Long-Term Eutrophication and Dynamics of Bloom-Forming Microbial Communities during Summer HAB in Large Arctic Lake

Harmful algal blooms (HABs) in arctic lakes are recent phenomena. In our study, we performed a long-term analysis (1990–2017) of the eutrophication of Lake Imandra, a large subarctic lake, and explored the biodiversity of bloom-forming microorganisms of a 2017 summer HAB. We performed a 16Sr rRNA metabarcoding study of microbial communities, analysed the associations between N, P, C, and chlorophyll concentrations in the lake water, and developed models for the prediction of HABs based on total P concentration. We have demonstrated that blooms in Lake Imandra occur outside of optimal Redfield ratios and have a nonlinear association with P concentrations. We found that recent summer HABs in a lake occur as simultaneous blooms of a diatom Aulacoseira sp. and cyanobacteria Dolichospermum sp. We have studied the temporal dynamics of microbial communities during the bloom and performed an analysis of the publicly available Dolichospermum genomes to outline potential genetic mechanisms beneath simultaneous blooming. We found genetic traits requisite for diatom-diazotroph associations, which may lay beneath the simultaneous blooming of Aulacoseira sp. and Dolichospermum sp. in Lake Imandra. Both groups of organisms have the ability to store nutrients and form a dormant stage. All of these factors will ensure the further development of the HABs in Lake Imandra and the dispersal of these bloom-forming species to neighboring lakes.

Geologic controls on phytoplankton elemental composition

Although the nutrient content of planktonic organic matter (C:N:Porg) plays a crucial role in marine metazoan evolution and global biogeochemistry (1–3), its geologic history is poorly constrained, and it is often regarded as a constant “Redfield” ratio of C:N:Porg~106:16:1. We calculate C:N:Porg through the Phanerozoic by including nutrient- and temperature-dependent C:N:Porg parameterizations (4–6) in a model of long-term biogeochemical cycles (7). We infer a decrease from high Paleozoic C:Porg and N:Porg to present-day Redfield ratios. This gradual nutrient enrichment of marine organic matter stems from a decrease in the global average temperature and an increase in seawater phosphate availability, which are driven by various Phanerozoic events, mainly the middle to late Paleozoic emergence and expansion of land plants and the Triassic breakup of the supercontinent Pangaea. The nutrient enrichment of planktonic organic matter likely impacted the evolution of marine fauna and global biogeochemistry.

Characterization of ocean biogeochemical processes: a generalized total least-squares estimator of the Redfield ratios

The chemical composition of the global ocean is governed by biological, chemical and physical processes. These processes interact with each other so that the concentrations of carbon dioxide, oxygen, nitrate and phosphate vary in constant proportions, referred to as the Redfield ratios. We build here the Generalized Total Least-Squares estimator of these ratios. The interest of our approach is twofold: it respects the hydrological characteristics of the studied areas, and it can be applied identically in any area where enough data are available. The tests performed on the Atlantic Ocean highlight a variability of the Redfield ratios, both with geographical location and with depth. This variability emphasizes the importance of local and accurate estimates of Redfield ratios.

Stark Contrast in Denitrification and Anammox across the Deep Norwegian Trench in the Skagerrak

ABSTRACTEnvironmental anaerobic ammonium oxidation (anammox) was demonstrated for the first time in 2002, using15N labeling, in homogenized sediment from the Skagerrak, where it accounted for up to 67% of N2production. We returned to some of these original sites in 2010 to make measurements of nitrogen and carbon cycling under conditions more representative of thosein situ, quantifying anammox and denitrification, together with oxygen penetration and consumption, in intact sediment cores. Overall, oxygen consumption and N2production decayed with water depth, as expected, but the drop in N2production was relatively more pronounced. Whereas we confirmed the dominance of N2production by anammox (72% and 77%) at the two deepest sites (∼700 m of water), anammox was conspicuously absent from two shallower sites (∼200 m and 400 m). At the shallower sites, we could measure no anammox activity with either intact or homogeneous sediment, and quantitative PCR (16S rRNA) gave a negligible abundance of anammox bacteria in the anoxic layers. Such an absence of anammox, especially at one locale where it was originally demonstrated, is hard to reconcile. Despite the dominance of anammox at the deepest sites, anammox activity could not make up for the drop in denitrification, and assuming Redfield ratios for the organic matter being mineralized, the estimated retention of fixed N actually increased to 90% to 97% of that mineralized, whereas it was 80% to 86% at the shallower sites.

