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  • 1
    Publikationsdatum: 2021-03-30
    Beschreibung: Anthropogenic atmospheric loading of CO2 raises concerns about combined effects of increasing ocean temperature and acidification, on biological processes. In particular, the response of appendicularian zooplankton to climate change may have significant ecosystem implications as they can alter biogeochemical cycling compared to classical copepod dominated food webs. However, the response of appendicularians to multiple climate drivers and effect on carbon cycling are still not well understood. Here, we investigated how gelatinous zooplankton (appendicularians) affect carbon cycling of marine food webs under conditions predicted by future climate scenarios. Appendicularians performed well in warmer conditions and benefited from low pH levels, which in turn altered the direction of carbon flow. Increased appendicularians removed particles from the water column that might otherwise nourish copepods by increasing carbon transport to depth from continuous discarding of filtration houses and fecal pellets. This helps to remove CO2 from the atmosphere, and may also have fisheries implications.
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  • 2
    Publikationsdatum: 2021-04-23
    Beschreibung: Nitrogen (N) is the major limiting nutrient for phytoplankton growth and productivity in large parts of the world's oceans. Differential preferences for specific N substrates may be important in controlling phytoplankton community composition. To date, there is limited information on how specific N substrates influence the composition of naturally occurring microbial communities. We investigated the effect of nitrate ( math formula), ammonium ( math formula), and urea on microbial and phytoplankton community composition (cell abundances and 16S rRNA gene profiling) and functioning (photosynthetic activity, carbon fixation rates) in the oligotrophic waters of the North Pacific Ocean. All N substrates tested significantly stimulated phytoplankton growth and productivity. Urea resulted in the greatest (〉300%) increases in chlorophyll a (〈0.06 μg L−1 and ∼0.19 μg L−1 in the control and urea addition, respectively) and productivity (〈0.4 μmol C L−1 d−1 and ∼1.4 μmol C L−1 d−1 in the control and urea addition, respectively) at two experimental stations, largely due to increased abundances of Prochlorococcus (Cyanobacteria). Two abundant clades of Prochlorococcus, High Light I and II, demonstrated similar responses to urea, suggesting this substrate is likely an important N source for natural Prochlorococcus populations. In contrast, the heterotrophic community composition changed most in response to math formula. Finally, the time and magnitude of response to N amendments varied with geographic location, likely due to differences in microbial community composition and their nutrient status. Our results provide support for the hypothesis that changes in N supply would likely favor specific populations of phytoplankton in different oceanic regions and thus, affect both biogeochemical cycles and ecological processes.
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  • 3
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 37 (6). pp. 1146-1163.
    Publikationsdatum: 2018-03-20
    Beschreibung: Denitrification was investigated in the Baltic proper at two stations with different conditions in the deep water. The Gotland Deep was examined as an example of a basin with anoxic, H2S‐containing deep water and station T was taken as an example of low‐oxygen (〈0.2 ml liter−1), sulfide‐free deep water. Denitrification was measured by the acetylene blockage method; in addition, N2O reduction was followed in samples without acetylene. To shed light on the factors limiting denitrification, we compared in situ rates to denitrification after adding nitrate or electron donors. Denitrification was restricted to the layer of the oxic‐anoxic interface in the Gotland Deep and to the water layer near the sediment of station T. For both stations it could be shown that denitrification was not limited by nitrate availability. A lack of available organic C seemed to limit denitrification rates and growth of denitrifiers. As a result of C limitation in the water column, denitrification was restricted to energy‐rich interfaces. In the low‐oxygen water away from energy‐rich interfaces, the less C‐demanding nitrification‐denitrification coupling (NH4+ → N2O → N2) seemed to be favored. Denitrification in the water of the central Baltic seems to be subjected to strong variability due to changing C supply during the course of the year. However, limitation by C availability can be assumed for most of the year and should be taken into account in calculating the N budget of the Baltic.
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  • 4
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 61 (S1). S300-S323.
    Publikationsdatum: 2019-09-23
    Beschreibung: Methane concentration and isotopic composition was measured in ice-covered and ice-free waters of the Arctic Ocean during 11 surveys spanning the years of 1992–1995 and 2009. During ice-free periods, methane flux from the Beaufort shelf varies from 0.14 mg CH4 m−2 d−1 to 0.43 mg CH4 m−2 d−1. Maximum fluxes from localized areas of high methane concentration are up to 1.52 mg CH4 m−2 d−1. Seasonal buildup of methane under ice can produce short-term fluxes of methane from the Beaufort shelf that varies from 0.28 mg CH4 m−2 d−1 to 1.01 mg CH4 m−2 d−1. Scaled-up estimates of minimum methane flux from the Beaufort Sea and pan-Arctic shelf for both ice-free and ice-covered periods range from 0.02 Tg CH4 yr−1 and 0.30 Tg CH4 yr−1, respectively to maximum fluxes of 0.18 Tg CH4 yr−1 and 2.2 Tg CH4 yr−1, respectively. A methane flux of 0.36 Tg CH4 yr−1 from the deep Arctic Ocean was estimated using data from 1993 to 1994. The flux can be as much as 2.35 Tg CH4 yr−1 estimated from maximum methane concentrations and wind speeds of 12 m/s, representing only 0.42% of the annual atmospheric methane budget of ∼ 560 Tg CH4 yr−1. There were no significant changes in methane fluxes during the time period of this study. Microbial methane sources predominate with minor influxes from thermogenic methane offshore Prudhoe Bay and the Mackenzie River delta and may include methane from gas hydrate. Methane oxidation is locally important on the shelf and is a methane sink in the deep Arctic Ocean.
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  • 5
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    ASLO (Association for the Sciences of Limnology and Oceanography) | Wiley
    In:  Limnology and Oceanography, 63 (2). pp. 968-984.
    Publikationsdatum: 2021-06-23
    Beschreibung: Overfishing, excess nutrient load, and invasion of Mnemiopsis leidyi acted on the Black Sea from 1960s to 1990s. Under the effect of these drivers, the ecosystem underwent several transformations that culminated in the shift from a planktonic food chain to a network with most of the energy diverted to jellyfish. The interplay between multiple stressors and the intricate web of trophic interactions make it difficult to understand which causative mechanisms linked the sources of change to the observed variations. To study such interplay, we focused on the structure of the trophic interactions and applied loop analysis to qualitatively predict the response of variables to stressors. Significant variations in biomass trends were identified with statistical analysis and considered as benchmark to validate loop analysis predictions. The results of the comparisons were used to select the most influential trophic interactions that explain the changes observed between 1960 and 1990. The models were applied to test (1) the importance of various environmental drivers and (2) the mechanisms that produced the observed changes. The results suggested that the changes observed before M. leidyi invasion were strongly influenced by the excess nutrient addition, an outcome that challenges the relevance of the trophic cascade as described in literature. The concurrent effect of overfishing, climate, and nutrient enrichment likely triggered the outburst of M. leidyi in the late 1980s. Our work shows the potential of loop analysis to grasp the causal relationships between the drivers, the structure of the interactions, and the responses of the variables.
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  • 6
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    ASLO (Association for the Sciences of Limnology and Oceanography) | Wiley
    In:  Limnology and Oceanography, 63 (1). pp. 3-21.
    Publikationsdatum: 2021-02-08
    Beschreibung: Ocean acidification (OA) is generally assumed to negatively impact calcification rates of marine organisms. At a local scale however, biological activity of macrophytes may generate pH fluctuations with rates of change that are orders of magnitude larger than the long-term trend predicted for the open ocean. These fluctuations may in turn impact benthic calcifiers in the vicinity. Combining laboratory, mesocosm and field studies, such interactions between OA, the brown alga Fucus vesiculosus, the sea grass Zostera marina and the blue mussel Mytilus edulis were investigated at spatial scales from decimetres to 100s of meters in the western Baltic. Macrophytes increased the overall mean pH of the habitat by up to 0.3 units relative to macrophyte-free, but otherwise similar, habitats and imposed diurnal pH fluctuations with amplitudes ranging from 0.3 to more than 1 pH unit. These amplitudes and their impact on mussel calcification tended to increase with increasing macrophyte biomass to bulk water ratio. At the laboratory and mesocosm scales, biogenic pH fluctuations allowed mussels to maintain calcification even under acidified conditions by shifting most of their calcification activity into the daytime when biogenic fluctuations caused by macrophyte activity offered temporal refuge from OA stress. In natural habitats with a low biomass to water body ratio, the impact of biogenic pH fluctuations on mean calcification rates of M. edulis was less pronounced. Thus, in dense algae or seagrass habitats, macrophytes may mitigate OA impact on mussel calcification by raising mean pH and providing temporal refuge from acidification stress.
