Biological invasion has become a major threat to economy, ecology, global biodiversity and ecosystem function of aquatic ecosystems. The main aim of the thesis was to study the effects of the zebra mussel (Dreissena polymorpha), a versatile invasive species, on phytoplankton dynamics and ecosystem function of lakes. In a first attempt, I compared the density of Dreissena and the physicochemical data of ecosystems that it invaded among North American and European lakes to identify important factors in its invasion success. Secondly, I investigated the impact of zebra mussels on phytoplankton community composition in a natural lake. Thirdly, I evaluated whether zebra mussel feeding behavior were affected by the presence of predatory waterborne cues. Finally, I examined the effect of Dreissena on seston stoichiometry. A Generalized Additive Model revealed that a joint effect of surface area, mean depth, total phosphorus and calcium concentrations can explain the variability in Dreissena density. Selective grazing by zebra mussels varied in relation to seasonal phytoplankton dynamics. Risk cues released by predators affected both feeding rate and prey selection of the mussels and had cascading indirect effects on phytoplankton biomass and community structure. I found that the flux in nutrients caused by differences in zebra mussel consumption lead to a variation in phytoplankton nutrient limitation. The flexibility of zebra mussel feeding behavior and variation in susceptibility among phytoplankton groups to mussel ingestion indicate that invading zebra mussels could alter phytoplankton community composition of lakes and have important ecosystem consequences. The results of this thesis contribute to the growing evidence that predators indirectly affect resource dynamics and food web structure through their non-lethal effects on consumers. The results suggested that zebra mussel can indirectly both reduce and increase the energy transfer efficiency from primary producers to upper trophic levels in the pelagic and benthic food webs, respectively.

Anthropogenic activities are increasing inorganic nitrogen (N) loadings to lakes in the northern hemisphere. In many boreal lakes phytoplankton are N limited, wherefore enhanced N input may affect the productivity of pelagic food webs. Simultaneously, global change causes increased inflows of terrestrial dissolved organic carbon (DOC) to boreal lakes. Between clear and humic lakes, whole lake primary and consumer production naturally differs. However, research is inconclusive as to what controls pelagic production in these lakes. Further, it is unclear how DOC affects the response of the pelagic food web to enhanced inorganic N availability. The overarching goal of this thesis was to study the effects of inorganic N and organic C for pelagic food webs in boreal lakes. In the thesis, I first identified the main drivers of pelagic production during summer in eight non-manipulated Swedish boreal lakes with naturally low or high DOC. Then I investigated how increased N availability affects the pelagic food chain, and how the response differs with DOC. Therefore, whole lake inorganic N fertilization experiments were conducted in six Swedish boreal lakes across a DOC gradient (low, medium, high) divided into three lake pairs (control, N enriched) with one reference and two impact years. In each lake, I also investigated the response of zooplankton growth using in situ mesocosm experiments excluding planktivores. I found that humic boreal lakes had lower phytoplankton production and biomass than clear water lakes. Further, phytoplankton community composition and food quality differed with DOC. However, high DOC did not reduce pelagic energy mobilization or zooplankton biomass, but promoted a higher dominance of cladoceran relative to copepod species. N addition clearly enhanced phytoplankton biomass and production in the experimental lakes. However, this stimulating N effect decreased with DOC as caused by light limitation. Further, the newly available phytoplankton energy derived from N addition was not efficiently transferred to zooplankton, which indicates a mismatch between producer energy supply and consumer energy use. Indeed, the mesocosm experiment revealed that decreased food quality of phytoplankton in response to N addition resulted in reduced food web performance, especially in clearer lakes. In humic lakes, zooplankton production and food web efficiency were clearly more resilient to N addition. In summary, my thesis suggests that any change in the landscape that enhances inorganic N availability will especially affect pelagic food webs in clear water lakes. In contrast, brownification will result in more lakes being resilient to eutrophication caused by enhanced N deposition.

<p>Biological invasion has become a major threat to economy, ecology, global biodiversity and ecosystem function of aquatic ecosystems. The main aim of the thesis was to study the effects of the zebra mussel <i>(Dreissena polymorpha)</i>, a versatile invasive species, on phytoplankton dynamics and ecosystem function of lakes. </p><p>In a first attempt, I compared the density of <i>Dreissena</i> and the physicochemical data of ecosystems that it invaded among North American and European lakes to identify important factors in its invasion success. Secondly, I investigated the impact of zebra mussels on phytoplankton community composition in a natural lake. Thirdly, I evaluated whether zebra mussel feeding behavior were affected by the presence of predatory waterborne cues. Finally, I examined the effect of <i>Dreissena</i> on seston stoichiometry.</p><p>A Generalized Additive Model revealed that a joint effect of surface area, mean depth, total phosphorus and calcium concentrations can explain the variability in <i>Dreissena</i> density. Selective grazing by zebra mussels varied in relation to seasonal phytoplankton dynamics. Risk cues released by predators affected both feeding rate and prey selection of the mussels and had cascading indirect effects on phytoplankton biomass and community structure. I found that the flux in nutrients caused by differences in zebra mussel consumption lead to a variation in phytoplankton nutrient limitation.</p><p>The flexibility of zebra mussel feeding behavior and variation in susceptibility among phytoplankton groups to mussel ingestion indicate that invading zebra mussels could alter phytoplankton community composition of lakes and have important ecosystem consequences. The results of this thesis contribute to the growing evidence that predators indirectly affect resource dynamics and food web structure through their non-lethal effects on consumers. The results suggested that zebra mussel can indirectly both reduce and increase the energy transfer efficiency from primary producers to upper trophic levels in the pelagic and benthic food webs, respectively. </p>