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Changes in the complexity of planktonic food webs may be expected in future aquatic systems due to increases in sea surface temperature and an enhanced stratification of the water column. Under these conditions, the growth of unpalatable, filamentous, N2‐fixing cyanobacterial blooms, and their effect on planktonic food webs will become increasingly important. The planktonic food web structure in aquatic ecosystems at times of filamentous cyanobacterial blooms is currently unresolved, with discordant lines of evidence suggesting that herbivores dominate the mesozooplankton or that mesozooplankton organisms are mainly carnivorous. Here, we use a set of proxies derived from amino acid nitrogen stable isotopes from two mesozooplankton size fractions to identify changes in the nitrogen source and the planktonic food web structure across different microplankton communities. A transition from herbivory to carnivory in mesozooplankton between more eutrophic, near‐coastal sites and more oligotrophic, offshore sites was accompanied by an increasing diversity of microplankton communities with aging filamentous cyanobacterial blooms. Our analyses of 124 biotic and abiotic variables using multivariate statistics confirmed salinity as a major driver for the biomass distribution of non‐N2‐fixing microplankton species such as dinoflagellates. However, we provide strong evidence that stratification, N2 fixation, and the stage of the cyanobacterial blooms regulated much of the microplankton diversity and the mean trophic position and size of the metabolic nitrogen pool in mesozooplankton. Our empirical, macroscale data set consistently unifies contrasting results of the dominant feeding mode in mesozooplankton during blooms of unpalatable, filamentous, N2‐fixing cyanobacteria by identifying the at times important role of heterotrophic microbial food webs. Thus, carnivory, rather than herbivory, dominates in mesozooplankton during aging and decaying cyanobacterial blooms with hitherto uncharacterized consequences for the biogeochemical functions of mesozooplankton. R/V Meteor passing a decaying bloom of unpalatable, filamentous, N2‐fixing cyanobacteria in the Baltic Sea in August 2015 that are expected to increase in large areas of the global ocean in the future. While herbivory dominated the trophic structure of mesozooplankton in earlier bloom stages, direct estimates of carnivory in field mesozooplankton identified the at times important role of heterotrophic microbial food webs namely in such aging and decaying cyanobacterial blooms with hitherto uncharacterized consequences for the biogeochemical functions of mesozooplankton. Photo: Andreas Raeke, DWD. [ABSTRACT FROM AUTHOR]

Abstract: Cyanobacteria are the main autotrophs and N2‐fixing (diazotrophic) organisms in large parts of the oligotrophic global ocean, where generally all heterotrophic production depends on their activity. Amino acids (AAs) from cyanobacteria are essential macronutrients for these heterotrophic food webs, yet little is known about the de novo synthesis of AAs during N2 fixation. Through a combination of bulk and amino acid nitrogen (AAN) specific analyses of field based N2 fixation experiments, we demonstrate that the de novo synthesis of 13 AAs accounted for the majority of bulk N2 fixation rates at four stations in the central Baltic Sea in July 2015. Slow AA turnover times of 87 ± 14 d coincided with low phosphate concentrations and high cell‐carbon biomasses of unicellular cyanobacteria. Very fast turnover times of 17 ± 3 d coincided with high phosphate concentrations and undecayed Nodularia spumigena cells, but unexpectedly also with phosphate depletion and decayed N. spumigena cells. In a decayed bloom, volumetric N2 fixation rates into AAN provided a much better estimate of the net incorporation of N2 into biomass than fixation into bulk nitrogen that rather reflected gross N2 fixation. In an undecayed bloom, the turnover times of 13 AAs can be predicted from a single bulk N2 fixation rate. This is the first direct evidence that the very late, decayed stage of a cyanobacteria bloom can be a flashpoint of very fast AA turnover during N2 fixation with hitherto uncharacterized consequences for heterotrophic food webs and diazotroph N inputs to the global ocean. [ABSTRACT FROM AUTHOR]

