Phytoplankton Species Richness along Coastal and Estuarine Salinity Continua.

The high number of freshwater species at low salinity and the correspondingly high number of marine species at high salinity enveloping a conspicuous richness minimum at intermediate salinities has shaped our basic understanding of biodiversity along a coastal salinity gradient for almost 80 years. Visualized as the Remane curve, this iconic concept was originally based on sedentary macroinvertebrates in the Baltic Sea. To what extent the concept can be generalized, particularly to free-drifting organisms, is currently debated. Here we use approximately 16,000 phytoplankton samples from two large coastal ecosystems-the Baltic Sea and Chesapeake Bay-to analyze the relationship between salinity and phytoplankton species richness. Alpha diversity showed a consistent variation along the salinity gradient, with a minimum at mesohaline salinities of around 7-9. Rarefied species pools at narrow salinity intervals also showed reduced diversity at intermediate salinities, surrounded by high richness toward both ends of the gradient. The cumulative likelihood of species presence validated the minimum at intermediate salinities. Community composition changed abruptly at the α diversity minimum in the Baltic Sea, while it changed gradually along the salinity gradient in Chesapeake Bay. We conclude that the Remane concept is in every respect valid for phytoplankton.

Alexandrium fundyense cyst viability and germling survival in light vs. dark at a constant low temperature.

Both observations and models suggest that large-scale coastal blooms of Alexandrium fundyense in the Gulf of Maine are seeded by deep-bottom cyst accumulation zones ("seed beds") where cysts germinate from the sediment surface or the overlying near-bottom nepheloid layers at water depths exceeding 100 m. The germling cells and their vegetative progeny are assumed to be subject to mortality while in complete darkness, as they swim to illuminated surface waters. To test the validity of this assumption we conducted laboratory investigations of cyst viability and the survival of the germling cells and their vegetative progeny during prolonged exposure to darkness at a temperature of 6 °C, simulating the conditions in deep Gulf of Maine waters. We isolated cysts from bottom sediments collected in the Gulf of Maine under low red light and incubated them in 96-well tissue culture-plates in culture medium under a 10:14 h light:dark cycle and under complete darkness. Cyst viability was high, with excystment frequency reaching 90% in the illuminated treatment after 30 days and in the dark treatment after 50 days. Average germination rates were 0.062 and 0.038 d-1 for light and dark treatments, respectively. The dark treatment showed an approximately 2-week time lag in maximum germination rates compared to the light treatment. Survival of germlings was considerably lower in the dark treatment. In the light treatments, 47% of germinated cysts produced germlings that were able to survive for 7 days and produce vegetative progeny, i.e., there were live cells in the well along with an empty cyst at least once during the experiment. In the dark treatments 12% of the cysts produced germlings that were able to survive for the same length of time. When dark treatments are scaled to take into account non-darkness related mortality, approximately 28% of the cysts produced germlings that were able to survive for at least 7 days. Even though cysts are able to germinate in darkness, the lack of illumination considerably reduces survival rate of germling cells. In addition to viability of cysts in surface sediments and the near-bottom nepheloid layer, survivability of germling cells and their vegetative progeny at aphotic depths is an important consideration in assessing the quantitative role of deep-coastal cyst seed beds in bloom formation.

Half-century trends in alpha and beta diversity of phytoplankton summer communities in the Helsinki Archipelago, the Baltic Sea.

We analyzed phytoplankton biodiversity trends in a 52 year (1967-2018) monitoring time-series from the archipelago of Helsinki, Gulf of Finland, the Baltic Sea. The community ordination revealed strong ordering of samples along the time axis (generalized additive model-gam fit: R 2 = 0.9). Species richness increased in time and was the most influential alpha diversity descriptor related to the community structure (gam fit: R 2 = 0.56-0.70). Changes in species richness accounted for 35-36% of the mean between-sample beta diversity. The remaining 64-65% was due to species turnover-the dominant component of the biodiversity trend. The temporal beta diversity trend reflected the eutrophication history of the geographically confined region, with a turning point in mid-1990s demarking the adaptation and recovery phases of the phytoplankton community. Trends in spatial beta diversity revealed homogenization of the communities in the outer archipelago zone, but not in the inner bays. The temporal decay of community similarity revealed high turnover rate, with 23.6 years halving time in the outer archipelago and 11.3 years in the inner bays, revealing the differences in eutrophication strength. The observed phytoplankton trends manifest the regional eutrophication history, and dispersal of new species to the unsaturated brackish species pool.

Environmental safety data on CuO and TiO2 nanoparticles for multiple algal species in natural water: Filling the data gaps for risk assessment.

