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.

Dataset from a mesocosm experiment on brownification in the Baltic Sea.

Climate change is projected to cause brownification of some coastal seas due to increased runoff of terrestrially derived organic matter. We carried out a mesocosm experiment over 15 days to test the effect of this on the planktonic ecosystem. The experiment was set up in 2.2 m3 plastic bags moored outside the Tvärminne Zoological Station at the SW coast of Finland. We used four treatments, each with three replicates: control (Contr) without any manipulation; addition of a commercially available organic carbon additive called HuminFeed (Hum; 2 mg L-1); addition of inorganic nutrients (Nutr; 5.7 µM NH4 and 0.65µM PO4); and a final treatment of combined Nutr and Hum (Nutr+Hum) additions. Water samples were taken daily, and measured variables included water transparency, organic and inorganic nutrient pools, chlorophyll a (Chla), primary and bacterial production and particle counts by flow cytometry.

Brownification affects phytoplankton community composition but not primary productivity in eutrophic coastal waters: A mesocosm experiment in the Baltic Sea.

Climate change is projected to cause brownification of some coastal seas due to increased runoff of terrestrially derived organic matter. We carried out a mesocosm experiment (15 d) to test the effect of this on the planktonic ecosystem expecting reduced primary production and shifts in the phytoplankton community composition. The experiment was set up in 2.2 m3 mesocosm bags using four treatments, each with three replicates: control (Contr) without any manipulation, organic carbon additive HuminFeed (Hum; 2 mg L-1), inorganic nutrients (Nutr; 5.7 µM NH4 and 0.65 µM PO4), and combined Nutr and Hum (Nutr + Hum) additions. Measured variables included organic and inorganic nutrient pools, chlorophyll a (Chla), primary and bacterial production and particle counts by flow cytometry. The bags with added inorganic nutrients developed a phytoplankton bloom that depleted inorganic N at day 6, followed by a rapid decline in Chla. Brownification did not reduce primary production at the tested concentration. Bacterial production was lowest in the Contr, but similar in the three treatments receiving additions likely due to increased carbon available for heterotrophic bacteria. Picoeukaryotes clearly benefited by brownification after inorganic N depletion, which could be due to more effective nutrient recycling, nutrient affinity, light absorption, or alternatively lower grazing pressure. In conclusion, brownification shifted the phytoplankton community composition towards smaller species with potential effects on carbon fluxes, such as sinking rates and export to the sea floor.

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.

Nile Red staining of phytoplankton neutral lipids: species-specific fluorescence kinetics in various solvents.

Nile Red (NR) staining potentially offers a simple method for monitoring lipid accumulation in microalgal cultivation. However, variable staining efficiencies and methods have been reported. The effect of dimethyl sulfoxide (DMSO), ethylene glycol (EG) and glycerol on NR penetration with four different phytoplankton species representing different taxonomical groups was studied. Treatment with the solvents enhanced the NR fluorescence of the diatom Phaeodactylum tricornutum during kinetic fluorescence measurements, but high concentrations of solvents were needed. None of the solvents improved NR staining of the green alga Chlorella pyrenoidosa and Scenedesmus obliquus, which are known to be difficult to stain due to their thick and rigid cell walls. The naked Isochrysis sp. cells stained best without solvents. The results confirm that NR staining protocol needs to be optimized for each species.

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.

Functional group richness: implications of biodiversity for light use and lipid yield in microalgae.

Currently, very few studies address the relationship between diversity and biomass/lipid production in primary producer communities for biofuel production. Basic studies on the growth of microalgal communities, however, provide evidence of a positive relationship between diversity and biomass production. Recent studies have also shown that positive diversity-productivity relationships are related to an increase in the efficiency of light use by diverse microalgal communities. Here, we hypothesize that there is a relationship between diversity, light use, and microalgal lipid production in phytoplankton communities. Microalgae from all major freshwater algal groups were cultivated in treatments that differed in species richness and functional group richness. Polycultures with high functional group richness showed more efficient light use and higher algal lipid content with increasing species richness. There was a clear correlation between light use and lipid production in functionally diverse communities. Hence, a powerful and cost-effective way to improve biofuel production might be accomplished by incorporating diversity related, resource-use-dynamics into algal biomass production.

