Marine bacterioplankton are essential in global nutrient cycling and organic matter turnover. Time-series analyses, often at monthly sampling frequencies, have established the paramount role of abiotic and biotic variables in structuring bacterioplankton communities and productivities. However, fine-scale seasonal microbial activities, and underlying biological principles, are not fully understood. We report results from four consecutive years of high-frequency time-series sampling in the Baltic Proper. Pronounced temporal dynamics in most investigated microbial variables were observed, including bacterial heterotrophic production, plankton biomass, extracellular enzyme activities, substrate uptake rate constants of glucose, pyruvate, acetate, amino acids, and leucine, as well as nutrient limitation bioassays. Spring blooms consisting of diatoms and dinoflagellates were followed by elevated bacterial heterotrophic production and abundances. During summer, bacterial productivity estimates increased even further, coinciding with an initial cyanobacterial bloom in early July. However, bacterial abundances only increased following a second cyanobacterial bloom, peaking in August. Uptake rate constants for the different measured carbon compounds varied seasonally and inter-annually and were highly correlated to bacterial productivity estimates, temperature, and cyanobacterial abundances. Further, we detected nutrient limitation in response to environmental conditions in a multitude of microbial variables, such as elevated productivities in nutrient bioassays, changes in enzymatic activities, or substrate preferences. Variations among biotic variables often occurred on time scales of days to a few weeks, yet often spanning several sampling occasions. Such dynamics might not have been captured by sampling at monthly intervals, as compared to more predictable transitions in abiotic variables such as temperature or nutrient concentrations. Our study indicates that high resolution analyses of microbial biomass and productivity parameters can help out in the development of biogeochemical and food web models disentangling the microbial black box.

The microbial part of the pelagic food web is seldom characterized in models despite its major contribution to biogeochemical cycles. In the Baltic Sea, spatial and temporal high frequency sampling over three years revealed changes in heterotrophic bacteria and phytoplankton coupling (biomass and production) related to hydrographic properties of the ecosystem. Phyto- and bacterioplankton were bottom-up driven in both coastal and offshore areas. Cold winter temperature was essential for phytoplankton to conform to the successional sequence in temperate waters. In terms of annual carbon production, the loss of the spring bloom (diatoms and dinoflagellates) after mild winters tended not to be compensated for by other taxa, not even summer cyanobacteria. These results improve our ability to project Baltic Sea ecosystem response to short- and long-term environmental changes.Electronic supplementary materialThe online version of this article (doi:10.1007/s13280-015-0662-8) contains supplementary material, which is available to authorized users.

Vitamin B 1 is an essential exogenous micronutrient for animals. Mass death and reproductive failure in top aquatic consumers caused by vitamin B 1 deficiency is an emerging conservation issue in Northern hemisphere aquatic ecosystems. We present for the first time a model that identifies conditions responsible for the constrained flow of vitamin B 1 from unicellular organisms to planktivorous fishes. The flow of vitamin B 1 through the food web is constrained under anthropogenic pressures of increased nutrient input and, driven by climatic change, increased light attenuation by dissolved substances transported to marine coastal systems. Fishing pressure on piscivorous fish, through increased abundance of planktivorous fish that overexploit mesozooplankton, may further constrain vitamin B 1 flow from producers to consumers. We also found that key ecological contributors to the constrained flow of vitamin B 1 are a low mesozooplankton biomass, picoalgae prevailing among primary producers and low fluctuations of population numbers of planktonic organisms.

Top predators in several aquatic food webs regularly display elevated reproductive failure, caused by thiamin (vitamin B1) deficiency. The reasons for these low‐thiamin levels are not understood and information about the transfer of thiamin from the producers (bacteria and phytoplankton) to higher trophic levels is limited. One main concern is whether cyanobacterial blooms could negatively affect thiamin transfer in aquatic systems. Laboratory experiments with Baltic Sea plankton communities and single phytoplankton species were used to study the effect of filamentous cyanobacteria on the transfer of thiamin from phytoplankton to zooplankton. Experiments showed that the thiamin content in copepods was reduced when exposed to elevated levels of cyanobacteria, although filamentous cyanobacteria had higher levels of thiamin than any other analyzed phytoplankton species. Filamentous cyanobacteria also had a negative effect on copepod egg production despite high concentrations of non‐cyanobacterial food. Phytoplankton species composition affected overall thiamin concentration with relatively more thiamin available for transfer when the relative abundance of Dinophyceae was higher. Finally, phytoplankton thiamin levels were lower when copepods were abundant, indicating that grazers affect thiamin levels in phytoplankton community, likely by selective feeding. Overall, high levels of thiamin in phytoplankton communities are not reflected in the copepod community. We conclude that presence of filamentous cyanobacteria during summer potentially reduces the transfer of thiamin to higher trophic levels by negatively affecting phytoplankton and copepod thiamin content as well as copepod reproduction, thereby lowering the absolute capacity of the food web to transfer thiamin through copepods to higher trophic levels.