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  • 7
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    ASLO (Association for the Sciences of Limnology and Oceanography) | Wiley
    In:  Limnology and Oceanography, 63 (1). pp. 445-458.
    Publikationsdatum: 2021-02-08
    Beschreibung: Despite its delicate morphology, the lobate ctenophore Mnemiopsis leidyi thrives in coastal ecosystems as an influential zooplankton predator. Coastal ecosystems are often characterized as energetic systems with high levels of natural turbulence in the water column. To understand how natural wind-driven turbulence affects the feeding ecology of M. leidyi, we used a combination of approaches to quantify how naturally and laboratory generated turbulence affects the behavior, feeding processes and feeding impact of M. leidyi. Experiments using laboratory generated turbulence demonstrated that turbulence can reduce M. leidyi feeding rates on copepods and Artemia nauplii by 〉 50%. However, detailed feeding data from the field, collected during highly variable surface conditions, showed that wind-driven turbulence did not affect the feeding rates or prey selection of M. leidyi. Additional laboratory experiments and field observations suggest that the feeding process of M. leidyi is resilient to wind-driven turbulence because M. leidyi shows a behavioral response to turbulence by moving deeper in the water column. Seeking refuge in deeper waters enables M. leidyi to maintain high feeding rates even under high turbulence conditions generated by wind driven mixing. As a result, M. leidyi exerted a consistently high predatory impact on prey populations during highly variable and often energetic wind-driven mixing conditions. This resilience adds to our understanding of how M. leidyi can thrive in a wide spectrum of environments around the world. The limits to this resilience also set boundaries to its range expansion into novel areas.
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  • 8
    Publikationsdatum: 2020-02-06
    Beschreibung: Thaumarchaea are among the most abundant microbial groups in the ocean, but controls on their abundance and the distribution and metabolic potential of different subpopulations are poorly constrained. Here, two ecotypes of ammonia-oxidizing thaumarchaea were quantified using ammonia monooxygenase (amoA) genes across the equatorial Pacific Ocean. The shallow, or water column “A” (WCA), ecotype was the most abundant ecotype at the depths of maximum nitrification rates, and its abundance correlated with other biogeochemical indicators of remineralization such as NO3 : Si and total Hg. Metagenomes contained thaumarchaeal genes encoding for the catalytic subunit of the urease enzyme (ureC) at all depths, suggesting that members of both WCA and the deep, water column “B” (WCB) ecotypes may contain ureC. Coupled urea hydrolysis-ammonia oxidation rates were similar to ammonia oxidation rates alone, suggesting that urea could be an important source of ammonia for mesopelagic ammonia oxidizers. Potential inducement of metal limitation of both ammonia oxidation and urea hydrolysis was demonstrated via additions of a strong metal chelator. The water column inventory of WCA was correlated with the depth-integrated abundance of WCB, with both likely controlled by the flux of sinking particulate organic matter, providing strong evidence of vertical connectivity between the ecotypes. Further, depth-integrated amoA gene abundance and nitrification rates were correlated with particulate organic nitrogen flux measured by contemporaneously deployed sediment traps. Together, the results refine our understanding of the controls on thaumarchaeal distributions in the ocean, and provide new insights on the relationship between material flux and microbial communities in the mesopelagic.
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  • 9
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 60 (1). pp. 100-109.
    Publikationsdatum: 2018-01-01
    Beschreibung: The influential predatory role of the lobate comb jellyfish Mnemiopsis leidyi has largely been attributed to the generation of a hydrodynamically silent feeding current to entrain and initiate high encounter rates with prey. However, for high encounter rates to translate to high ingestion rates, M. leidyi must effectively capture the entrained prey. To investigate the capture mechanisms, we recorded and quantified, using three-dimensional videography, the outcome of encounter events with slow swimming Artemia prey. The auricles, which produce the feeding current of M. leidyi, were the primary encounter structures, first contacting 59% of the prey in the feeding current. Upon detection, the auricles manipulated the Artemia to initiate captures on the tentillae, which are coated with sticky cells (colloblasts). Using this mechanism of sensory-scanning to capture prey entrained in the feeding current, M. leidyi uses a similar foraging strategy to that of feeding-current foraging copepods. As such, M. leidyi has a higher capture efficiency than do medusae, contributing to the greater predatory effect of M. leidyi in both its endemic and invasive ecosystems.
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  • 10
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 60 (4). pp. 1392-1410.
    Publikationsdatum: 2019-09-23
    Beschreibung: The Arctic Ocean faces multiple environmental changes induced by climate change on both global and regional scale. In addition to global changes in seawater temperature and pH, Arctic waters receive organic matter enrichment due to increasing pelagic primary production, enhanced sea ice melting and increasing terrestrial carbon loads. We experimentally tested individual and combined effects of warming, acidification and organic matter amendment on growth, biomass production and extracellular enzyme activities of bacterioplankton in Fram Strait during early summer. Results reveal pH optima of 6.7–7.6 for extracellular leucine-aminopeptidase and below pH 6.0 for beta-glucosidase in the West Spitsbergen Current. These optima well below the current seawater pH imply increasing hydrolytic activity with ongoing ocean acidification. However, the new synthesis of extracellular enzymes during 4-d incubations obscured the biochemical pH effects. Elevated temperature and carbohydrate supply had strongly interactive effects on bacterial biomass production in both Atlantic Water of the West Spitsbergen Current and Polar Water of the East Greenland Current. Activation energies ranged from 45 kJ mol−1 to 52 kJ mol−1 at in situ substrate concentration, while substantially higher values of 122–174 kJ mol−1 could be estimated from incubations with carbohydrate addition. The net loss of total amino acids in carbohydrate-amended incubations was significantly reduced at elevated temperature in all experiments, suggesting enhanced de novo synthesis. Our findings show that the complexity of combined effects must be considered to better assess the potential of climate change to alter biogenic carbon and energy fluxes in marine systems.
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  • 11
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 60 (3). pp. 901-919.
    Publikationsdatum: 2018-06-01
    Beschreibung: Carbon uptake and partitioning of two globally abundant diatom species, Thalassiosira weissflogii and Dactyliosolen fragilissimus, was investigated in batch culture experiments under four conditions: ambient (15 degrees C, 400 atm), high CO2 (15 degrees C, 1000 atm), high temperature (20 degrees C, 400 atm), and combined (20 degrees C, 1000 atm). The experiments were run from exponential growth into the stationary phase (six days after nitrogen depletion), allowing us to track biogeochemical dynamics analogous to bloom situations in the ocean. Elevated CO2 had a fertilizing effect and enhanced uptake of dissolved inorganic carbon (DIC) by about 8% for T. weissflogii and by up to 39% for D. fragilissimus. This was also reflected in higher cell numbers, build-up of particulate and dissolved organic matter, and transparent exopolymer particles. The CO2 effects were most prominent in the stationary phase when nitrogen was depleted and CO2(aq) concentrations were low. This indicates that diatoms in the high CO2 treatments could take up more DIC until CO2 concentrations in seawater became so low that carbon limitation occurs. These results suggest that, contrary to common assumptions, diatoms could be highly sensitive to ongoing changes in oceanic carbonate chemistry, particularly under nutrient limitation. Warming from 15 to 20 degrees C had a stimulating effect on one species but acted as a stressor on the other species, highlighting the importance of species-specific physiological optima and temperature ranges in the response to ocean warming. Overall, these sensitivities to CO2 and temperature could have profound impacts on diatoms blooms and the biological pump.
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  • 12
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 60 (5). pp. 1619-1633.
