Made available in DSpace on 2014-12-17T14:03:42Z (GMT). No. of bitstreams: 1 JoseEGJ.pdf: 604842 bytes, checksum: a34879bd40606d248f800a49ed111824 (MD5) Previous issue date: 2006-12-06 / This report shows 2232 times purification of a βNAcetylhexosaminidase from hepatic extracts from the sea mammal Sotalia fluviatilis homogenate with final recovery of 8,4%. Sequenced steps were utilized for enzyme purification: ammonium sulfate fractionation, Biogel A 1.5 m, chitin, DEAESepharose and hydroxyapatite chromatographies. The protein molecular mass was estimated in 10 kDa using SDSPAGE and confirmed by MALDITOF. It was found to have an optimal pH of 5.0 and a temperature of 60?C. Using pnitrophenylNAcetylβDglycosaminide apparent Km and Vmax values were of 2.72 mM and 0.572 nmol/mg/min, respectively. The enzyme was inhibited by mercury chloride (HgCl2) and sodium dodecil sulfate (SDS) / Este trabalho mostra a purifica??o de 2232 vezes de uma βNAcetilhexosaminidase obtida a partir dos extratos hep?ticos do mam?fero marinho Sotalia fluviatilis com recupera??o final de 8,4%. Passos seq?enciais foram utilizados para a purifica??o enzim?tica: fracionamento com sulfato de am?nio e as cromatografias de Biogel A 1.5 m, Quitina, DEAESepharose e Hidroxiapatita. A massa molecular prot?ica foi estimada em 10 kDa usando SDSPAGE e confirmada por MALDITOF. Foi encontrado como pH e temperatura ?timos, 5,0 e 60?C, respectivamente. Os valores de Km e Vm?x aparentes foram 2,72 mM e 0,572 nmol/mg/min, sendo utilizado o pnitrofenilNAcetilβDglicosamin?deo como substrato. A enzima foi inibida pelo cloreto de merc?rio (HgCl2) e dodecil sulfato de s?dio (SDS)

Dams play an important role in our society, but their negative impacts on aquatic ecosystems have led to a need for conservation efforts to mitigate the negative effects. In some instances, dam removal has been suggested to be the most economically and ecologically beneficial restoration method to improve longitudinal connectivity. This, however, has increased the need of more research about the effects dam removal have on aquatic ecosystems, since the outcomes of this restoration method can be unpredictable. In this study, I investigated ecological impacts of dam removals in Swedish inland waters. I collected field data from five different lakes in Värmland and Dalarna, two lakes with dam removal in 2017/2018 and three without. I studied how dam removal affected different trophic levels of the aquatic ecosystem, where I predicted that a top-down effect would increase abundances of piscivorous European perch (Perca fluviatilis) and zooplankton, but decrease phytoplankton, whereas a bottom-up effect would lead to increases in biomass at all trophic levels (with most pronounced effects on nutrient concentration). If both top-down and bottom-up effects would operate in parallel, all the above mentioned effects could occur. I also predicted that if dam removal affected the fish community, there would be a change in species diversity and fish biomass. Moreover, if perch demography was affected, there would be a change in perch length and abundance of different age classes. To investigate this, I collected data on fish, plankton, nutrients (P, N), DOC, pH and conductivity in the five lakes and compared this with already existing historical data using a BACI design. Two-way ANOVAs were used to study the effects of dam removals on these variables. There were not enough historical data on P concentration or zoo-and phytoplankton to be analyzed statistically. I did, however, conduct correlation analyses with zoo- and phytoplankton. Contradictory to my prediction, the results showed no effect of dam removal on fish biomass or diversity, water chemistry or perch demography in the studied lakes. There was, however, a statistically significant effect of time on Tot-N, with lower values after the years of 2017/2018 (regardless of dam removal). There were also some differences between the lakes in species diversity, concentration of Tot-N, potentially piscivorous perch and perch length. The outcomes of dam removal are unpredictable, as are the expected time-lapse of the effects, which means that changes may yet occur in the studied lakes. Knowledge about ecosystem responses to dam removal enables managers to better predict the effects of dam removal, and thereby facilitate decision makings related to this restoration method. / Dammar spelar en viktig roll i vårt samhälle, men deras negativa påverkan på akvatiska ekosystem har ökat behovet av bevarandeinsatser för att minska de negativa effekterna. I somliga fall har dammutrivning föreslagits vara den mest ekonomiskt och ekologiskt gynnsamma bevarandemetoden för att förbättra longitudinell konnektivitet. Detta har dock ökat behovet av mer forskning gällande dammutrivningars effekter på akvatiska ekosystem, eftersom utfallen av denna bevarandemetod kan vara oförutsägbara. I denna studie undersökte jag dammutrivningars ekologiska effekter i svenska inlandsvatten. Jag samlade data i fält från fem olika sjöar i Värmland och Dalarna, två sjöar med dammutrivningar under 2017/2018 och tre utan. Jag undersökte hur dammutrivning påverkade olika trofiska nivåer i det akvatiska ekosystemet, med förväntningarna att en top-down-effekt skulle öka abundansen av fiskätande abborre (Perca fluviatilis) samt djurplankton, men minska växtplankton, medan en bottom-up-effekt skulle leda till ökad biomassa inom samtliga trofiska nivåer (men med mest effekter på näringskoncentration). Om både top-down- och bottom-up-effekter skulle verka parallellt kunde samtliga ovanstående nämnda effekter förekomma. Jag förväntade även att om dammutrivning påverkade fisksamhället skulle det ske en förändring i artdiversitet och fiskbiomassa. Därtill bör en effekt på demografi hos abborre leda till förändringar i abborrlängd- och abundans i olika åldersklasser. För att undersöka detta samlade jag in data över fisk, plankton, näringsämnen (P, N), DOC, pH och konduktivitet i de fem sjöarna och jämförde detta med redan existerande historiska data med en BACI design. Tvåvägs-ANOVAs användes för att studera effekterna av dammutrivningar på dessa variabler. Det fanns inte tillräckligt med historiska data över P-koncentration eller djur- och växtplankton för att analyseras statistiskt. Jag utförde dock korrelationsanalyser med djur- och växtplankton. I kontrast till min förväntan visade resultaten ingen effekt av dammutrivning på fiskbiomassa- eller diversitet, vattenkemi eller demografi hos abborre i de studerade sjöarna. Det var däremot en statistiskt signifikant effekt av tid på Tot-N, med lägre värden efter 2017/2018 (oberoende av dammutrivning). Det var även en del skillnader mellan sjöarna i artdiversitet, koncentration av Tot-N, potentiellt fiskätande abborre samt abborrlängder. Utfallen av damutrivningar är oförutsägbara, och detsamma gäller tidsintervallet för effekterna, vilket innebär att det fortfarande kan ske förändringar i de studerade sjöarna. Kunskap om ekosystemrespons till dammutrivning möjliggör för förvaltare att lättare bedöma effekter av dammutrivningar, och underlättar därav beslutsfattning relaterat till denna bevarandemetod.