Investigating microplankton biomass and diversity under different climatological conditions is key to the understanding of cascading effects of climate change on nutrient cycles and biological productivity. Here we have used data collected during two contrasting summers along the coast of Viet Nam to show how climatological-driven changes can have a significant influence on the distribution of microplankton communities and their biomass via its impact on nutrient concentrations in the water column. The first summer in July 2003 followed a weak El-Nino Southern Oscillation (ENSO) event and was characterized by weak coastal upwelling, in the second summer during July 2004, upwelling was normal. Very low silicate (SiO 4 ) concentrations and SiO 4 :DIN ratios characterized the source water mass for upwelling in July 2004, and dynamic SiO 4 to dissolved inorganic nitrogen ratios (SiO 4 :DIN) mainly below the Redfield-Brzezinski ratio and DIN to phosphate ratios (DIN:PO 4 3− ) below the Redfield ratio were a common feature off Viet Nam. Much higher particle concentrations and PSi/PC ratios during normal upwelling revealed major changes in the microplankton community structure among summers. Small dinoflagellates (10–20 µm) prevailed ubiquitously during reduced upwelling. During normal upwelling, the diatom Rhizosolenia sp. dominated the cell-carbon biomass in the silicate poor upwelling waters. Trichodesmium erythraeum dominated in the Mekong-influenced and nutrient depleted offshore waters, where it co-occurred with Rhizosolenia sp. Both species were directly associated with the much higher primary production (PP) and N 2 fixation rates that were quantified in earlier studies, as well as with much higher diversities at these offshore sites. Along the coast, the correlation between Rhizosolenia sp. and PP rates was less clear and the factors regulating the biomass of Rhizosolenia sp. in the upwelling waters are discussed. The very low silicate concentrations in the source water mass for upwelling and the offshore deflection of the Mekong river plume likely triggered the observed ecological differences in the microplankton communities off Viet Nam in the normal upwelling season compared to the post ENSO conditions. [ABSTRACT FROM AUTHOR]

The plasticity of nitrogen specific net growth efficiency (NGE) in marine mesozooplankton is currently unresolved, with discordant lines of evidence suggesting that NGE is constant, or that it varies with nitrogen source, food availability, and food quality in marine ecosystems. Specifically, the fate of nitrogen from nitrogen fixation is poorly known. We use 15N : 14N ratios in plankton in combination with hydrological data, nutrient profiles, and nitrogen fixation rate measurements to investigate the relationship between new nitrogen sources and the nitrogen specific NGE in three plankton communities along the outer Amazon River plume. The NGE of small (200-500 μm) mesozooplankton was estimated from the δ15N differences between particulate nitrogen and zooplankton using an open system Rayleigh fractionation model. The transfer efficiency of nitrogen among larger (> 500 μm) mesozooplankton was estimated from the change in δ15N as a function of zooplankton size. The Amazon River was not a significant source of bioavailable nitrogen anywhere in our study region, and subsurface nitrate was the primary new nitrogen source for the outer shelf community, which was dominated by diatoms. N2 fixation was the principal new nitrogen source at sites of high diatom diazotroph association abundance and at oceanic sites dominated by Trichodesmium spp. and Synechococcus spp. Although we found clear spatial differences in food quantity, food quality, and diazotroph inputs into mesozooplankton, our data show no significant differences in mesozooplankton nitrogen transfer efficiency and NGE (for latter, mean ± SD: 59 ± 10%) among sites. [ABSTRACT FROM AUTHOR]

Abstract: A redefinition of characteristic water masses in the Vietnamese upwelling area is presented based on five interdisciplinary cruises during different seasons and 331 CTD casts. In contrast to the previous definition of Rojana-anawat et al. (2001), we clearly can define water masses which serve as end members of mixing. This new definition is useful for an improved understanding of geographical positions of different water masses with respect to climate variability, the phytoplankton distribution and complex nutrient cycle in this area. [Copyright &y& Elsevier]

Eutrophication, increased temperatures and stratification can lead to massive, filamentous, N2‐fixing cyanobacterial (FNC) blooms in coastal ecosystems with largely unresolved consequences for the mass and energy supply in food webs. Mesozooplankton adapt to not top‐down controlled FNC blooms by switching diets from phytoplankton to microzooplankton, resulting in a directly quantifiable increase in its trophic position (TP) from 2.0 to as high as 3.0. If this process in mesozooplankton, we call trophic lengthening, was transferred to higher trophic levels of a food web, a loss of energy could result in massive declines of fish biomass. We used compound‐specific nitrogen stable isotope data of amino acids (CSIA) to estimate and compare the nitrogen (N) sources and TPs of cod and flounder from FNC bloom influence areas (central Baltic Sea) and areas without it (western Baltic Sea). We tested if FNC‐triggered trophic lengthening in mesozooplankton is carried over to fish. The TP of cod from the western Baltic (4.1 ± 0.5), feeding mainly on decapods, was equal to reference values. Only cod from the central Baltic, mainly feeding on zooplanktivorous pelagics, had a significantly higher TP (4.6 ± 0.4), indicating a strong carry‐over effect trophic lengthening from mesozooplankton. In contrast, the TP of molluscivorous flounder, associated with the benthic food web, was unaffected by trophic lengthening and quite similar reference values of 3.2 ± 0.2 in both areas. This suggests that FNC blooms lead to a large loss of energy in zooplanktivorous but not in molluscivorous mesopredators. If FNC blooms continue to trigger the detour of energy at the base of the pelagic food web due to a massive heterotrophic microbial system, the TP of cod will not return to lower TP values and the fish stock not recover. Monitoring the TP of key species can identify fundamental changes in ecosystems and provide information for resource management. [ABSTRACT FROM AUTHOR]