    Publikationsdatum: 2018-06-20
    Beschreibung: We present the first set of dissolved silicon isotope data in seawater (delta Si-30(Si(OH)4)) from the East China Sea, a large and productive marginal sea significantly influenced by the Kuroshio Current and freshwater inputs from the Changjiang (Yangtze River). In summer (August 2009), the lowest surface delta Si-30(Si(OH)4) signatures of +2.1 parts per thousand corresponding to the highest Si(OH)(4) concentrations (similar to 30.0 mu mol L-1) were observed nearshore in Changjiang Diluted Water. During advection on the East China Sea inner shelf, surface delta Si-30(Si(OH)4) increased rapidly to +3.2 parts per thousand while Si(OH)(4) became depleted, indicating increasing biological utilization of the Si(OH)(4) originating from the Changjiang Diluted Water. This is also reflected in the water column profiles characterized by a general decrease of delta Si-30(Si(OH)4) and an increase of Si(OH)(4) with depth on the East China Sea mid-shelf and slope. In winter (December 2009-January 2010), however, the delta Si-30(Si(OH)4) was nearly constant at +1.9 parts per thousand throughout the water column on the East China Sea shelf beyond the nearshore, which was a consequence of enhanced vertical mixing of the Kuroshio subsurface water. Horizontal admixture of Kuroshio surface water, which is highly fractionated in Si isotopes, was observed only beyond the shelf break. Significant seasonal differences in delta Si-30(Si(OH)4) were detected in the surface waters beyond the Changjiang Diluted Water-influenced region on the East China Sea shelf, where the winter values were similar to 1.0 parts per thousand lower than those in summer, despite the same primary Si(OH)(4) supply from the Kuroshio subsurface water during both seasons. This demonstrates significantly higher biological consumption and utilization of Si(OH)(4) in summer than in winter.
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  • 13
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 60 (6). pp. 2145-2157.
    Publikationsdatum: 2018-10-01
    Beschreibung: Global change leads to a multitude of simultaneous modifications in the marine realm among which shoaling of the upper mixed layer, leading to enhanced surface layer light intensities, as well as increased carbon dioxide (CO2) concentration are some of the most critical environmental alterations for phytoplankton. In this study, we investigated the responses of growth, photosynthetic carbon fixation and calcification of the coccolithophore Gephyrocapsa oceanica to elevated inline image (51 Pa, 105 Pa, and 152 Pa) (1 Pa ≈ 10 μatm) at a variety of light intensities (50–800 μmol photons m−2 s−1). By fitting the light response curve, our results showed that rising inline image reduced the maximum rates for growth, photosynthetic carbon fixation and calcification. Increasing light intensity enhanced the sensitivity of these rate responses to inline image, and shifted the inline image optima toward lower levels. Combining the results of this and a previous study (Sett et al. 2014) on the same strain indicates that both limiting low inline image and inhibiting high inline image levels (this study) induce similar responses, reducing growth, carbon fixation and calcification rates of G. oceanica. At limiting low light intensities the inline image optima for maximum growth, carbon fixation and calcification are shifted toward higher levels. Interacting effects of simultaneously occurring environmental changes, such as increasing light intensity and ocean acidification, need to be considered when trying to assess metabolic rates of marine phytoplankton under future ocean scenarios.
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  • 14
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 60 (2). pp. 402-410.
    Publikationsdatum: 2015-07-03
    Beschreibung: The effect of submarine carbon dioxide (CO2) vents on seawater carbonate chemistry have been determined using hydrographical and marine carbonate data obtained from two submarine hydrothermal vent fields, as well as a reference station, all near the Jan Mayen Island in the Norwegian-Greenland Sea. We have shown that one can successfully determine the excess carbon that enters the seawater from the vents by applying a modified version of a back-calculation technique, which is traditionally used to study the invasion of excess atmospheric CO2 in the surface ocean. As a result of this excess carbon, total dissolved inorganic carbon (CT) in the seawater surrounding the vents was on average 12 μmol kg−1 (1-30 μmol kg−1) higher compared to samples obtained from a reference station outside the venting areas. The observed excess CT was most significant between 100 m and 200 m but was noticeable in all depths with the exception of the upper 10-20 m. The absence of a venting CO2 signal in the surface water and the realism of the results are discussed. We believe the present method is promising for monitoring (detection and quantification) of CO2 leakage into the water column due to its high sensitivity and readiness for automation.
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  • 15
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography: Methods, 13 (11). pp. 601-608.
    Publikationsdatum: 2021-04-21
    Beschreibung: High quality, reference measurements of chemical and physical properties of seawater are of great importance for a wide research community, including the need to validate models and attempts to quantify spatial and temporal variability. Whereas data precision has been improved by technological advances, the data accuracy has improved mainly by the use of certified reference materials (CRMs). However, since CRMs are not available for all variables, and use of CRMs does not guarantee bias-free data, we here present a recently developed Matlab toolbox for performing so-called secondary quality control on oceanographic data by the use of crossover analysis. This method and how it has been implemented in this toolbox is described in detail. This toolbox is developed mainly for use by sea-going scientists as a tool for quickly assessing possible bias in the measurements that can-hopefully-be remedied during the expedition, but also for possible post-cruise adjustment of data to be consistent with previous measurements in the region. The toolbox, and reference data, can be downloaded from the Carbon Dioxide Information Analysis Center (CDIAC): http://cdiac.ornl.gov/ftp/oceans/2nd_QC_Tool_V2/.
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  • 16
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 61 (3). pp. 853-868.
    Publikationsdatum: 2019-02-01
    Beschreibung: Increasing seawater temperature and CO2 concentrations both are expected to increase coastal phytoplankton biomass and carbon to nutrient ratios in nutrient limited seasonally stratified summer conditions. This is because temperature enhances phytoplankton growth while grazing is suggested to be reduced during such bottom-up controlled situations. In addition, enhanced CO2 concentrations potentially favor phytoplankton species, that otherwise depend on costly carbon concentrating mechanisms (CCM). The trophic consequences for consumers under such conditions, however, remain little understood. We set out to experimentally explore the combined effects of increasing temperature and CO2 concentration for phytoplankton biomass and stoichiometry and the consequences for trophic transfer (here for copepods) on a natural nutrient limited Baltic Sea summer plankton community. The results show, that warming effects were translated to the next trophic level by switching the system from a bottom-up controlled to a mainly top-down controlled one. This was reflected in significantly down-grazed phytoplankton and increased zooplankton abundance in the warm temperature treatment (22.5°C). Additionally, at low temperature (16.5°C) rising CO2 concentrations significantly increased phytoplankton biomass. The latter effect however, was due to direct negative impact of CO2 on copepod nauplii which released phytoplankton from grazing in the cold but not in the warm treatments. Our results suggest that future seawater warming has the potential to switch trophic relations between phytoplankton and their grazers under nutrient limited conditions with the consequence of potentially disguising CO2 effects on coastal phytoplankton biomass.
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  • 17
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 47 . pp. 1324-1335.
    Publikationsdatum: 2014-01-30
    Beschreibung: EisenEx�the second in situ iron enrichment experiment in the Southern Ocean�was performed in the Atlantic sector over 3 weeks in November 2000 with the overarching goal to test the hypothesis that primary productivity in the Southern Ocean is limited by iron availability in the austral spring. Underwater irradiance, chlorophyll a (Chl a), photochemical efficiency, and primary productivity were measured inside and outside of an iron-enriched patch in order to quantify the response of phytoplankton to iron fertilization. Chl a concentration and photosynthetic rate (14C uptake in simulated in situ incubations) were measured in pico-, nano-, and microphytoplankton. Photochemical efficiency was studied with fast repetition rate fluorometry and xenon-pulse amplitude modulated fluorometry. The high-nutrient low-chlorophyll waters outside the Fe-enriched patch were characterized by deep euphotic zones (63-72 m), low Chl a (48-56 mg m-2), low photosynthetic efficiency (Fv/Fm ~ 0.3), and low daily primary productivity (130-220 mg C m-2 d-1). Between 70 and 90% of Chl a was found in pico- and nanophytoplankton. During the induced bloom, Fv/Fm increased up to ;0.55, primary productivity and Chl a reached the maximum values of 790 mg C m-2 d-1 and 231 mg Chl a m-2, respectively. As a consequence, the euphotic depth decreased to ~41 m. Picophytoplankton biomass hardly changed. Nano- and microphytoplankton biomass increased. In the first 2 weeks of the experiment, when the depth of the upper mixed layer was mostly 〈40 m, primary productivity was highly correlated with Chl a. In the third week, productivity was much lower than predicted from Chl a, probably because of a reduction in photosynthetic capacity as a consequence of increased physical variability in the upper water column. These results provide unequivocal evidence that iron supply is the central factor controlling phytoplankton primary productivity in the Southern Ocean, even if the mixing depth is 〉80 m.