The stoichiometric ratio during biological removal of inorganic carbon and nutrient in the Mississippi River plume and adjacent continental shelf

The stoichiometric ratios of dissolved inorganic carbon (DIC) and nutrients during biological removal have been widely assumed to follow the Redfield ratios (especially the C/N ratio) in large river plume ecosystems. However, this assumption has not been systematically examined and documented because DIC and nutrients are rarely studied simultaneously in a river plume area, a region in which they can be affected by strong river-ocean mixing as well as intense biological activity. We examined stoichiometric ratios of DIC, total alkalinity (TA), and nutrients (NO3, PO43− and Si(OH)4) data during biological removal in the Mississippi River plume and adjacent continental shelf in June 2003 and August 2004 with biological removals defined as the difference between measured values and values predicted on the basis of conservative mixing determined using a multi-endmember mixing model. Despite complex physical and biogeochemical influences, relationships between DIC and nutrients were strongly dependent on salinity range and geographic location, and influenced by biological removal. Lower C/Si and N/Si ratios in one nearshore area were attributed to a potential silicate source induced by water exchange with coastal salt marshes. When net biological uptake was separated from river-ocean mixing and the impact of marshes and bays excluded, stoichiometric ratios of C/N/Si were similar to the Redfield ratios, thus supporting the applicability of the Redfield-type C/N/Si ratios as a principle in river-plume biogeochemical models.

Seasonal variation in marine C:N:P stoichiometry: can the composition of seston explain stable Redfield ratios?

Seston is suspended particulate organic matter, comprising a mixture of autotrophic, heterotrophic and detrital material. Despite variable proportions of these components, marine seston often exhibits relatively small deviations from the Redfield ratio (C:N:P = 106:16:1). Two time-series from the Norwegian shelf in Skagerrak are used to identify drivers of the seasonal variation in seston elemental ratios. An ordination identified water mass characteristics and bloom dynamics as the most important drivers for determining C:N, while changes in nutrient concentrations and biomass were most important for the C:P and N:P relationships. There is no standardized method for determining the functional composition of seston and the fractions of POC, PON and PP associated with phytoplankton, therefore any such information has to be obtained by indirect means. In this study, a generalized linear model was used to differentiate between the live autotrophic and non-autotrophic sestonic fractions, and for both stations the non-autotrophic fractions dominated with respective annual means of 76 and 55%. This regression model approach builds on assumptions (e.g. constant POC:Chl-a ratio) and the robustness of the estimates were explored with a bootstrap analysis. In addition the autotrophic percentage calculated from the statistical model was compared with estimated phytoplankton carbon, and the two independent estimates of autotrophic percentage were comparable with similar seasonal cycles. The estimated C:nutrient ratios of live autotrophs were, in general, lower than Redfield, while the non-autotrophic C:nutrient ratios were higher than the live autotrophic ratios and above, or close to, the Redfield ratio. This is due to preferential remineralization of nutrients, and the P content mainly governed the difference between the sestonic fractions. Despite the seasonal variability in seston composition and the generally low contribution of autotrophic biomass, the variation observed in the total seston ratios was low compared to the variation found in dissolved and particulate pools. Sestonic C:N:P ratios close to the Redfield ratios should not be used as an indicator of phytoplankton physiological state, but could instead reflect varying contributions of sestonic fractions that sum up to an elemental ratio close to Redfield.

Seasonal variation in marine C:N:P stoichiometry: can the composition of seston explain stable Redfield ratios?

Seston is suspended particulate organic matter, comprising a mixture of autotrophic, heterotrophic and detrital material. Despite variable proportions of these components, marine seston often exhibit relatively small deviations from the Redfield ratio (C:N:P = 106:16:1). Two time-series from the Norwegian shelf in Skagerrak are used to identify drivers of the seasonal variation in seston elemental ratios. An ordination identified water mass characteristics and bloom dynamics as the most important drivers for determining C:N, while changes in nutrient concentrations and biomass were most important for the C:P and N:P relationships. A generalized linear model was used to differentiate between the live autotrophic and non-autotrophic sestonic fractions, and for both stations the non-autotrophic fractions dominated with respective annual means of 24 and 45 % live autotrophs. The autotrophic percentage calculated from the statistical model was compared with estimated phytoplankton carbon, and the two independent estimates of autotrophic percentage were comparable with similar seasonal cycles. The estimated C:nutrient ratios of live autotrophs were in general lower than Redfield, while the non-autotrophic C:nutrient ratios were higher than the live autotrophic ratios and above or close to the Redfield ratio. This is due to preferential remineralization of nutrients (especially phosphorus), while carbon gradually builds up in the detritus pool. Despite the seasonal variability in seston composition and the generally low contribution of autotrophic biomass, the variation observed in the total seston ratios was low compared to the variation found in dissolved and particulate pools. This study shows that sestonic Redfield ratios cannot automatically be interpreted as phytoplankton with "balanced growth", but could instead reflect varying contributions of sestonic compartments that sum up to an elemental ratio close to Redfield.