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.

Diversity predicts stability and resource use efficiency in natural phytoplankton communities.

The relationship between species diversity and ecosystem functioning has been debated for decades, especially in relation to the "macroscopic" realm (higher plants and metazoans). Although there is emerging consensus that diversity enhances productivity and stability in communities of higher organisms; however, we still do not know whether these relationships apply also for communities of unicellular organisms, such as phytoplankton, which contribute approximately 50% to the global primary production. We show here that phytoplankton resource use, and thus carbon fixation, is directly linked to the diversity of phytoplankton communities. Datasets from freshwater and brackish habitats show that diversity is the best predictor for resource use efficiency of phytoplankton communities across considerable environmental gradients. Furthermore, we show that the diversity requirement for stable ecosystem functioning scales with the nutrient level (total phosphorus), as evidenced by the opposing effects of diversity (negative) and resource level (positive) on the variability of both resource use and community composition. Our analyses of large-scale observational data are consistent with experimental and model studies demonstrating causal effects of microbial diversity on functional properties at the system level. Our findings point at potential linkages between eutrophication and pollution-mediated loss of phytoplankton diversity. Factors reducing phytoplankton diversity may have direct detrimental effects on the amount and predictability of aquatic primary production.

Internal ecosystem feedbacks enhance nitrogen-fixing cyanobacteria blooms and complicate management in the Baltic Sea.

Eutrophication of the Baltic Sea has potentially increased the frequency and magnitude of cyanobacteria blooms. Eutrophication leads to increased sedimentation of organic material, increasing the extent of anoxic bottoms and subsequently increasing the internal phosphorus loading. In addition, the hypoxic water volume displays a negative relationship with the total dissolved inorganic nitrogen pool, suggesting greater overall nitrogen removal with increased hypoxia. Enhanced internal loading of phosphorus and the removal of dissolved inorganic nitrogen leads to lower nitrogen to phosphorus ratios, which are one of the main factors promoting nitrogenfixing cyanobacteria blooms. Because cyanobacteria blooms in the open waters of the Baltic Sea seem to be strongly regulated by internal processes, the effects of external nutrient reductions are scale-dependent. During longer time scales, reductions in external phosphorus load may reduce cyanobacteria blooms; however, on shorter time scales the internal phosphorus loading can counteract external phosphorus reductions. The coupled processes inducing internal loading, nitrogen removal, and the prevalence of nitrogen-fixing cyanobacteria can qualitatively be described as a potentially self-sustaining "vicious circle." To effectively reduce cyanobacteria blooms and overall signs of eutrophication, reductions in both nitrogen and phosphorus external loads appear essential.

Allelopathic effects of toxic haptophyte Prymnesium parvum lead to release of dissolved organic carbon and increase in bacterial biomass.

The haptophyte Prymnesium parvum has lytic properties, and it affects coexisting phytoplankton species through allelopathy. We studied the effect of P. parvum allelochemicals on the lysis of the nontoxic and nonaxenic cryptomonad Rhodomonas salina and the consequent release of dissolved organic carbon (DOC). Changes in production, cell density, and biomass of associated bacteria were measured over 12 h. Six different combinations of P. parvum and R. salina cultures, their cell- and bacteria-free filtrates, and growth media as controls were used in the experiments. When P. parvum and R. salina cells were mixed, a significant increase in DOC concentration was measured within 30 min. Bacterial biomass increased significantly during the next 6 to 12 h when R. salina was mixed either with the P. parvum culture or the cell-free P. parvum filtrates (allelochemicals only). In contrast, bacterial biomass did not change in the treatments without the allelopathic action (without R. salina cells). Blooms of P. parvum alter the functioning of the planktonic food web by increasing carbon transfer through the microbial loop. In addition, P. parvum may indirectly benefit from the release of DOC as a result of its ability to ingest bacteria, by which it can acquire nutrients during limiting conditions.