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  • 18
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 49 . pp. 1435-1445.
    Publikationsdatum: 2019-09-23
    Beschreibung: Microzooplankton have received increased attention as an important trophic link between the microbial loop and calanoid copepods. On the basis of food size spectra overlap in some microzooplankton groups and calanoid copepods, however, such microzooplankton could function as competitors rather than as food for calanoid copepods (intraguild prey). Mixotrophic flagellates presumably represent a link between the microbial loop and the micro and mesozooplankton. We investigated the effects of microzooplankton and mixotrophy by altering the presence of a heterotrophic dinoflagellate and of a mixotrophic nanoflagellate in artificial food webs with calanoid copepods as terminal consumers. Overall system productivity was manipulated by two levels of nutrient enrichment. The heterotrophic dinoflagellate drastically reduced the nanophytoplankton and enhanced the reproduction of the copepods, suggesting that its role as a competitor is negligible compared to its function as a trophic link. In spite of the presence of heterotrophic nanoflagellates, the mixotroph had a strong negative effect on the picophytoplankton and (presumably) on bacterial biomass. At the same time, the mixotroph enhanced the atomic C:N ratio of the seston biomass, indicating a higher efficiency in overall primary production. Copepod reproduction was enhanced in the presence of the mixotrophic nanoflagellate. Results did not support predictions of the intraguild predation theory: The ratios of the intraguild predators and their preys were not affected by overall system productivity
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  • 19
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 48 (5). pp. 1825-1830.
    Publikationsdatum: 2018-06-25
    Beschreibung: Environmental evidence suggests that Aureococcus anophagefferens (Pelagophyceae), a eukaryotic picoplankton that blooms in coastal seawaters, can outcompete other organisms because of its ability to use abundant dissolved organic nitrogen (DON). To test this hypothesis, we isolated A. anophagefferens in axenic culture and monitored its growth on high-molecular weight (HMW) DON collected from sediment pore waters, a putative source for DON in bays where blooms occur. HMW DON originating from pore water had a substantially higher protein content than surface seawater DON. We found that A. anophagefferens could deplete 25-36% of the available nitrogen in cultures with HMW DON as the sole source of nitrogen and that this corresponded well with the protein fraction in pore-water HMW DON. High rates of cell surface peptide hydrolysis and no detectable N-acetyl polysaccharide hydrolysis, together with the high percentage of hydrolyzable amino acids compared to hydrolyzable aminosugars present in the HMW DON, pointed to the protein fraction as the more likely source of nitrogen used for growth. Whether or not nitrogen scavenging from protein is a common mechanism in phytoplankton is at present unknown but needs to be investigated
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  • 20
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 61 (3). pp. 1120-1133.
    Publikationsdatum: 2019-02-01
    Beschreibung: Marine organisms in the Mediterranean Sea experience the highest temperatures, salinities and oligotrophic conditions in its easternmost part along the eastern shores of the Levantine basin. Over the past three decades this region has warmed by ca. 1.5–3.0°C with current winter and summer extremums of 17°C and 31°C, respectively. In this study, we tested the response of the native abundant articulated coralline red alga Ellisolandia elongata to this warming. Coralline algae play a key role in coastal ecosystems by structuring marine habitats, providing shelter for a myriad of species, and substantially influencing the coastal carbon budget. Despite being ubiquitous along the Levantine coasts, coralline's ecology, physiology, and biogeochemical role are nearly unknown as well as their performance under different temperatures. Measurements of primary production, respiration and calcification in the temperatures range 15–35°C, which represent past, present and predicted local annual conditions, indicated two physiological tipping points: 1) metabolic breakdown above 31°C; 2) metabolic shift at 23°C, possibly promoting seasonal algal heterotrichy (perennation of the alga without its fronds). Annual production rates were evaluated under the current and predicted temperature regimes indicating a loss of ca. one third of the organic carbon and carbonate production by corallines contributed to the shallow Levantine coast in the upcoming decades. We predict that with continued warming, Eastern Mediterranean corallines will experience a westward range contraction, initiating with phenological shifts, followed by performance declines and population decreases, ending with local extinctions.
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  • 21
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 61 (5). pp. 1661-1676.
    Publikationsdatum: 2019-09-23
    Beschreibung: We present a comparison of the dissolved stable isotope composition of silicate (δ30Si(OH)4) and nitrate (δ15 inline image) to investigate the biogeochemical processes controlling nutrient cycling in the upwelling area off Peru, where one of the globally largest Oxygen Minimum Zones (OMZs) is located. Besides strong upwelling of nutrient rich waters mainly favoring diatom growth, an anticyclonic eddy influenced the study area. We observe a tight coupling between the silicon (Si) and nitrogen (N) cycles in the study area. Waters on the shelf showed high Si(OH)4 concentrations accompanied by diminished inline image concentration as a consequence of intense remineralization, high Si fluxes from the shelf sediments, and N-loss processes such as anammox/denitrification within the OMZ. Correspondingly, the surface waters show low δ30Si(OH)4 values (+2‰) due to low Si utilization but relatively high δ15 inline image (+13‰) values due to upwelling of waters influenced by N-loss processes. In contrast, as a consequence of the deepening of the thermocline in the eddy center, a pronounced Si(OH)4 depletion led to the highest δ30Si(OH)4 values (+3.7‰) accompanied by high δ15 inline image values (+16‰). In the eddy center, high inline image: Si(OH)4 ratios favored the growth of non-siliceous organisms (Synechococcus). Our data show that upwelling processes and the presence of eddies play important roles controlling the nutrient cycles and therefore also exert a major influence on the phytoplankton communities in the Peruvian Upwelling. Our findings also show that the combined approach of δ30Si(OH)4 and δ15 inline image can improve our understanding of paleo records as it can help to disentangle utilization and N-loss processes.
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  • 22
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 48 (5). pp. 1903-1912.
    Publikationsdatum: 2014-01-30
    Beschreibung: According to a recent dynamical model, the depth of a well-mixed water column should have contrasting effects on the abundances of sinking and nonsinking phytoplankton taxa. Because of increasing light limitation, nonsinking taxa should decline monotonically with increasing mixing depth, and because of sinking loss limitation at low mixing depths, sinking taxa should peak at intermediate mixing depths. Along a gradient of mixing depths, the position of this maximum should increase with increasing taxon-specific sinking velocity and decrease with increasing background turbidity. In two field-enclosure experiments, we investigated the effects of mixing depth and background turbidity on a variety of sinking and nonsinking phytoplankton taxa. We exposed the natural, 100-µm screened phytoplankton community of a clear, unproductive, but silica-rich lake to a gradient of mixing depths (1.5- 15 m) during 4-6 weeks. To mimic two different background turbidities, the transparent enclosure walls were surrounded by either white or black foliage. Although diatoms suffered from high sedimentation losses at low mixing depths, they dominated biomass at all mixing depths throughout both experiments. Results were largely in accordance with model predictions. Specific gross growth rates of most common taxa were negatively related to mixing depth. In both experiments, the abundances of most sinking taxa showed a unimodal pattern along the mixing depth gradient, while two of three motile taxa declined monotonically with mixing depth. The depths where these taxa reached their maximal abundances were positively related to taxon-specific sinking velocity and negatively related to background turbidity.
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  • 23
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 48 (1). pp. 179-188.
    Publikationsdatum: 2014-01-30
    Beschreibung: The uptake and efflux of 64Cu was studied in the marine cyanobacterium Synechoccous strain WH7803 (DC2). Uptake followed classical Michaelis-Menten type kinetics in metal-buffered seawater. The maximum uptake rate, Vmax, was 0.236 ± 0.016 × 10-18 mol Cu cell-1 h-1, with the half-saturation constant, KS, of 10-10.81±0.11 mol L-1. An efflux mechanism was also observed in WH7803, whose growth was inhibited by high internal Cu concentrations. Efflux of Cu enabled WH7803 to maintain homeostasis for Cu at typical seawater ambient free copper concentrations ([Cu2+]f). The sensitivity of WH7803 growth to Cu was related to a simple inability to regulate internal Cu concentrations when external concentrations were 〉10-11 mol L-1.
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  • 24
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 46 . pp. 1378-1391.
