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Climate change is projected to cause elevated precipitation in northern Europe, leading to increased runoff of terrestrial matter to coastal areas. The consequences for food web production and ecosystem function remain unclear. A mesocosm experiment was performed to investigate the impacts of elevated terrestrial matter input, using a natural plankton community from the northern Baltic Sea with added young-of-the-year perch as planktivorous top consumer. Addition of terrestrial matter caused water browning and increased dissolved organic carbon and inorganic nutrient concentrations. Phytoplankton primary production showed a negative response to terrestrial matter due to decreased light availability, while heterotrophic bacterial production increased. The trophic balance, calculated as the difference between primary production and heterotrophic bacterial production, indicated that net-heterotrophy was triggered by terrestrial matter enrichment. Primary production was identified as the main basal energy source for fish. Addition of terrestrial matter reduced the food web efficiency, calculated as the ratio between fish production and basal production (primary production?+?heterotrophic bacterial production). Furthermore, stable isotope analysis of seston and fish indicated that the added terrestrial matter was not efficiently incorporated in the food web and only marginally altered the food web trophic positions. The results suggest that the main food chain consisted of phytoplankton, mesozooplankton, and fish, and that the ecosystem production was overall light driven. Under a changing climate, several negative effects can be expected, including a poorer light climate, reduced ecosystem production and net-heterotrophy. These alterations have potentially significant consequences for ecosystem functioning, fish production, and thus ecosystem services.

Climate change scenarios project that precipitation will increase in northern Europe, causing amplified inflows of terrestrial matter (tM) and inorganic nutrients to coastal areas. How this will affect the plankton community is poorly understood. A mesocosm experiment was carried out to investigate the influence of two levels of tM inputs on the composition, size-structure and productivity of a natural coastal phytoplankton community from the northern Baltic Sea. The tM addition caused browning of the water and decreased underwater light levels, while the concentrations of dissolved organic carbon (DOC) and inorganic nutrients increased. Microphytoplankton were promoted by tM addition, while in the controls picophytoplankton dominated the phytoplankton community. Inorganic nutrient availability was instrumental in defining the phytoplankton community composition and size-structure. As a response to tM addition, the phytoplankton increased their chlorophyll a content. This physiological adaptation helped to maintain high primary production rates at the low tM enrichment, but at the high tM load the primary production decreased as did the biomass of mesozooplankton. The ciliate biomass was high when the mesozooplankton biomass was low, indicating that a trophic cascade occurred in the system. Structural equation modeling showed that tM borne DOC promoted ciliates, while primary and bacterial production were disfavored. Thus, DOC originating from soils had an indirect negative effect on the mesozooplankton by reducing their food availability. Although, a positive correlation between heterotrophic bacteria and phytoplankton suggested coupling between phytoplankton produced carbon and heterotrophs growth. The results from our study indicate that river-borne DOC and inorganic nutrients have a large impact on the phytoplankton community, driving the system to the dominance of large diatoms. However, since river-borne humic substances cause browning of the water, phytoplankton increase their light harvesting pigments. At moderate inflow this helps to support the primary production, but at high inflows of terrestrial material the primary production will decrease. As high river inflows have been projected to be a consequence of climate change, we foresee that primary production will decrease in coastal areas in the future, and the impacts of such changes on the food web could be significant.

Climate change projections indicate increased precipitation in northern Europe, leading to increased inflow of allochthonous organic matter to aquatic systems. The food web responses are poorly known, and may differ depending on the trophic structure. We performed an experimental mesocosm study where effects of labile dissolved organic carbon (DOC) on two different pelagic food webs were investigated, one having zooplankton as highest trophic level and the other with planktivorous fish as top consumer. In both food webs, DOC caused higher bacterial production and lower food web efficiency, i.e., energy transfer efficiency from the base to the top of the food web. However, the top-level response to DOC addition differed in the zooplankton and the fish systems. The zooplankton production increased due to efficient channeling of energy via both the bacteria land the phytoplankton pathway, while the fish production decreased due to channeling of energy mainly via the longer and less efficient bacterial pathway. We conclude that the added DOC either acted as a subsidy by increasing the production of the top trophic level (mesozooplankton), or as a sink causing decreased top consumer production (planktivorous fish).