Most research on nanoparticle (NP) ecotoxicological effects has been conducted on single species in laboratory conditions that are not environmentally representative. We compared the effects of CuO NPs, CuSO4 (ionic control) and TiO2 NPs in nutrient-adjusted natural water (ANW) and in the OECD201 standard medium to four different algal species: green algae Raphidocelis subcapitata and Chlamydomonas reinhardtii, a diatom Fistulifera pelliculosa, and a cyanobacterium Synechocystis sp. Biomass and the effective quantum yield of photosystem II (Fv/Fm) were used as toxicity endpoints. CuO NPs were very toxic across taxa in the OECD201 assay (biomass-based 72 h EC50 0.2-0.9 mg l-1). Toxicity of CuO NPs was explained by shedding of ions from particles as Cu2+ is highly toxic: 72 h EC50 in the OECD201 medium was 0.01-0.03 mg l-1 in three species and 0.003 mg l-1 in the case of the cyanobacterium. Toxicity of copper compounds was overall reduced in ANW, presumably because of reduced bioavailability due to metal ions binding to natural organic matter. Copper compounds were more toxic to the cyanobacterium than to other algae and this effect was not amended in ANW. TiO2 NPs did not inhibit the biomass production and photosynthesis of the diatom or the cyanobacterium up to 100 mg l-1, but inhibited biomass production of green algae in the OECD201 medium (EC50 14-15 mg l-1). TiO2 NPs also did not significantly inhibit Fv/Fm up to 100 mg l-1, suggesting a general lack of effect on photosynthesis. Adverse effects of TiO2 NPs were at least in part due to cell-NP heteroagglomeration. Our data are informative for the complete risk assessment of engineered NPs by filling data gaps about NP effects in environmentally realistic conditions.

Potency of (doped) rare earth oxide particles and their constituent metals to inhibit algal growth and induce direct toxic effects.

Use of rare earth elements (REEs) has increased rapidly in recent decades due to technological advances. It has been accompanied by recurring rare earth element anomalies in water bodies. In this work we (i) studied the effects of eight novel doped and one non-doped rare earth oxide (REO) particles (aimed to be used in solid oxide fuel cells and gas separation membranes) on algae, (ii) quantified the individual adverse effects of the elements that constitute the (doped) REO particles and (iii) attempted to find a discernible pattern to relate REO particle physicochemical characteristics to algal growth inhibitory properties. Green algae Raphidocelis subcapitata (formerly Pseudokirchneriella subcapitata) were used as a test species in two different formats: a standard OECD201 algal growth inhibition assay and the algal viability assay (a 'spot test') that avoids nutrient removal effects. In the 24h 'spot' test that demonstrated direct toxicity, algae were not viable at REE concentrations above 1mgmetal/L. 72-hour algal growth inhibition EC50 values for four REE salts (Ce, Gd, La, Pr) were between 1.2 and 1.4mg/L, whereas the EC50 for REO particles ranged from 1 to 98mg/L. The growth inhibition of REEs was presumably the result of nutrient sequestration from the algal growth medium. The adverse effects of REO particles were at least in part due to the entrapment of algae within particle agglomerates. Adverse effects due to the dissolution of constituent elements from (doped) REO particles and the size or specific surface area of particles were excluded, except for La2NiO4. However, the structure of the particles and/or the varying effects of oxide composition might have played a role in the observed effects. As the production rates of these REO particles are negligible compared to other forms of REEs, there is presumably no acute risk for aquatic unicellular algae.

Rehabilitating the cyanobacteria - niche partitioning, resource use efficiency and phytoplankton community structure during diazotrophic cyanobacterial blooms.

Blooms of nitrogen-fixing cyanobacteria are recurrent phenomena in marine and freshwater habitats, and their supplying role in aquatic biogeochemical cycles is generally considered vital. The objective of this study was to analyse whether an increasing proportion of nitrogen-fixing cyanobacteria affects (i) the composition of the non-diazotrophic component of ambient phytoplankton communities and (ii) resource use efficiency (RUE; ratio of Chl a to total nutrients) - an important ecosystem function. We hypothesize that diazotrophs increase community P use and decrease N use efficiencies, as new N is brought into the system, relaxing N, and concomitantly aggravating P limitation. We test this by analysing an extensive data set from the Baltic Sea (> 3700 quantitative phytoplankton samples), known to harbour conspicuous and recurrent blooms of Nodularia spumigena and Aphanizomenon sp.System-level phosphorus use efficiency (RUEP) was positively related to high proportion of diazotrophic cyanobacteria, suggesting aggravation of phosphorus limitation. However, concomitant decrease of nitrogen use efficiency (RUEN) was not observed. Nodularia spumigena, a dominant diazotroph and a notorious toxin producer, had a significantly stronger relationship with RUEP, compared to the competing non-toxic Aphanizomenon sp., confirming niche differentiation in P acquisition strategies between the major bloom-forming cyanobacterial species in the Baltic Sea. Nodularia occurrences were associated with stronger temperature stratification in more offshore environments, indicating higher reliance on in situ P regeneration.By using constrained and unconstrained ordination, permutational multivariate analysis of variance and local similarity analysis, we show that diazotrophic cyanobacteria explained no more than a few percentage of the ambient phytoplankton community variation. The analyses furthermore yielded rather evenly distributed negative and positive effects on individual co-occurring phytoplankton taxa, with no obvious phylogenetic or functional trait-based patterns. Synthesis. Our study reveals that despite the widely acknowledged noxious impacts of cyanobacterial blooms, the overall effect on phytoplankton community structure is minor. There are no predominantly positive or negative associations with ambient phytoplankton species. Species-specific niche differences in cyanobacterial resource acquisition affect important ecosystem functions, such as biomass production per unit limiting resource.