    Publikationsdatum: 2014-01-30
    Beschreibung: Rates of cellular uptake of CO2 and HCO3- during steady-state photosynthesis were measured in the marine diatoms Thalassiosira weissflogii and Phaeodactylum tricornutum, acclimated to CO2 partial pressures of 36, 180, 360, and 1,800 ppmv. In addition, in vivo activity of extracellular (eCA) and intracellular (iCA) carbonic anhydrase was determined in relation to CO2 availability. Both species responded to diminishing CO2 supply with an increase in eCA and iCA activity. In P. tricornutum, eCA activity was close to the detection limit at higher CO2 concentrations. Simultaneous uptake of CO2 and HCO3- was observed in both diatoms. At air-equilibrated CO2 levels (360 ppmv), T. weissflogii took up CO2 and HCO3- at approximately the same rate, whereas CO2 uptake exceeded HCO3- uptake by a factor of two in P. tricornutum. In both diatoms, CO2 :HCO3- uptake ratios progressively decreased with decreasing CO2 concentration, whereas substrate affinities of CO2 and HCO3- uptake increased. Half-saturation concentrations were always 〈=5 mM CO2 for CO2 uptake and 〈700 mM HCO3- for HCO3- uptake. Our results indicate the presence of highly efficient uptake systems for CO2 and HCO3- in both diatoms at concentrations typically encountered in ocean surface waters and the ability to adjust uptake rates to a wide range of inorganic carbon supply.
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  • 25
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 46 . pp. 964-970.
    Publikationsdatum: 2014-01-30
    Beschreibung: Redfield ratios of remineralization are calculated based on chemical data analysis on isopycnal surfaces. The concentrations of dissolved inorganic carbon used in this study were corrected for the anthropogenic CO2 content as estimated with a back-calculation technique. The corrections increased the apparent carbon remineralization by 25-30%, thus proving important for the reliable estimation of Redfield carbon ratios in the presence of anthropogenic CO2. Best estimates from this study largely confirm the more recently published Redfield ratios of remineralization. The following results were obtained for the latitude range 3-41°N along 20-29°W in the Northeast Atlantic Ocean: Corg: P ratio = 123 ± 10; Corg : N ratio = 7.2 ± 0.8; -O2 :Corg ratio = 1.34 ± 0.06; -O2 : P ratio = 165 ± 15; N: P ratio = 17.5 ± 2.0. These ratios are in close agreement with the average composition of phytoplankton and represent respiration of organic matter consisting on average of 52% protein, 36% polysaccharide, and 12% lipid.
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  • 26
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 46 . pp. 749-757.
    Publikationsdatum: 2018-06-25
    Beschreibung: In contrast to most pelagic primary producers, benthic macrophytes pass through morphologically distinct life stages, which can be subject to different ecological controls. Using factorial field experiments, we investigated how grazing pressure (three levels) and nutrient supply (four levels) interact in controlling the passage of marine macroalgae through an apparent recruitment bottleneck at the germling stage. In comparative experiments, we asked whether relative bottom-up and top-down effects on early life stages (〈4 week germlings) vary (1) between the eutrophic Baltic Sea and the oligotrophic NW Atlantic, (2) across seasons in the NW Atlantic, and (3) among annual and perennial macroalgae. In both systems nutrient enrichment favored and grazers suppressed recruitment of green and brown annual algae; however, enrichment effects were much more pronounced in the Baltic, whereas grazer effects dominated in the NW Atlantic. Grazers induced a shift from grazer-susceptible green to more resistant brown algae in the Baltic without reducing total germling density. In the NW Atlantic, grazers strongly reduced overall recruitment rate throughout all seasons. Effects on perennials were similar in both systems with moderate losses to grazing and no effects of nutrient enrichment. Recruit densities and species composition shifted with season in the NW Atlantic. We conclude that the relative effects of grazers and nutrient enrichment depended on the nutrient status of the system, algal life history strategy, and season. Strong bottom-up and top-down controls shape benthic community composition before macroalgae reach visible size
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  • 27
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 47 (3). pp. 753-761.
    Publikationsdatum: 2014-01-30
    Beschreibung: Flows of the major biogeochemical elements (C, N, P, Si) and of transparent exopolymer particles (TEP) were traced during a bloom of a natural assemblage of marine diatoms in a mesocosm (l m(3)) to determine whether the exudation and subsequent gelation of carbon-rich phytoplankton exopolymers can account for the formation and potential export of carbon in excess of that predicted by Redfield ratios. Exponential growth of the phytoplankton community in the mesocosm extended for 10 d until nitrate concentration fell below detection and concentrations of dissolved inorganic and particulate organic nitrogen and phosphorus remained stable. Tight covariation of particulate organic elements occurred as long as nutrients were replete. But, after nitrate depletion, decoupling of carbon dynamics from that of nitrogen and phosphorus was observed, with a large flow of carbon into TEP An uptake of 72% more dissolved inorganic carbon (DIC) than inferred from nitrate supply and Redfield stoichiometry (referred to as carbon overconsumption) occurred during the study, largely during the postbloom phase, and was almost entirely traced to the particulate organic matter (POM) pool. Marine snow (aggregates 〉0.5 mm) appeared at the onset of nitrate depletion and coincided with rapid increase in TEP concentrations. Elemental composition of marine snow differed from the Redfield ratio by an enrichment in carbon and a depletion in phosphorus relative to nitrogen. It is suggested that sinking of TEP-rich marine snow could be a possible mechanism for export of carbon above calculations that are based on the Redfield stoichiometry.
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  • 28
    Publikationsdatum: 2019-09-24
    Beschreibung: Large quantities of methane are stored in hydrates and permafrost within shallow marine sediments in the Arctic Ocean. These reservoirs are highly sensitive to climate warming, but the fate of methane released from sediments is uncertain. Here, we review the principal physical and biogeochemical processes that regulate methane fluxes across the seabed, the fate of this methane in the water column, and potential for its release to the atmosphere. We find that, at present, fluxes of dissolved methane are significantly moderated by anaerobic and aerobic oxidation of methane. If methane fluxes increase then a greater proportion of methane will be transported by advection or in the gas phase, which reduces the efficiency of the methanotrophic sink. Higher freshwater discharge to Arctic shelf seas may increase stratification and inhibit transfer of methane gas to surface waters, although there is some evidence that increased stratification may lead to warming of sub-pycnocline waters, increasing the potential for hydrate dissociation. Loss of sea-ice is likely to increase wind speeds and seaair exchange of methane will consequently increase. Studies of the distribution and cycling of methane beneath and within sea ice are limited, but it seems likely that the sea-air methane flux is higher during melting in seasonally ice-covered regions. Our review reveals that increased observations around especially the anaerobic and aerobic oxidation of methane, bubble transport, and the effects of ice cover, are required to fully understand the linkages and feedback pathways between climate warming and release of methane from marine sediments.
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  • 29
    Publikationsdatum: 2019-02-01
    Beschreibung: Large quantities of the greenhouse gas methane (CH4) are stored in the seafloor. The flux of CH4 from the sediments into the water column and finally to the atmosphere is mitigated by a series of microbial methanotrophic filter systems of unknown efficiency at highly active CH4-release sites in shallow marine settings. Here, we studied CH4-oxidation and the methanotrophic community at a high-CH4-flux site in the northern North Sea (well 22/4b), where CH4 is continuously released since a blowout in 1990. Vigorous bubble emanation from the seafloor and strongly elevated CH4 concentrations in the water column (up to 42 µM) indicated that a substantial fraction of CH4 bypassed the highly active (up to ∼2920 nmol cm−3 d−1) zone of anaerobic CH4-oxidation in sediments. In the water column, we measured rates of aerobic CH4-oxidation (up to 498 nM d−1) that were among the highest ever measured in a marine environment and, under stratified conditions, have the potential to remove a significant part of the uprising CH4 prior to evasion to the atmosphere. An unusual dominance of the water-column methanotrophs by Type II methane-oxidizing bacteria (MOB) is partially supported by recruitment of sedimentary MOB, which are entrained together with sediment particles in the CH4 bubble plume. Our study thus provides evidence that bubble emission can be an important vector for the transport of sediment-borne microbial inocula, aiding in the rapid colonization of the water column by methanotrophic communities and promoting their persistence close to highly active CH4 point sources.
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  • 30
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 62 (1). pp. 334-347.
