Aim To test if a phytoplankton bloom is panmictic, or whether geographical and environmental factors cause spatial and temporal genetic structure.Location Baltic Sea.Method During four cruises, we isolated clonal strains of the diatom Skeletonema marinoi from 9 to 10 stations along a 1132 km transect and analysed the genetic structure using eight microsatellites. Using F-statistics and Bayesian clustering analysis we determined if samples were significantly differentiated. A seascape approach was applied to examine correlations between gene flow and oceanographic connectivity, and combined partial Mantel test and RDA based variation partitioning to investigate associations with environmental gradients. ResultsThe bloom was initiated during the second half of March in the southern and the northern-parts of the transect, and later propagated offshore. By mid-April the bloom declined in the south, whereas high phytoplankton biomass was recorded northward. We found two significantly differentiated populations along the transect. Genotypes were significantly isolated by distance and by the southnorth salinity gradient, which illustrated that the effects of distance and environment were confounded. The gene flow among the sampled stations was significantly correlated with oceanographic connectivity. The depletion of silica during the progression of the bloom was related to a temporal population genetic shift.Main conclusions A phytoplankton bloom may propagate as a continuous cascade and yet be genetically structured over both spatial and temporal scales. The Baltic Sea spring bloom displayed strong spatial structure driven by oceanographic connectivity and geographical distance, which was enhanced by the pronounced salinity gradient. Temporal transition of conditions important for growth may induce genetic shifts and different phenotypic strategies, which serve to maintain the bloom over longer periods.

In temperate systems, phytoplankton spring blooms deplete inorganic nutrients and are major sources of organic matter for the microbial loop. In response to phytoplankton exudates and environmental factors, heterotrophic microbial communities are highly dynamic and change their abundance and composition both on spatial and temporal scales. Yet, most of our understanding about these processes comes from laboratory model organism studies, mesocosm experiments or single temporal transects. Spatial-temporal studies examining interactions of phytoplankton blooms and bacterioplankton community composition and function, though being highly informative, are scarce. In this study, pelagic microbial community dynamics (bacteria and phytoplankton) and environmental variables were monitored during a spring bloom across the Baltic Proper (two cruises between North Germany to Gulf of Finland). To test to what extent bacterioplankton community composition relates to the spring bloom, we used next generation amplicon sequencing of the 16S rRNA gene, phytoplankton diversity analysis based on microscopy counts and population genotyping of the dominating diatom Skeletonema marinoi. Several phytoplankton bloom related and environmental variables were identified to influence bacterial community composition. Members of Bacteroidetes and Alphaproteobacteria dominated the bacterial community composition but the bacterial groups showed no apparent correlation with direct bloom related variables. The less abundant bacterial phyla Actinobacteria, Planctomycetes, and Verrucomicrobia, on the other hand, were strongly associated with phytoplankton biomass, diatom:dinoflagellate ratio, and colored dissolved organic matter (cDOM). Many bacterial operational taxonomic units (OTUs) showed high niche specificities. For example, particular Bacteroidetes OTUs were associated with two distinct genetic clusters of S. marinoi. Our study revealed the complexity of interactions of bacterial taxa with inter- and intraspecific genetic variation in phytoplankton. Overall, our findings imply that biotic and abiotic factors during spring bloom influence bacterial community dynamics in a hierarchical manner.

In the Baltic Sea, floating blooms of nitrogen-fixing cyanobacteria occur yearly during late summer. These blooms can sometimes be limited by iron. Due to extensive foresting around the Baltic Sea, iron is entering the Baltic Sea partly bound to dissolved organic material (DOM) via rivers. An experiment was performed in 300 l laboratory mesocosms to test the hypothesis that riverine highmolecular weight dissolved organic matter (HMWDOM), extracted by tangential flow filtration >1000 Da, stimulates the biomass of nitrogen-fixing cyanobacteria, by increasing the availability of iron. The addition of iron/EDTA and of DOM resulted in 5 to 10 times higher biomass of nitrogenfixing cyanobacteria. Accordingly, higher primary production and particulate nitrogen concentration at the end of the experiment were observed in those treatments compared to the control. The removal of mesozooplankton grazers did not have a significant effect on the microphytoplankton biomass and species composition. Nodularia spumigena biomass was highest in the treatments receiving DOM, but addition of iron alone had no significant effect on this. N. spumigena was less positively affected by iron addition than Anabaena cf. inaequalis, suggesting that N. spumigena is a better competitor for iron. Separate microcosms comparing additions of iron, manganese and cobalt showed that iron was limiting for cyanobacterial biomass development. The results strongly suggest that iron bound to DOM can contribute to the iron demands of nitrogen-fixing cyanobacteria in the Baltic Sea.

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