Heavy rainfall events causing significant terrestrial runoff into coastal marine ecosystems are predicted to become more frequent with climate change in the Mediterranean. To simulate the effects of soil runoff on the pelagic food web of an oligotrophic Mediterranean coastal lagoon, we crossed soil extract addition (increasing nutrient availability and turbidity) and fish presence in a full factorial design to coastal mesocosms containing a natural pelagic community. Soil extract addition increased both bacteria and phytoplankton biomass. Diatoms however profited most from soil extract addition, especially in the absence of fish. In contrast zooplankton and fish did not profit from soil extract addition. Furthermore, our data indicate that nutrients (instead of light or carbon) limited basal production. Presumed changes in carbon availability are relatively unimportant to primary and secondary production in strongly nutrient limited systems like the Thau Lagoon. We conclude that in shallow Mediterranean coastal ecosystems, heavy rainfall events causing soil runoff will (1) increase the relative abundance of phytoplankton in relation to bacteria and zooplankton, especially in the absence of fish (2) not lead to higher biomass of zooplankton and fish, possibly due to the brevity of the phytoplankton bloom and the slow biomass response of higher trophic levels.

Predicted increases in runoff of terrestrial dissolved organic matter (DOM) and sea surface temperatures implicate substantial changes in energy fluxes of coastal marine ecosystems. Despite marine bacteria being critical drivers of marine carbon cycling, knowledge of compositional responses within bacterioplankton communities to such disturbances is strongly limited. Using 16S rRNA gene pyrosequencing, we examined bacterioplankton population dynamics in Baltic Sea mesocosms with treatments combining terrestrial DOM enrichment and increased temperature. Among the 200 most abundant taxa, 62 % either increased or decreased in relative abundance under changed environmental conditions. For example, SAR11 and SAR86 populations proliferated in combined increased terrestrial DOM/temperature mesocosms, while the hgcI and CL500-29 clades (Actinobacteria) decreased in the same mesocosms. Bacteroidetes increased in both control mesocosms and in the combined increased terrestrial DOM/temperature mesocosms. These results indicate considerable and differential responses among distinct bacterial populations to combined climate change effects, emphasizing the potential of such effects to induce shifts in ecosystem function and carbon cycling in the future Baltic Sea.

The three-spined stickleback Gasterosteus aculeatus is a small omnivorous fish, widely distributed in the northern hemisphere. In the Baltic Sea, recently observed increases in their population densities have been attributed to declines of piscivorous predators. Concurrent with this predator release, an alternative hypothesis is that past and present consumption rates and resource limitation thresholds may have contributed to the recent increases in stickleback abundance. To evaluate this hypothesis, we estimated the size- and temperature-dependent attack rate and the critical resource density (CRD) of three-spined sticklebacks. We incorporated laboratory results with time series of zooplankton abundance to estimate historical trends in degrees of resource limitation in sticklebacks and evaluate if increases in individual consumption rates could be a plausible mechanism facilitating the suggested population increase. Attack rates increased with body size and temperature in laboratory experiments. Estimated CRD increased with size but decreased with temperature, suggesting that stickleback scope for individual and population growth might increase at temperatures above 15 degrees C. Our results further suggest that sticklebacks have been living closer to maximum consumption capacity in the coastal areas of the Bothnian Sea (BS) and Bothnian Bay (BB). Moreover, decreasing levels of resource limitation in the corresponding off-shore zones may have facilitated increases in stickleback densities for these areas. However, in the coastal zones of the Baltic proper (BP), resource levels have declined and are approaching the CRD, suggesting that stickleback populations in BP may not increase further. The decrease in CRD with temperature implies that increasing summer temperatures will increase the scope of individual and population growth in the three-spined stickleback and may favor the three-spined stickleback's competitive ability over other species under a warmer climate. (C) 2013 Elsevier B.V. All rights reserved.

Both temperature and terrestrial organic matter have strong impacts on aquatic food-web dynamics and production. Temperature affects vital rates of all organisms, and terrestrial organic matter can act both as an energy source for lower trophic levels, while simultaneously reducing light availability for autotrophic production. As climate change predictions for the Baltic Sea and elsewhere suggest increases in both terrestrial matter runoff and increases in temperature, we studied the effects on pelagic food-web dynamics and food-web efficiency in a plausible future scenario with respect to these abiotic variables in a large-scale mesocosm experiment. Total basal (phytoplankton plus bacterial) production was slightly reduced when only increasing temperatures, but was otherwise similar across all other treatments. Separate increases in nutrient loads and temperature decreased the ratio of autotrophic:heterotrophic production, but the combined treatment of elevated temperature and terrestrial nutrient loads increased both fish production and food-web efficiency. CDOM: Chl a ratios strongly indicated that terrestrial and not autotrophic carbon was the main energy source in these food webs and our results also showed that zooplankton biomass was positively correlated with increased bacterial production. Concomitantly, biomass of the dominant calanoid copepod Acartia sp. increased as an effect of increased temperature. As the combined effects of increased temperature and terrestrial organic nutrient loads were required to increase zooplankton abundance and fish production, conclusions about effects of climate change on food-web dynamics and fish production must be based on realistic combinations of several abiotic factors. Moreover, our results question established notions on the net inefficiency of heterotrophic carbon transfer to the top of the food web.