    Publikationsdatum: 2020-02-06
    Beschreibung: Marine phytoplankton is simultaneously affected by multiple environmental drivers. To-date integrative assessments of multiple combined effects are rare on the relationship between elemental stoichiometry and biochemicals in marine phytoplankton. We investigated responses of stoichiometric (N:C and P:C ratios) and fatty acid-based (polyunsaturated fatty acid, PUFA) indicators of nutritional quality to three N:P supply ratios (10:1, 24:1, and 63:1 mol mol−1), three temperatures (12, 18, and 24°C) and two pCO2 levels (560 and 2400 μatm) in the marine phytoplankters Rhodomonas sp. and Phaeodactylum tricornutum. Overall, warming and nutrient deficiency showed dramatic effects, but increased pCO2 had modest effects on the two indicators of nutritional quality. Specifically, warming showed strong positive effects on N:C and P:C ratios in Rhodomonas sp. but negative effects on PUFAs in both species. The low N- and low P-media led to low contents of both nutrients but high contents of PUFAs in the biomass of Rhodomonas sp., while the response of P. tricornutum was more complex: N:C ratios were lowest at the intermediate N:P supply but P:C ratios responded negatively to P deficiency and positively to N deficiency. Large variations in the two indicators of nutritional quality can be attributed to species-specific physiological optima and interactions between the three manipulated variables. Our results suggest that stoichiometric and FA-based indicators of nutritional quality may change differentially in response to warming and nutrient deficiency in marine phytoplankton, highlighting the relevance of simultaneous considerations of the two indicators of nutritional quality, when assessing food web dynamics under future ocean scenarios.
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  • 31
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 61 (5). pp. 1891-1899.
    Publikationsdatum: 2019-02-01
    Beschreibung: Coastal marine ecosystems have been under high anthropogenic pressure and it can be assumed that prevalent local perturbation interacts with rising global stressors under proceeding climate change. Understanding their effective pathways and cumulative effects is of high relevance not only with regard to future risk assessment, but also for current ecosystem management. In benthic mesocosms, we factorially tested the effects of one global (combined elevated seawater temperature and CO2 concentration) and one local (nutrient enrichment) stressor on a common coastal Baltic seaweed system (Fucus vesiculosus). Both treatments in combination had additive negative impacts on the seaweed—epiphyte—mesograzer system by altering its regulatory mechanisms. That is, warming decreased the biomass of two mesograzer species (weakened top-down control), whereas moderate nutrient enrichment increased epiphyte biomass (intensified bottom-up control), which ultimately resulted in a significant biomass reduction of the foundation seaweed. Our results suggest that climate change impacts might be underestimated if local pressures are disregarded. Furthermore, they give implication for local ecological management as the mitigation of local perturbation may limit climate change impacts on marine ecosystems.
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  • 32
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 49 (1). pp. 168-179.
    Publikationsdatum: 2019-03-06
    Beschreibung: We assess population dynamics of picophytoplankton groups (≤2 μm diameter; Prochlorococcus, Synechococcus, and picoeukaryote) at a Pacific Ocean coastal site in the Southern California Bight. Weekly sampling (August 2000 to January 2002), dilution experiments, and flow cytometric analysis were combined with an instrument-specific calibration for cell size determination, allowing biovolume and carbon biomass estimation. Synechococcus was almost always numerically dominant, accounting for 60 ± 12 of the total picoplankton cells over time. It had moderately high growth rates (0.52-0.86 d-1) and was subject to low grazing mortality (-0.14 to -0.39 d-1). Prochlorococcus growth and mortality rates were roughly balanced (0.33 ± 0.14 d-1 and -0.36 ± 0.06 d-1, respectively). Picoeukaryotes had the highest growth rates (0.71-1.29 d-1) and were responsible for, on average, 76 of net carbon production (NCP), amounting in up to 32.05 ± 1.31 μg C L-1 d-1 produced and 28.31 ± 2.61 μg C L-1 d-1 consumed. In order to better define the eukaryotic component of these populations, an isolate was characterized via small subunit rRNA gene sequencing, transmission electron microscopy, and growth experiments and was identified as the prasinophyte Ostreococcus, not previously known to the Pacific Ocean. Our results show that although picoeukaryotes do not stand out as particularly important players in this system on the basis of cell abundance, they dominate in terms of picophytoplankton biomass and trophic transfer potential of carbon in this size class.
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  • 33
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 38 (8). pp. 1803-1812.
    Publikationsdatum: 2019-09-23
    Beschreibung: Sediments from the River Elbe estuary and incubated sediments were extracted with 1 N HCI for 24 h at room temperature. The extracted ferric and ferrous iron was determined with DC and AC polarography. Acid-volatile sulfide was determined from H2S trapped in aqueous zinc acetate solution. Analysis of sediment samples and extraction residues with Mossbauer spectroscopy demonstrated that the Fe oxidation state was conserved during extraction and polarographic determination, siderite and vivianite were completely dissolved, and Fe(II) in chlorite was partially extracted with HCl. Incubation experiments showed that extractable Fe was almost completely oxidized to Fe(III) at the oxic sediment surface and reduced to Fe(II) in deeper anoxic layers within a few weeks. Reactive Fe(III), i.e. that fraction of Fe which was reducible on the time scale of the incubation experiment, was completely extracted with HCl.
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  • 34
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 62 (1). pp. 75-88.
    Publikationsdatum: 2020-02-06
    Beschreibung: Western boundary currents support high primary production and carbon export. Here, we performed a survey of photosynthetic picoeukaryotes in the North Pacific Ocean in four transects crossing the Kuroshio Front. Prasinophyte algae comprised 85% of 18S rRNA gene sequences for photosynthetic taxa in the 〈5 μm size fraction. The picoplanktonic (〈2 μm) genera Micromonas and Ostreococcus comprised 30% and 51% of the total photosynthetic 18S rDNA sequences from five stations. Phylogenetic analysis showed that two Ostreococcus ecotypes, until now rarely found to co‐occur, were both present in the majority of samples. Ostreococcus ecotype OI reached 6,830 ± 343 gene copies mL−1, while Ostreococcus ecotype OII reached 50,190 ± 971 gene copies mL−1 based on qPCR analysis of the 18S rRNA gene. These values are higher than in studies of other oceanographic regions by a factor of 10 for OII. The data suggest that meso‐ and finer‐scale physical dynamics had a significant impact on the populations at the front, either by mingling ecotypes from different source regions at fine scales (∼10s km) or by stimulating their growth through vertical nutrient injections. We investigate this hypothesis with an idealized diffusion‐reaction model, and find that only a combination of mixing and positive net growth can explain the observed distributions and overlap of the two Ostreococcus ecotypes. Our field observations support larger‐scale numerical ocean simulations that predict enhanced biodiversity at western boundary current fronts, and suggest a strategy for systematically testing that hypothesis.
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  • 35
    Publikationsdatum: 2021-05-19
    Beschreibung: Gelatinous zooplankton hold key functions in the ocean and have been shown to significantly influence the transport of organic carbon to the deep sea. We discovered a gelatinous, flux‐feeding polychaete of the genus Poeobius in very high abundances in a mesoscale eddy in the tropical Atlantic Ocean, where it co‐occurred with extremely low particle concentrations. Subsequent analysis of an extensive in situ imaging dataset revealed that Poeobius sp. occurred sporadically between 5°S–20°N and 16°W–46°W in the upper 1000 m. Abundances were significantly elevated and the depth distribution compressed in anticyclonic modewater eddies (ACMEs). In two ACMEs, high Poeobius sp. abundances were associated with strongly reduced particle concentrations and fluxes in the layers directly below the polychaete. We discuss possible reasons for the elevated abundances of Poeobius sp. in ACMEs and provide estimations showing that a single zooplankton species can completely intercept the downward particle flux by feeding with their mucous nets, thereby substantially altering the biogeochemical setting within the eddy.
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  • 36
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 47 . pp. 120-128.