Thiamin (vitamin B 1) is mainly produced by bacteria and phytoplankton and then transferred to zooplankton and higher trophic levels but knowledge on the dynamics of these processes in aquatic ecosystems is lacking. Hence, the seasonal variation in thiamin content was assessed in field samples of copepods and in pico-, nano-and micro-plankton of two size classes (0.7-3 µm and > 3 µm) collected monthly in the Baltic Sea during 3 years and in the Skagerrak during 1 year. Copepods exhibited species-specific concentrations of thiamin and Acartia sp. had the highest carbon-specific thiamin content, at both locations. Even members of the same genus, but from different systems contained different levels of thiamin, with higher thiamin content per specimen in copepods from the Skagerrak compared to congeners from the Baltic Sea. Furthermore, our results show that the small plankton (0.7-3 µm) had a higher carbon-specific thiamin content compared to the large (> 3 µm). Additionally, there was a large seasonal variation and thiamin content was highly correlated comparing the two size fractions. Finally, there was an overall positive correlation between thiamin content in copepods and plankton. However, for periods of high thiamin content in the two size fractions, this correlation was negative. This suggests a decoupling between thiamin availability in pico-, nano-and micro-plankton and zooplankton in the Baltic Sea. Knowledge about concentrations of this essential micronutrient in the aquatic food web is limited and this study constitutes a foundation for further understanding the dynamics of thiamin in aquatic environments.

Thiamin (vitamin B1) is primarily produced by bacteria and phytoplankton in aquatic food webs and transferred by ingestion to higher trophic levels. However, much remains unknown regarding production, content and transfer of this water-soluble, essential micronutrient. Hence, the thiamin content of six phytoplankton species from different taxa was investigated, along with the effect of thiamin amendment on thiamin content. Furthermore, thiamin transfer to copepods was estimated in feeding experiments. Prey type, not phytoplankton thiamin content per se, was the most important factor for the transfer of thiamin, as it was lowest from filamentous Cyanophyceae and highest from more easily ingested prey like Dunaliella tertiolecta and Rhodomonas salina. Cyanophyceae had the highest thiamin content of the investigated species, eightfold higher than the lowest. Phytoplankton varied in thiamin content related to the supply of thiamin, where thiamin addition enabled higher thiamin content in some species, while copepod thiamin content was less variable. In all, thiamin transfer is not only dependent on the prey thiamin content, but also the edibility and/or digestibility is of importance. Thiamin is essential for all organisms, and this study constitutes an important building block to understanding the dynamics and transfer of thiamin in the aquatic food web.

Vitamin B 1 (thiamin) deficiency is an issue periodically affecting a wide range of taxa worldwide. In aquatic pelagic systems, thiamin is mainly produced by bacteria and phytoplankton and is transferred to fish and birds via zooplankton, but there is no general consensus on when or why this transfer is disrupted. We focus on the occurrence in salmon (Salmo salar) of a thiamin deficiency syndrome (M74), the incidence of which is highly correlated among populations derived from different spawning rivers. Here, we show that M74 in salmon is associated with certain large-scale abiotic changes in the main common feeding area of salmon in the southern Baltic Sea. Years with high M74 incidence were characterized by stagnant periods with relatively low salinity and phosphate and silicate concentrations but high total nitrogen. Consequently, there were major changes in phytoplankton and zooplankton, with, e.g., increased abundances of Cryptophyceae, Dinophyceae, Diatomophyceae and Euglenophyceae and Acartia spp. during high M74 incidence years. The prey fish communities also had increased stocks of both herring and sprat in these years. Overall, this suggests important changes in the entire food web structure and nutritional pathways in the common feeding period during high M74 incidence years. Previous research has emphasized the importance of the abundance of planktivorous fish for the occurrence of M74. By using this 27-year time series, we expand this analysis to the entire ecosystem and discuss potential mechanisms inducing thiamin deficiency in salmon.

Although free-living (FL) and particle-attached (PA) bacteria are recognized as ecologically distinct compartments of marine microbial food-webs, few, if any, studies have determined their dynamics in abundance, function (production, respiration and substrate utilization) and taxonomy over a yearly cycle. In the Baltic Sea, abundance and production of PA bacteria (defined as the size-fraction >3.0 μm) peaked over 3 months in summer (6 months for FL bacteria), largely coinciding with blooms of Chitinophagales (Bacteroidetes). Pronounced changes in the growth efficiency (range 0.05–0.27) of FL bacteria (defined as the size-fraction <3.0 μm) indicated the magnitude of seasonal variability of ecological settings bacteria experience. Accordingly, 16S rRNA gene analyses of bacterial community composition uncovered distinct correlations between taxa, environmental variables and metabolisms, including Firmicutes associated with elevated hydrolytic enzyme activity in winter and Verrucomicrobia with utilization of algal-derived substrates during summer. Further, our results suggested a substrate-controlled succession in the PA fraction, from Bacteroidetes using polymers to Actinobacteria and Betaproteobacteria using monomers across the spring to autumn phytoplankton bloom transition. Collectively, our findings emphasize pronounced seasonal changes in both the composition of the bacterial community in the PA and FL size-fractions and their contribution to organic matter utilization and carbon cycling. This is important for interpreting microbial ecosystem function-responses to natural and human-induced environmental changes.