ALBERT

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 deep waters make eastern boundary upwelling systems (EBUSs), such as the Humboldt Current system, hot spots of marine productivity. Associated settling of organic matter to depth and consecutive aerobic decomposition results in large subsurface water volumes being oxygen depleted. Under these circumstances, organic matter remineralisation can continue via denitrification, which represents a major loss pathway for bioavailable nitrogen. Additionally, anaerobic ammonium oxidation can remove significant amounts of nitrogen in these areas. Here we assess the interplay of suboxic water upwelling and nitrogen cycling in a manipulative offshore mesocosm experiment. Measured denitrification rates in incubations with water from the oxygen-depleted bottom layer of the mesocosms (via 15N label incubations) mostly ranged between 5.5 and 20 nmol N2 L−1 h−1 (interquartile range), reaching up to 80 nmol N2 L−1 h−1. However, actual in situ rates in the mesocosms, estimated via Michaelis–Menten kinetic scaling, did most likely not exceed 0.2–4.2 nmol N2 L−1 h−1 (interquartile range) due to substrate limitation. In the surrounding Pacific, measured denitrification rates were similar, although indications of substrate limitation were detected only once. In contrast, anammox (anaerobic ammonium oxidation) made only a minor contribution to the overall nitrogen loss when encountered in both the mesocosms and the Pacific Ocean. This was potentially related to organic matter C / N stoichiometry and/or process-specific oxygen and hydrogen sulfide sensitivities. Over the first 38 d of the experiment, total nitrogen loss calculated from in situ rates of denitrification and anammox was comparable to estimates from a full nitrogen budget in the mesocosms and ranged between ∼ 1 and 5.5 µmol N L−1. This represents up to ∼  20 % of the initially bioavailable inorganic and organic nitrogen standing stocks. Interestingly, this loss is comparable to the total amount of particulate organic nitrogen that was exported into the sediment traps at the bottom of the mesocosms at about 20 m depth. Altogether, this suggests that a significant portion, if not the majority of nitrogen that could be exported to depth, is already lost, i.e. converted to N2 in a relatively shallow layer of the surface ocean, provided that there are oxygen-deficient conditions like those during coastal upwelling in our study. Published data for primary productivity and nitrogen loss in all EBUSs reinforce such conclusion.

Description: Ocean deoxygenation due to climate change may alter redox-sensitive nutrient cycles in the marine environment. The productive eastern tropical North Atlantic (ETNA) upwelling region may be particularly affected when the relatively moderate oxygen minimum zone (OMZ) deoxygenates further and microbially driven nitrogen (N) loss processes are promoted. Consequently, water masses with a low nitrogen to phosphorus (N : P) ratio could reach the euphotic layer, possibly influencing primary production in those waters. Previous mesocosm studies in the oligotrophic Atlantic Ocean identified nitrate availability as a control of primary production, while a possible co-limitation of nitrate and phosphate could not be ruled out. To better understand the impact of changing N : P ratios on primary production and N2 fixation in the ETNA surface ocean, we conducted land-based mesocosm experiments with natural plankton communities and applied a broad range of N : P ratios (2.67–48). Silicic acid was supplied at 15 µmol L−1 in all mesocosms. We monitored nutrient drawdown, biomass accumulation and nitrogen fixation in response to variable nutrient stoichiometry. Our results confirmed nitrate to be the key factor determining primary production. We found that excess phosphate was channeled through particulate organic matter (POP) into the dissolved organic matter (DOP) pool. In mesocosms with low inorganic phosphate availability, DOP was utilized while N2 fixation increased, suggesting a link between those two processes. Interestingly this observation was most pronounced in mesocosms where nitrate was still available, indicating that bioavailable N does not necessarily suppress N2 fixation. We observed a shift from a mixed cyanobacteria–proteobacteria dominated active diazotrophic community towards a diatom-diazotrophic association of the Richelia-Rhizosolenia symbiosis. We hypothesize that a potential change in nutrient stoichiometry in the ETNA might lead to a general shift within the diazotrophic community, potentially influencing primary productivity and carbon export.