    Publikationsdatum: 2014-01-30
    Beschreibung: The carbon isotopic composition of marine phytoplankton varies significantly with growth conditions. Aqueous CO2 concentration [CO2] and algal growth rate (µ) have been suggested to be important factors determining isotope fractionation (ep). Here we examine ep of the coccolithophorid Emiliania huxleyi in relation to CO2 concentration and light conditions in dilute batch cultures. Cells were incubated at different irradiance cycles, photon flux densities (PFDs), and [CO2]. Isotope fractionation varied between 6.7 and 12.3‰ under 16 : 8 h light : dark cycle (L :D) and between 14.7 and 17.8‰ at continuous light. ep was largely independent of ambient [CO2], varying generally by less than 2‰ over a range of [CO2] from 5 to 34 mmol L-1. Instantaneous carbon-specific growth rates (µC) and PFDs, ranging from 15 to 150 mmol m-2 s-1, positively correlated with ep. This result is inconsistent with theoretical considerations and experimental results obtained under constant light conditions, suggesting an inverse relationship between ep and µ. In the present study the effect of PFDs on ep was stronger than that of mand thus resulted in a positive relationship between µ and ep. In addition, the L:D cycle of 16 : 8 h resulted in significantly lower ep values compared to continuous light. Since the observed offset of about 8‰ could not be related to daylength dependent changes in µC, this implies a direct influence of the irradiance cycle on ep. These findings are best explained by invoking active carbon uptake in E. huxleyi. If representative for the natural environment, these results complicate the interpretation of carbon isotope data in geochemical and paleoceanographic applications.
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  • 37
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 46 (7). pp. 1802-1808.
    Publikationsdatum: 2018-06-25
    Beschreibung: Iron supply is thought to regulate primary production in high nitrate, low chlorophyll (HNLC) regions of the sea in both the past and the present. A critical aspect of this relationship is acquisition of iron (Fe) by phytoplankton, which occurs through a complex series of extracellular reactions that are influenced by Fe chemistry and speciation. During the first in situ mesoscale Fe-enrichment experiment in the Southern Ocean (Southern Ocean iron release experiment [SOIREE]), we monitored the uptake of Fe by three size classes of plankton and their ensuing physiological response to the Fe enrichment. Rates of Fe uptake from both inorganic Fe (Fe') and organic Fe complexes (FeL) were initially fast, indicative of Fe-limitation. After Fe enrichment phytoplankton down-regulated Fe uptake and optimized physiological performance, but by day 12 they had greatly increased their capacity to acquire Fe from FeL. The increase in Fe uptake from FeL coincided with a sixfold decrease in Fe' that followed the production of Fe-binding organic ligands. Phytoplankton were able to use organically bound Fe at rates sufficient to maintain net growth for more than 42 d. Adaptation to such shifts in Fe chemistry may contribute to bloom longevity in these polar HNLC waters.
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  • 38
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 48 . pp. 55-67.
    Publikationsdatum: 2014-01-30
    Beschreibung: Carbon acquisition in relation to CO2 supply was investigated in three marine bloom-forming microalgae, the diatom Skeletonema costatum, the flagellate Phaeocystis globosa, and the coccolithophorid Emiliania huxleyi. In vivo activities of extracellular (eCA) and intracellular (iCA) carbonic anhydrase activity, photosynthetic O2 evolution, CO2 and HCO uptake rates were measured by membrane inlet mass spectrometry in cells acclimated to pCO2 levels of 36, 180, 360, and 1,800 ppmv. Large differences were obtained between species both with regard to the efficiency and regulation of carbon acquisition. While eCA activity increased with decreasing CO2 concentration in S. costatum and P. globosa, consistently low values were obtained for E. huxleyi. No clear trends with pCO2 were observed in iCA activity for any of the species tested. Half saturation concentrations (K1/2) for photosynthetic O2 evolution, which were highest for E. huxleyi and lowest for S. costatum, generally decreased with decreasing CO2 concentration. In contrast, K1/2 values for P. globosa remained unaffected by pCO2 of the incubation. CO2 and HCO3- were taken up simultaneously by all species. The relative contribution of HCO3- to total carbon uptake generally increased with decreasing CO2, yet strongly differed between species. Whereas K1/2 for CO2 and HCO3- uptake was lowest at the lowest pCO2 for S. costatum and E. huxleyi, it did not change as a function of pCO2 in P. globosa. The observed taxon-specific differences in CO2 sensitivity, if representative for the natural environment, suggest that changes in CO2 availability may influence phytoplankton species succession and distribution. By modifying the relative contribution of different functional groups, e.g., diatomaceous versus calcareous phytoplankton, to the overall primary production this could potentially affect marine biogeochemical cycling and air-sea gas exchange.
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  • 39
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 48 (2). pp. 764-776.
    Publikationsdatum: 2014-01-30
    Beschreibung: The chemical speciation of dissolved Cu was investigated by voltammetric methods in Gullmar Fjord, Sweden, over the course of a year from September 1996 until August 1997. Sampling was carried out on a roughly monthly basis, with an intensive survey carried out in May 1997. Surface water temperatures ranged from 21 to 22°C, whereas bottom waters in the fjord were approximately 6°C throughout. Macronutrient concentrations in the fjord during the period of the survey were investigated independently by the Göteborgs och Bohus läns Vattenvårdsförbund (Water Quality Association of Göteborg and Bohus). Surface phosphate concentrations were highest in early spring with low levels (〈0.1 mmol kg-1) over the late spring and summer. Nitrate and silicate showed a similar pattern to phosphate with the exception of high concentrations encountered in surface waters when low salinity plumes caused by runoff were encountered. A period of calm, sunny weather in January 1997 saw the initiation of the spring bloom some 2 months earlier than usual. Dissolved Cu speciation was dominated by organic complexation (over 99.8%) throughout this study. Strong Cu binding ligands (log K 〉 12.5) were not detected during the winter or early spring and could be related to the temperature-related seasonal appearance of the cyanobacterium Synechoccocus in these waters. The appearance of the strong Cu ligands led to a decrease in the concentration of free copper, resulting in a seasonal cycle for free copper in the fjord. This is the first study to examine Cu speciation over an annual cycle in a coastal environment
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  • 40
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 46 . pp. 497-504.
    Publikationsdatum: 2014-01-30
    Beschreibung: Blooms of the marine diatom Skeletonema costatum were initiated in closed-system batch cultures with P-deficient medium under two different initial concentrations of dissolved molecular CO2([CO2,aq]: 20.6 and 4.5 µmol L-1). Algal C: N: P ratios strongly increased with decreasing P concentration. In the exponential growth phase, C: N ratios were 1.3 mol mol-1 higher in the low relative to the high [CO2,aq] treatment. There was no [CO2,aq] effect on C: N: P ratios during P-limited growth. Carbon isotope fractionation («p ) was 2-3‰ higher in the high [CO2,aq] treatment. With growth rate decreasing due to P limitation, ep increased in both [CO2,aq] treatments by 2-3‰ despite decreasing [CO2,aq]. Under these conditions the effect of decreasing growth rate on isotope fractionation strongly dominated over that of declining CO2 availability. When extrapolated to the natural environment, these results imply that systematic changes in algal growth, as occurring during the course of phytoplankton blooms, may affect algal isotope fractionation. These results severely complicate the interpretation of carbon isotope measurements in suspended and sedimentary organic matter
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  • 41
    Publikationsdatum: 2018-06-25
    Beschreibung: Although N2-fixing cyanobacteria contribute significantly to oceanic sequestration of atmospheric CO2, little is known about how N2 fixation and carbon fixation (primary production) interact in natural populations of marine cyanobacteria. In a developing cyanobacterial bloom in the Baltic Sea, rates of N2 fixation (acetylene reduction) showed both diurnal and longer-term fluctuations. The latter reflected fluctuations in the nitrogen status of the cyanobacterial population and could be correlated with variations in the ratio of acetylene reduced to 15N2 assimilated. The value of this ratio may provide useful information about the release of newly fixed nitrogen by a cyanobacterial population. However, although the diurnal fluctuations in N2 fixation broadly paralleled diurnal fluctuations in carbon fixation, the longer-term fluctuations in these two processes were out of phase.
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  • 42
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    Unbekannt
    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 45 (2). pp. 339-349.
