<p>An influence at one trophic level can result in dynamic impacts also on other components of a food web. These dynamics are known as trophic cascades, and can be both top-down and bottom-up. After a near-collapse of the Baltic cod <i>Gadus morhua</i> stock in the 1980s, its main prey sprat <i>Sprattus sprattus</i> increased dramatically. The main food of sprat, marine copepods, decreased during the same time period, likely a combined effect of increased predation pressure from sprat and decreasing salinities. This shortage of food for sprat resulted in decreasing quality of sprat as a food source for common guillemots <i>Uria aalge</i>. However, a recent increase in fishing for sprat has again resulted in better feeding conditions for guillemots.</p><p>Human impacts on this simple food web can be complex. In the early 20th century, marine mammals were abundant and nutrient levels were low in the Baltic Sea. This thesis illustrate that this situation corresponded to lower fish biomass. A reduction of seals early in the century led to reduced top-down control, which resulted in increasing fish stocks. Later, in the 1950s, the largest inflow of salt water during the century mobilized accumulated phosphorus from the deep sediments, which stimulated nitrogen fixation. Combined with increasing anthropogenic nutrient loads, this led to increased primary production and a rapid change from an oligotrophic to a eutrophicated state. This change can be termed a regime shift, which also stimulated fish production. Subsequent over-fishing of cod likely caused a second regime shift, from a cod- to a clupeid- dominated state, which led to the described effects on the common guillemots.</p><p>Several factors affect the life-history of this long-lived seabird. Bycatches in gillnet fisheries is one factor directly affecting guillemot survival, and the proportion of bycatches increased during a period of increasing fishing effort. Surprisingly, avian cholera, a previously undocumented disease in common guillemots, was found at times to cause considerable adult mortality. Common guillemot life-history information can communicate the diversity of factors influencing marine ecosystems – hopefully this can increase our understanding of how complex even "simple" food webs are.</p>

The aim of this Thesis was to improve our understanding and assessment of feeding conditions for zooplanktivorous fish in the Baltic Sea. We investigated (papers I, II) the usefulness of biochemical proxies for assessments of growth and metabolic rates in the dominant Baltic copepod Acartia bifilosa. A predictive model (paper I) for egg production rate (EPR), based on body size, RNA content, and water temperature, was established using females of different geographical origin. This model demonstrates the usefulness of RNA content as a proxy for growth in zooplankton and, together with abundance data, it could be used to evaluate fish feeding conditions. Further (paper II), using A. bifilosa exposed to a food gradient, we evaluated responses of physiological rates and other biochemical proxies for growth and established correlations between physiological and biochemical variables. EPR and ingestion rate were most significantly correlated with RNA content. As assayed variables saturated at different food concentrations, food availability may affect assessments of physiological rates using proxies. In paper III, we explored the effect of high EPR and ingestion rate on astaxanthin content in A. bifilosa. We found that the astaxanthin content decreased at high feeding rates, most likely due to decreased assimilation efficiency. This may impact the quality of zooplankton as prey. The invasion of Cercopagis pengoi, a zooplanktivorous cladoceran, has altered the trophic linkages in the Baltic Sea food web. In paper IV, we evaluated the feeding of zooplanktivorous fish on C. pengoi and found that irrespective of size both herring and sprat feed on it, with large herring being more selective. In turn, C. pengoi feeds mainly on older copepods (paper V), which are acknowledged important in fish nutrition. These results indicate that C. pengoi may compete with fish due to the diet overlap. / <p>At the time of the doctoral defense, the following papers were unpublished and had a status as follows: Paper 2: In progress. Paper 3: Submitted</p>

An influence at one trophic level can result in dynamic impacts also on other components of a food web. These dynamics are known as trophic cascades, and can be both top-down and bottom-up. After a near-collapse of the Baltic cod Gadus morhua stock in the 1980s, its main prey sprat Sprattus sprattus increased dramatically. The main food of sprat, marine copepods, decreased during the same time period, likely a combined effect of increased predation pressure from sprat and decreasing salinities. This shortage of food for sprat resulted in decreasing quality of sprat as a food source for common guillemots Uria aalge. However, a recent increase in fishing for sprat has again resulted in better feeding conditions for guillemots. Human impacts on this simple food web can be complex. In the early 20th century, marine mammals were abundant and nutrient levels were low in the Baltic Sea. This thesis illustrate that this situation corresponded to lower fish biomass. A reduction of seals early in the century led to reduced top-down control, which resulted in increasing fish stocks. Later, in the 1950s, the largest inflow of salt water during the century mobilized accumulated phosphorus from the deep sediments, which stimulated nitrogen fixation. Combined with increasing anthropogenic nutrient loads, this led to increased primary production and a rapid change from an oligotrophic to a eutrophicated state. This change can be termed a regime shift, which also stimulated fish production. Subsequent over-fishing of cod likely caused a second regime shift, from a cod- to a clupeid- dominated state, which led to the described effects on the common guillemots. Several factors affect the life-history of this long-lived seabird. Bycatches in gillnet fisheries is one factor directly affecting guillemot survival, and the proportion of bycatches increased during a period of increasing fishing effort. Surprisingly, avian cholera, a previously undocumented disease in common guillemots, was found at times to cause considerable adult mortality. Common guillemot life-history information can communicate the diversity of factors influencing marine ecosystems – hopefully this can increase our understanding of how complex even "simple" food webs are.

The thesis combines laboratory experiments and field expeditions to study production, transfer and consumption of non-enzymatic antioxidants and thiamine in an aquatic food web. In particular, I (1) documented spatial and seasonal variation of tocopherols and carotenoids in the Baltic Sea pelagic food web, and (2) examined the effects of abiotic and biotic factors on tocopherol, carotenoid and thiamine concentrations in phytoplankton, zooplankton and fish. Moderate differences in temperature and salinity affected α-tocopherol, β-carotene and thiamine production in microalgae. Furthermore, the results suggest that acute stress favors the expression of non-enzymatic antioxidants rather than enzymatic antioxidants. Because production of α-tocopherol, β-carotene and thiamine differ markedly between microalgae, the availability of non-enzymatic antioxidants and thiamine is likely to be highly variable in the Baltic Sea and is difficult to predict. The transfer of non-enzymatic antioxidants from phytoplankton to zooplankton was biomass dependent. The field expeditions revealed that phytoplankton biomass was negatively associated with α-tocopherol concentration in mesozooplankton. Thus, increased eutrophication of the Baltic Sea followed by an increase in phytoplankton biomass could decrease the transfer of essential biochemicals to higher levels in the pelagic food web. This could lead to deficiency syndromes, of the kind already observed in the Baltic Sea. Astaxanthin is synthesized from precursors provided by the phytoplankton community. Thus biomass dependent transfer of astaxanthin precursors from phytoplankton to zooplankton could be responsible for astaxanthin deficiency in zooplanktivorous herring. Astaxanthin in herring consists mostly of all-Z-isomers, which are characterized by low bioavailability. Therefore, astaxanthin deficiency in salmon could be explained by the low concentration of this substance and its isomeric composition in herring.