Both temperature and terrestrial dissolved organic carbon (TDOC) have strong impacts on aquatic food web dynamics and production. Temperature affects vital rates of all organisms and terrestrial carbon has been shown to alter the dynamics of phytoplankton and bacterial production and affect the trophic structure of planktonic food webs. As climate change predictions for the Baltic Sea suggests future increases in both terrestrial carbon run-off and increases in temperature, the aim of thesis was to adopt a system-ecological approach and study effects of these abiotic variables, not only on interactions within planktonic food webs, but also on the growth and consumption rates of one of the most common zooplanktivorous fish in the Baltic Sea, the three-spined stickleback Gasterosteus aculeatus. Results showed that three-spined sticklebacks display a high degree of resilience against increasing temperatures, as both growth rates as well as consumption rates on zooplankton were high at temperatures well over 20 °C. Furthermore, it was shown that the minimal resource densities required to sustain individual and population growth, actually decreased with increasing temperatures, implying that sticklebacks around their optimum temperature for growth at 21 °C will actually have an increased scope for growth. As stickleback population densities have increased over the last decade in the Baltic Sea and are now suggested to out-compete other coastal fish species for shared zooplankton resources, the results presented in this thesis suggest that increased water temperatures would only serve to increase sticklebacks competitive advantage. As the structuring role of this small zooplanktivore on pelagic communities might be considerable, further studies investigating competitive interactions as well as patterns of population abundances are definitely warranted. TDOC was overall shown to stimulate bacterial production and the microbial food web. Because of the longer trophic pathways required to transport carbon from bacterial production to higher trophic levels, the addition of TDOC always reduced food web transfer efficiency. However, it became apparent that the full effect of TDOC additions on pelagic food webs was complex and depended heavily not only on the existing trophic structure to which the carbon was introduced, but also on ambient temperature levels. When three-spined sticklebacks were part of food webs with significant TDOC inputs, the presence of fish, indirectly, through predator release of lower trophic levels, amplified the magnitude of the effects of carbon addition on bacterial production, turning the base of the system significantly more heterotrophic, which ultimately, impacted negatively on their own production. However, when a pelagic food web containing sticklebacks was simultaneously subjected to realistic increases in temperature and TDOC concentrations, food web efficiency and fish production increased compared to present day conditions. These results were explained by a temperature dependent increased production potential of zooplankton, sustained by an increased production of heterotropic microzooplankton via TDOC additions, which lead to higher fish production. Although the increased number of trophic linkages in heterotrophic food webs should have reduced energy transfer efficiency, these negative effects seem here to have been overridden by the positive increases in zooplankton production as a result of increased temperature. These results show that heterotrophic carbon transfer can be a viable pathway to top-consumers, but also indicates that in order to understand the full effects of climate change on trophic dynamics and fish production, abiotic variables cannot be studied in isolation. 

Specific growth rates of individually reared juvenile three-spined sticklebacks Gasterosteus aculeatus were investigated under laboratory conditions to parameterize a complete temperature-dependent growth model for this species. To test the applicability of experimentally derived optima in growth response rates to natural conditions, the effects of commercial pellets and natural prey on growth rates were investigated. In addition, to test for seasonal effects on growth, laboratory trials were performed in both spring and winter. Growth took place from 5 to 29° C with a temperature for optimum growth reaching a sharp peak at 21° C. Modelled optimal temperature for maximum growth was estimated to be 21.7° C and lower and upper temperatures for growth were estimated to be 3.6 and 30.7° C, respectively. There were no significant differences in growth rates between fish reared on invertebrates or commercial pellets. Seasonal effects on growth were pronounced, with reduced growth rates in the winter despite similar laboratory conditions. On average, 60% higher growth rates were achieved at the optimum temperature in summer compared to the winter. The strong seasonality in the growth patterns of G. aculeatus indicated here reduces the applicability of the model derived in this study to spring and summer conditions.