New production regulates export stoichiometry in the ocean.

The proportion in which carbon and growth-limiting nutrients are exported from the oceans' productive surface layer to the deep sea is a crucial parameter in models of the biological carbon pump. Based on >400 vertical flux observations of particulate organic carbon (POC) and nitrogen (PON) from the European Arctic Ocean we show the common assumption of constant C:N stoichiometry not to be met. Exported POC:PON ratios exceeded the classical Redfield atomic ratio of 6.625 in the entire region, with the largest deviation in the deep Central Arctic Ocean. In this part the mean exported POC:PON ratio of 9.7 (a:a) implies c. 40% higher carbon export compared to Redfield-based estimates. When spatially integrated, the potential POC export in the European Arctic was 10-30% higher than suggested by calculations based on constant POC:PON ratios. We further demonstrate that the exported POC:PON ratio varies regionally in relation to nitrate-based new production over geographical scales that range from the Arctic to the subtropics, being highest in the least productive oligotrophic Central Arctic Ocean and subtropical gyres. Accounting for variations in export stoichiometry among systems of different productivity will improve the ability of models to resolve regional patterns in carbon export and, hence, the oceans' contribution to the global carbon cycle will be predicted more accurately.

Decadal-scale changes of dinoflagellates and diatoms in the anomalous baltic sea spring bloom.

The algal spring bloom in the Baltic Sea represents an anomaly from the winter-spring bloom patterns worldwide in terms of frequent and recurring dominance of dinoflagellates over diatoms. Analysis of approximately 3500 spring bloom samples from the Baltic Sea monitoring programs revealed (i) that within the major basins the proportion of dinoflagellates varied from 0.1 (Kattegat) to >0.8 (central Baltic Proper), and (ii) substantial shifts (e.g. from 0.2 to 0.6 in the Gulf of Finland) in the dinoflagellate proportion over four decades. During a recent decade (1995-2004) the proportion of dinoflagellates increased relative to diatoms mostly in the northernmost basins (Gulf of Bothnia, from 0.1 to 0.4) and in the Gulf of Finland, (0.4 to 0.6) which are typically ice-covered areas. We hypothesize that in coastal areas a specific sequence of seasonal events, involving wintertime mixing and resuspension of benthic cysts, followed by proliferation in stratified thin layers under melting ice, favors successful seeding and accumulation of dense dinoflagellate populations over diatoms. This head-start of dinoflagellates by the onset of the spring bloom is decisive for successful competition with the faster growing diatoms. Massive cyst formation and spreading of cyst beds fuel the expanding and ever larger dinoflagellate blooms in the relatively shallow coastal waters. Shifts in the dominant spring bloom algal groups can have significant effects on major elemental fluxes and functioning of the Baltic Sea ecosystem, but also in the vast shelves and estuaries at high latitudes, where ice-associated cold-water dinoflagellates successfully compete with diatoms.

On the relative significance of bacteria for the distribution of polychlorinated biphenyls in Arctic Ocean surface waters.

This study presents the first field observations of polychlorinated biphenyls (PCB) in bacteria in oceanic waters. To contribute to the limited knowledge of what role bacteria play in the dynamics of hydrophobic organic contaminants (HOCs) in surface seawater, PCB concentrations were measured in bacteria (0.2-2 microm) collected at seven stations in the northern Barents Sea marginal ice zone (MIZ) and the central Arctic Ocean. Concentrations of individual PCB congeners in bacteria were 0.5-5 ng/g oc (organic carbon), which was as high as or higher than PCB concentrations in bulk particulate organic carbon (POC, "phytoplankton"; > 0.7 microm). Considering the relative biomasses of phytoplankton and bacteria, the amount of PCB in bacteria was generally 5-20% of that in phytoplankton, but at two stations the bacterial biomass contained more PCBs than the phytoplankton pool. This study further showed that efficient PCB uptake in bacteria may be described by an apparent equilibrium partitioning model with linear regressions between the organic-carbon-normalized partition coefficient and the octanol-water partition coefficient (log K(bact-oc)-log K(ow)).