Eurytemora affinis (Copepoda) were fed 15N-labeled Rhodomonas salina (Cryptophyta) or 15N-labeled Nodularia spumigena (Cyanobacteria) in excess under controlled laboratory conditions. Zooplankton collected from the Baltic Sea were fed natural phytoplankton amended with 15N-labeled N. spumigena. We quantified the direct incorporation of 15N tracer from N2-fixing N. spumigena (diazotroph nitrogen) and ammonium-utilizing R. salina into the amino acid nitrogen (AA-N) of zooplankton using complementary gas chromatography-combustion-isotope ratio mass spectrometry, gas chromatography-mass spectrometry, and elemental analysis-isotope ratio mass spectrometry approaches. Specific and mass-specific TN and AA-N incorporation rates of the 15N tracers were calculated for zooplankton. Highest incorporation of 15N was found in field zooplankton relying on N. spumigena and in E. affinis relying on R. salina. Lowest incorporation was found in E. affinis relying on N. spumigena. Decreasing specific and mass-specific rates during field experiments possibly were due to food shortage, whereas decreasing rates in E. affinis grazing on R. salina were more likely due to satiation. Specific and mass-specific rates were consistently low in E. affinis when exposed to N. spumigena, suggesting that these animals were reluctant to feed on N. spumigena. Essential isoleucine received most of the diazotroph nitrogen in field zooplankton, while nonessential amino acids received most 15N tracer in E. affinis. N. spumigena was clearly an important amino acid nitrogen source for Baltic Sea zooplankton. [ABSTRACT FROM AUTHOR]

Increasing sea surface temperatures (SST) and blooms of lipid‐poor, filamentous cyanobacteria can change mesozooplankton metabolism and foraging strategies in marine systems. Lipid shortage and imbalanced diet may challenge the build‐up of energy pools of lipids and proteins, and access to essential fatty acids (FAs) and amino acids (AAs) by copepods. The impact of cyanobacterial blooms on individual energy pools was assessed for key species temperate Temora longicornis and boreal Pseudo‐/Paracalanus spp. that dominated field mesozooplankton communities isolated by seasonal stratification in the central Baltic Sea during the hot and the cold summer. We looked at (a) total lipid and protein levels, (b) FA trophic markers and AA composition, and (c) compound‐specific stable carbon isotopes (δ13C) in bulk mesozooplankton and in a subset of parameters in particulate organic matter. Despite lipid‐poor cyanobacterial blooms, the key species were largely able to cover both energy pools, yet a tendency of lipid reduction was observed in surface animals. Omni‐ and carnivory feeding modes, FA trophic makers, and δ13C patterns in essential compounds emphasized that cyanobacterial FAs and AAs have been incorporated into mesozooplankton mainly via feeding on mixo‐ and heterotrophic (dino‐) flagellates and detrital complexes during summer. Foraging for essential highly unsaturated FAs from (dino‐) flagellates may have caused night migration of Pseudo‐/Paracalanus spp. from the deep subhalocline waters into the upper waters. Only in the hot summer (SST>19.0°C) was T. longicornis submerged in the colder subthermocline water (~4°C). Thus, the continuous warming trend and simultaneous feeding can eventually lead to competition on the preferred diet by key copepod species below the thermocline in stratified systems. A comparison of δ13C patterns of essential AAs in surface mesozooplankton across sub‐basins of low and high cyanobacterial biomasses revealed the potential of δ13C‐AA isoscapes for studies of commercial fish feeding trails across the Baltic Sea food webs. [ABSTRACT FROM AUTHOR]

Abstract: We report on methane enrichments that were observed during summer in the upper water column of the Gotland Basin, central Baltic Sea. In the eastern part of the basin, methane concentrations just below the thermocline varied between 15 nM and 77 nM, in contrast to the western part where no methane enrichments could be detected. Stable carbon isotope ratios of methane (δ13C‐CH4 of −67.6‰) indicated its in situ biogenic origin from CO2 reduction, which was supported by clonal sequences that clustered with Methanomicrobiaceae, a family of methanogenic Archaea. Incubation experiments with a Temora longicornis dominated seston fraction obtained from the relevant depth showed a positive correlation between seston concentration and methane production rates. Our results, in combination with previous literature outcomes, suggest that the methane enrichment in the eastern basin might be sustained by a diet‐consumer relationship between the dinoflagellate Dinophysis norvegica and the copepod T. longicornis. However, our mass balance indicates that a local methane production of 110 pmol L−1 d−1 was needed to maintain the methane enrichment, and that the estimated production rate from our incubation experiments of 0.3 pmol CH4 d−1 per adult T. longicornis (about 1 pmol L−1 d−1) was too low to maintain the methane enrichment by zooplankton associated methane production only. These calculations also showed that methane was consumed below the thermocline and not transported into the upper‐ocean, suggesting that other sources in the mixed layer in the range of 95 pmol L−1 d−1 are needed to maintain the observed methane air–sea flux. [ABSTRACT FROM AUTHOR]