    Publikationsdatum: 2018-06-25
    Beschreibung: By factorial field experiments we analyzed the relative effects of increased nutrient (N+P) loading and natural grazing pressure on species composition, carbon storage, and nitrogen retention in the Baltic Sea littoral food web, composed of macroalgae, grazers (snails, isopods, amphipods), and predators (shrimps, crabs, fish). Nitrogen was depleted relative to phosphorus throughout most of the year. Increasing nitrogen (6–200% over ambient concentrations) enhanced algal productivity and cover of fast-growing annual algae, grazer, and predator densities, suggesting a three-level bottom-up effect. With increasing nitrogen loading, annual algae increasingly blocked perennial algal recruitment (65–98% decrease) and growth. Grazers counteracted the effects of nutrient enrichment on algal species composition through selective consumption of annual algae. Grazer exclusion had equivalent negative effects on perennial recruitment as a 85% increase in nitrogen loading. Nutrient enrichment increased algal nitrogen content and decreased tissue C: N ratios in spring and summer but not in fall. Carbon storage and nitrogen retention, measured as C and N retained in plant biomass at the end of the growth season, were increased by grazers (C: 39%, N: 24%) but decreased with increasing nitrogen loading (C: -71%, N: -74%). Our results emphasize the important role of grazers in buffering moderate eutrophication effects and illustrate how food web interactions and shifts in species composition are tightly linked to coastal ecosystem function
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  • 43
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    ASLO (Association for the Sciences of Limnology and Oceanography)
    In:  Limnology and Oceanography, 61 (2). pp. 666-683.
    Publikationsdatum: 2019-02-01
    Beschreibung: A multitracer approach is applied to assess the impact of boundary fluxes (e.g., benthic input from sediments or lateral inputs from the coastline) on the acid-base buffering capacity, and overall biogeochemistry, of the North Sea. Analyses of both basin-wide observations in the North Sea and transects through tidal basins at the North-Frisian coastline, reveal that surface distributions of the δ13C signature of dissolved inorganic carbon (DIC) are predominantly controlled by a balance between biological production and respiration. In particular, variability in metabolic DIC throughout stations in the well-mixed southern North Sea indicates the presence of an external carbon source, which is traced to the European continental coastline using naturally occurring radium isotopes (224Ra and 228Ra). 228Ra is also shown to be a highly effective tracer of North Sea total alkalinity (AT) compared to the more conventional use of salinity. Coastal inputs of metabolic DIC and AT are calculated on a basin-wide scale, and ratios of these inputs suggest denitrification as a primary metabolic pathway for their formation. The AT input paralleling the metabolic DIC release prevents a significant decline in pH as compared to aerobic (i.e., unbuffered) release of metabolic DIC. Finally, long-term pH trends mimic those of riverine nitrate loading, highlighting the importance of coastal AT production via denitrification in regulating pH in the southern North Sea.
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  • 44
    Publikationsdatum: 2022-01-31
    Beschreibung: The subterranean flow of water through sand barriers between coastal lagoons and the sea, driven by a positive hydraulic gradient, is a net new pathway for solute transfer to the sea. On the sea side of sand barriers, seawater circulation in the swash-zone generates a flux of recycled and new solutes. The significance and temporal variability of these vectors to the French Mediterranean Sea is unknown, despite lagoons constituting ~ 50% of the coastline. A one-dimensional Ra-224(ex)/Ra-223 reactive-transport model was used to quantify water flow between a coastal lagoon (La Palme) and the sea over a 6-month period. Horizontal flow between the lagoon and sea decreased from ~ 85 cm d(-1) during May 2017 (0.3 m(3) d(-1) m(-1) of shoreline) to ~ 20 cm d(-1) in July and was negligible in the summer months thereafter due to a decreasing hydraulic gradient. Seawater circulation in the swash-zone varied from 10 to 52 cm d(-1) (0.4-2.1 m(3) d(-1) m(-1)), driven by short-term changes in the prevailing wind and wave regimes. Both flow paths supply minor dissolved silica fluxes on the order of ~ 3-10 mmol Si d(-1) m(-1). Lagoon-sea water exchange supplies a net dissolved inorganic carbon (DIC) flux (320-1100 mmol C d(-1) m(-1)) two orders of magnitude greater than seawater circulation and may impact coastal ocean acidification. The subterranean flow of water through sand barriers represents a significant source of new DIC, and potentially other solutes, to the Mediterranean Sea during high lagoon water-level periods and should be considered in seasonal element budgets.
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  • 45
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    ASLO (Association for the Sciences of Limnology and Oceanography) | Wiley
    In:  Limnology and Oceanography, 64 (2). pp. 661-678.
    Publikationsdatum: 2022-01-31
    Beschreibung: The trait‐based approach is increasingly used in plankton ecology to understand diversity, community dynamics, and biogeography. While on the global scale phytoplankton traits are fairly well established, zooplankton traits are only beginning to be understood. One taxa‐transcending aspect of zooplankton diversity is the distinction between ambush and active feeding strategies. We present a global‐scale empirical estimate of feeding strategy derived from copepod abundance observations, which for the first time suggests a distinct trait biogeography with ambush feeding as the dominant feeding strategy at higher, but not at lower latitudes. To explain this trait biogeography, we develop a minimalist trade‐off based model of feeding strategies based on encounter rates between zooplankton predators and their phyto‐ and zooplankton prey. Encounter rates are governed by the two traits, size and motility, that trade off against predation risk. Coupled to a three‐dimensional dynamic green ocean model, our idealized encounter model captures the observed feeding strategy biogeography. In the model, this pattern arises from competing dominant food chains within the food web and is shaped by a trophic trait cascade of active vs. passive feeding in adjacent trophic levels. The dominant feeding strategy structures the pathways and efficiency of energy and biomass transfer through the model food web, with consequences for primary production, export and higher trophic levels. Understanding feeding strategies is therefore important for fisheries, biogeochemical cycling, and long‐term predictions of ecosystem dynamics and functioning by global dynamic green ocean models.
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  • 46
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    ASLO (Association for the Sciences of Limnology and Oceanography) | Wiley
    In:  Limnology and Oceanography, 64 (1). pp. 302-316.
    Publikationsdatum: 2022-01-31
    Beschreibung: Coastal seas like the North Sea have been subject to major changes in nutrient inputs over the last decades, resulting in shifts of limiting nutrients for phytoplankton communities. Here, we investigated the seasonal and spatial distribution and synthesis patterns of individual amino acids and distinct fatty acid groups and show how these were affected by different nutrient limitations in natural coastal phytoplankton communities. Nitrogen limited communities exhibited substantially slower synthesis of essential amino acids compared to synthesis of nonessential amino acids. In short‐term nutrient addition experiments, this trend was reversed immediately after N addition to levels found under not limiting conditions. On the contrary, phosphorus limited communities showed no such shift in amino acids. Both N and P limitation induced a shift from structural to storage fatty acids with a concurrent decrease in the synthesis of poly‐unsaturated fatty acids. Reversed effects in fatty acid synthesis after N or P addition were only apparent after 72 h, when they could be found in both fatty acid biosynthesis and concentrations. The different strategies of qualitative and quantitative regulation of different biomolecule synthesis under nutrient scarcity may have far‐reaching consequences for the phytoplankton's nutritional value. Higher trophic levels may have to cope with the loss of essential amino acids and poly‐unsaturated fatty acids in nutrient limited phytoplankton, which could induce changes in the structure of food webs.
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  • 47
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    ASLO (Association for the Sciences of Limnology and Oceanography) | Wiley
    In:  Limnology and Oceanography, 64 (5). pp. 1883-1894.
    Publikationsdatum: 2022-01-31
    Beschreibung: Abyssal polymetallic nodule fields constitute an unusual deep-sea habitat. The mix of soft sediment and the hard substratum provided by nodules increases the complexity of these environments. Hard substrata typically support a very distinct fauna to that of seabed sediments, and its presence can play a major role in the structuring of benthic assemblages. We assessed the influence of seafloor nodule cover on the megabenthos of a marine conservation area (area of particular environmental interest 6) in the Clarion Clipperton Zone (3950–4250 m water depth) using extensive photographic surveys from an autonomous underwater vehicle. Variations in nodule cover (1–20%) appeared to exert statistically significant differences in faunal standing stocks, some biological diversity attributes, faunal composition, functional group composition, and the distribution of individual species. The standing stock of both the metazoan fauna and the giant protists (xenophyophores) doubled with a very modest initial increase in nodule cover (from 1% to 3%). Perhaps contrary to expectation, we detected little if any substantive variation in biological diversity along the nodule cover gradient. Faunal composition varied continuously along the nodule cover gradient. We discuss these results in the context of potential seabed-mining operations and the associated sustainable management and conservation plans. We note in particular that successful conservation actions will likely require the preservation of areas comprising the full range of nodule cover and not just the low cover areas that are least attractive to mining.
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