Description: In open-ocean regions, as is the Eastern Tropical North Atlantic (ETNA), pelagic production is the main source of dissolved organic matter (DOM) and is affected by dissolved inorganic nitrogen (DIN) and phosphorus (DIP) concentrations. Changes in pelagic production under nutrient amendments were shown to also modify DOM quantity and quality. However, little information is available about the effects of nutrient variability on chromophoric (CDOM) and fluorescent (FDOM) DOM dynamics. Here we present results from two mesocosm experiments ("Varied P" and "Varied N") conducted with a natural plankton community from the ETNA, where the effects of DIP and DIN supply on DOM optical properties were studied. CDOM accumulated proportionally to phytoplankton biomass during the experiments. Spectral slope (S) decreased over time indicating accumulation of high molecular weight DOM. In Varied N, an additional CDOM portion, as a result of bacterial DOM reworking, was determined. It increased the CDOM fraction in DOC proportionally to the supplied DIN. The humic-like FDOM component (Comp.1) was produced by bacteria proportionally to DIN supply. The protein-like FDOM component (Comp.2) was released irrespectively to phytoplankton or bacterial biomass, but depended on DIP and DIN concentrations. Under high DIN supply, Comp.2 was removed by bacterial reworking, leading to an accumulation of humic-like Comp.1. No influence of nutrient availability on amino acid-like FDOM component in peptide form (Comp.3) was observed. Comp.3 potentially acted as an intermediate product during formation or degradation of Comp.2. Our findings suggest that changes in nutrient concentrations may lead to substantial responses in the quantity and quality of optically active DOM and, therefore, might bias results of the applied in situ optical techniques for an estimation of DOC concentrations in open-ocean regions.

Description: Recent modeling results suggest that oceanic oxygen levels will decrease significantly over the next decades to centuries in response to climate change and altered ocean circulation. Hence the future ocean may experience major shifts in nutrient cycling triggered by the expansion and intensification of tropical oxygen minimum zones (OMZs). There are numerous feedbacks between oxygen concentrations, nutrient cycling and biological productivity; however, existing knowledge is insufficient to understand physical, chemical and biological interactions in order to adequately assess past and potential future changes. We investigated the pelagic biogeochemistry of OMZs in the eastern tropical North Atlantic and eastern tropical South Pacific during a series of cruise expeditions and mesocosm studies. The following summarizes the current state of research on the influence of low environmental oxygen conditions on marine biota, viruses, organic matter formation and remineralization with a particular focus on the nitrogen cycle in OMZ regions. The impact of sulfidic events on water column biogeochemistry, originating from a specific microbial community capable of highly efficient carbon fixation, nitrogen turnover and N2O production is further discussed. Based on our findings, an important role of sinking particulate organic matter in controlling the nutrient stochiometry of the water column is suggested. These particles can enhance degradation processes in OMZ waters by acting as microniches, with sharp gradients enabling different processes to happen in close vicinity, thus altering the interpretation of oxic and anoxic environments.

Description: Gel particles such as the polysaccharidic transparent exopolymer particles (TEP) and the proteinaceous Coomassie stainable particles (CSP) play an important role in marine biogeochemical and ecological processes like particle aggregation and export, or microbial nutrition and growth. So far, effects of nutrient availability or of changes in nutrient ratios on gel particle production and fate are not well understood. The tropical ocean includes large oxygen minimum zones, where nitrogen losses due to anaerobic microbial activity result in a lower supply of nitrate relative to phosphate to the euphotic zone. Here, we report of two series of mesocosm experiments that were conducted with natural plankton communities collected from the eastern tropical North Atlantic (ETNA) close to Cape Verde in October 2012. The experiments were performed to investigate how different phosphate (experiment 1, Varied P: 0.15–1.58 μmol L−1) or nitrate (experiment 2, Varied N: 1.9–21.9 μmol L−1) concentrations affect the abundance and size distribution of TEP and CSP. In the days until the bloom peak was reached, a positive correlation between gel particle abundance and Chl a concentration was determined, linking the release of dissolved gel precursors and the subsequent formation of gel particles to autotrophic production. After the bloom peak, gel particle abundance remained stable or even increased, implying a continued partitioning of dissolved into particulate organic matter after biomass production itself ceased. During both experiments, differences between TEP and CSP dynamics were observed; TEP were generally more abundant than CSP. Changes in size distribution indicated aggregation of TEP after the bloom, while newly formed CSP decomposed. Abundance of gel particles clearly increased with nitrate concentration during the second experiment, suggesting that changes in [DIN] : [DIP] ratios can affect gel particle formation with potential consequences for carbon and nitrogen cycling as well as food web dynamics in tropical ecosystems.

