 d -1 ± 0.18 (SD), 0.09 d -1 ± 0.12 (SD), and 0.11 d -1 ± 0.13 (SD), respectively. There was little temporal variation in grazing of nanophytoplankton but grazing of Synechococcus -like cells and picoeukaryotes were more variable. There were positive correlations between abundance of these groups and grazing rates, suggesting a response in the microzooplankton community to prey availability. The average phytoplankton to microzooplankton ratio (biovolume) increased with added deep water, and this increase was highest in the Singular treatment, reaching ~30 (m 3 m -3 ), whereas the phytoplankton to total zooplankton biomass ratio (weight) increased from ~1 under low upwelling to ~6 (g g -1 ) in the highest upwelling but without a difference between the Singular and the Recurring mode. Several smaller, recurring upwelling events increased the importance of microzooplankton compared with one large pulse of deep water. Our results demonstrate that microzooplankton would be an important component for trophic transfer if artificial upwelling would be carried out at scale in the oligotrophic ocean.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 9
    Publication Date: 2024-02-07
    Description: Reduction of anthropogenic CO2 emissions alone will not sufficiently restrict global warming and enable the 1.5°C goal of the Paris agreement to be met. To effectively counteract climate change, measures to actively remove carbon dioxide from the atmosphere are required. Artificial upwelling has been proposed as one such carbon dioxide removal technique. By fueling primary productivity in the surface ocean with nutrient-rich deep water, it could potentially enhance downward fluxes of particulate organic carbon (POC) and carbon sequestration. In this study we investigated the effect of different intensities of artificial upwelling combined with two upwelling modes (recurring additions vs. one singular addition) on POC export, sinking matter stoichiometry and remineralization depth. We carried out a 39 day-long mesocosm experiment in the subtropical North Atlantic, where we fertilized oligotrophic surface waters with different amounts of deep water. The total nutrient inputs ranged from 1.6 to 11.0 μmol NO3– L–1. We found that on the one hand POC export under artificial upwelling more than doubled, and the molar C:N ratios of sinking organic matter increased from values around Redfield (6.6) to ∼8–13, which is beneficial for potential carbon dioxide removal. On the other hand, sinking matter was remineralized at faster rates and showed lower sinking velocities, which led to shallower remineralization depths. Particle properties were more favorable for deep carbon export in the recurring upwelling mode, while in the singular mode the C:N increase of sinking matter was more pronounced. In both upwelling modes roughly half of the produced organic carbon was retained in the water column until the end of the experiment. This suggests that the plankton communities were still in the process of adjustment, possibly due to the different response times of producers and consumers. There is thus a need for studies with longer experimental durations to quantify the responses of fully adjusted communities. Finally, our results revealed that artificial upwelling affects a variety of sinking particle properties, and that the intensity and mode with which it is applied control the strength of the effects
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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  • 10
    Publication Date: 2024-02-07
    Description: The Peruvian upwelling system is a highly productive ecosystem with a large oxygen minimum zone (OMZ) close to the surface. In this work, we carried out a mesocosm experiment off Callao, Peru, with the addition of water masses from the regional OMZ collected at two different sites simulating two different upwelling scenarios. Here, we focus on the pelagic remineralization of organic matter by the extracellular enzyme activity of leucine aminopeptidase (LAP) and alkaline phosphatase activity (APA). After the addition of the OMZ water, dissolved inorganic nitrogen (N) was depleted, but the standing stock of phytoplankton was relatively high, even after N depletion (mostly 〉 4 µg chlorophyll a L−1). During the initial phase of the experiment, APA was 0.6 nmol L−1 h−1 even though the PO concentration was 〉 0.5 µmol L−1. Initially, the dissolved organic phosphorus (DOP) decreased, coinciding with an increase in the PO concentration that was probably linked to the APA. The LAP activity was very high, with most of the measurements in the range of 200–800 nmol L−1 h−1. This enzyme hydrolyzes terminal amino acids from larger molecules (e.g., peptides or proteins), and these high values are probably linked to the highly productive but N-limited coastal ecosystem. Moreover, the experiment took place during a rare coastal El Niño event with higher than normal surface temperatures, which could have affected enzyme activity. Using a nonparametric multidimensional scaling analysis (NMDS) with a generalized additive model (GAM), we found that biogeochemical variables (e.g., nutrient and chlorophyll-a concentrations) and phytoplankton and bacterial communities explained up to 64 % of the variability in APA. The bacterial community best explained the variability (34 %) in LAP. The high hydrolysis rates for this enzyme suggest that pelagic N remineralization, likely driven by the bacterial community, supported the high standing stock of primary producers in the mesocosms after N depletion.
    Type: Article , PeerReviewed , info:eu-repo/semantics/article
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