Description: Concerning their sensitivity to ocean acidification, coccolithophores, a group of calcifying single-celled phytoplankton, are one of the best-studied groups of marine organisms. However, in spite of the large number of studies investigating coccolithophore physiological responses to ocean acidification, uncertainties still remain due to variable and partly contradictory results. In the present study we have used all existing data in a meta-analysis to estimate the effect size of future pCO2 changes on the rates of calcification and photosynthesis and the ratio of particulate inorganic to organic carbon (PIC / POC) in different coccolithophore species. Our results indicate that ocean acidification has a negative effect on calcification and the cellular PIC / POC ratio in the two most abundant coccolithophore species: Emiliania huxleyi and Gephyrocapsa oceanica. In contrast, the more heavily calcified species Coccolithus braarudii did not show a distinct response when exposed to elevated pCO2/reduced pH. Photosynthesis in Gephyrocapsa oceanica was positively affected by high CO2, while no effect was observed for the other coccolithophore species. There was no indication that the method of carbonate chemistry manipulation was responsible for the inconsistent results regarding observed responses in calcification and the PIC / POC ratio. The perturbation method, however, appears to affect photosynthesis, as responses varied significantly between total alkalinity (TA) and dissolved inorganic carbon (DIC) manipulations. These results emphasize that coccolithophore species respond differently to ocean acidification, both in terms of calcification and photosynthesis. Where negative effects occur, they become evident at CO2 levels in the range projected for this century in the case of unabated CO2 emissions. As the data sets used in this meta-analysis do not account for adaptive responses, ecological fitness and ecosystem interactions, the question remains as to how these physiological responses play out in the natural environment.

Description: The upper 500 m of the water column and the sediment surface along an E–W transect in the Fram Strait were sampled for recent planktic foraminifera. The δ18O and δ13C values of the tests are compared to the stable isotope composition of water samples taken from the same depths, and related to the characteristics of the water column. The polar species Neogloboquadrina pachyderma (sin.) clearly dominates the species assemblage in the Fram Strait in the early summer, while the subpolar Turborotalita quinqueloba accounts only for 5–23%. In this area the average depth of calcification of N. pachyderma (sin.) lies between 70–150m water depth, T. quinqueloba shows a similar range with 50–120 m water depth. The δ18O values of N. pachyderma (sin.) show an average vital effect of about −1.5‰ compared to calculated equilibrium calcite values. Except for the upper ~ 75 m, the vertical profiles of δ13C of the net-sampled shells are nearly parallel to the values measured in the water column with an average offset of −1.6‰ and −3.6‰ for N. pachyderma (sin.) and T. quinqueloba, respectively. The discrepancy found in the upper ~ 75 m might indicate the influence of the "carbonate ion effect" on the carbon isotope incorporation in the tests. Oxygen and carbon isotopes from the sediment surface yield higher values than those from the water column for both species. This may be because specimens from the water column reflect a modern snapshot only, while tests from surface sediments record environmental parameters from the past ~ 1000 years.

Description: Epibenthic megafauna play an important role in the deep-sea environment and contribute significantly to benthic biomass, but their population dynamics are still understudied. We used a towed deep-sea camera system to assess the population densities of epibenthic megafauna in 2002, 2007, and 2012 at the shallowest station (HG I, ∼1300 m) of the deep-sea observatory HAUSGARTEN, in the eastern Fram Strait. Our results indicate that the overall density of megafauna was significantly lower in 2007 than in 2002, but was significantly higher in 2012, resulting in overall greater megafaunal density in 2012. Different species showed different patterns in population density, but the relative proportions of predator/scavengers and suspension-feeding individuals were both higher in 2012. Variations in megafaunal densities and proportions are likely due to variation in food input to the sea floor, which decreased slightly in the years preceding 2007 and was greatly elevated in the years preceding 2012. Both average evenness and diversity increased over the time period studied, which indicates that HG I may be food-limited and subject to bottom-up control. The community of HG I may be unique in its response to elevated food input, which resulted in higher evenness and diversity in 2012.

Description: Little is known about the climate evolution on the Kamchatka Peninsula during the last glacial–interglacial transition as existing climate records do not reach beyond 12 ka BP. In this study, a summer temperature record for the past 20 kyr is presented. Branched glycerol dialkyl glycerol tetraethers, terrigenous biomarkers suitable for continental air temperature reconstructions, were analyzed in a sediment core from the western continental margin off Kamchatka in the marginal northwest Pacific (NW Pacific). The record suggests that summer temperatures on Kamchatka during the Last Glacial Maximum (LGM) equaled modern temperatures. We suggest that strong southerly winds associated with a pronounced North Pacific High pressure system over the subarctic NW Pacific accounted for the warm conditions. A comparison with an Earth system model reveals discrepancies between model and proxy-based reconstructions for the LGM temperature and atmospheric circulation in the NW Pacific realm. The deglacial temperature development is characterized by abrupt millennial-scale temperature oscillations. The Bølling–Allerød warm phase and the Younger Dryas cold spell are pronounced events, suggesting a connection to North Atlantic climate variability.