<p>This thesis reports the results of biomarker measurements in three environmental monitoring projects. In the first project, which was part of the Swedish national environmental monitoring, biomarkers were measured annually in female perch (<i>Perca fluviatilis</i>) in two background areas in the Baltic Sea during 1988–2000, resulting in a unique 13-year series of measurements. The most important results were a strong decreasing temporal trend in the gonadosomatic index (GSI) and a strong increasing temporal trend in the hepatic ethoxyresorufin O-deethylase (EROD) activity in the Baltic Proper. In the second project, biomarkers and concentrations of classic pollutants were measured in female perch in the Stockholm recipient 1999–2001. This was the first time a large city was investigated as a point source of pollution, and the gradient was longer and included more stations than customary. Severe pollution conditions in central Stockholm were indicated by the poor health status of the perch: retarded growth, decreased frequency of sexually mature females, low GSI, disturbed visceral fat metabolism, increased hepatic EROD activity, decreased muscle acetylcholinesterase activity, increased frequency of hepatic DNA adducts, and a high concentration of biliary 1-pyrenol. Muscle ΣDDT and ΣPCB were measured as pollution indicators and were 10–28 respectively 12–35 times higher than the background levels in perch from the Baltic Proper. In the Stockholm archipelago two superimposed gradients were found. Whereas the response of several biomarkers consistently decreased with increasing distance from central Stockholm, the response of others first decreased from Stockholm to the middle archipelago and then increased to the open Baltic Sea. The latter biomarkers included the frequency of sexually mature females, GSI, hepatic EROD activity, and hepatic DNA adducts. In the third project, potential toxicity from munitions on the seafloor, at a dumpsite in the Stockholm archipelago, was analysed by the nanoinjection of sediment extracts into newly fertilised rainbow trout (<i>Oncorhynchus mykiss</i>) eggs, followed by the measurement of biomarkers in the developing larvae. No biological effects of the dumped munitions were found. The same stations in the Stockholm archipelago as in the second project were investigated as a positive control. The results of the three projects agreed well, which demonstrated the continuous pollution of the Baltic Sea and the severe pollution conditions and adverse biological effects in central Stockholm. Further investigations are urgently needed to identify which pollutants or other factors are causing the observed biological effects, both in the background areas in the Baltic Sea and in the Stockholm recipient.</p>

This thesis reports the results of biomarker measurements in three environmental monitoring projects. In the first project, which was part of the Swedish national environmental monitoring, biomarkers were measured annually in female perch (Perca fluviatilis) in two background areas in the Baltic Sea during 1988–2000, resulting in a unique 13-year series of measurements. The most important results were a strong decreasing temporal trend in the gonadosomatic index (GSI) and a strong increasing temporal trend in the hepatic ethoxyresorufin O-deethylase (EROD) activity in the Baltic Proper. In the second project, biomarkers and concentrations of classic pollutants were measured in female perch in the Stockholm recipient 1999–2001. This was the first time a large city was investigated as a point source of pollution, and the gradient was longer and included more stations than customary. Severe pollution conditions in central Stockholm were indicated by the poor health status of the perch: retarded growth, decreased frequency of sexually mature females, low GSI, disturbed visceral fat metabolism, increased hepatic EROD activity, decreased muscle acetylcholinesterase activity, increased frequency of hepatic DNA adducts, and a high concentration of biliary 1-pyrenol. Muscle ΣDDT and ΣPCB were measured as pollution indicators and were 10–28 respectively 12–35 times higher than the background levels in perch from the Baltic Proper. In the Stockholm archipelago two superimposed gradients were found. Whereas the response of several biomarkers consistently decreased with increasing distance from central Stockholm, the response of others first decreased from Stockholm to the middle archipelago and then increased to the open Baltic Sea. The latter biomarkers included the frequency of sexually mature females, GSI, hepatic EROD activity, and hepatic DNA adducts. In the third project, potential toxicity from munitions on the seafloor, at a dumpsite in the Stockholm archipelago, was analysed by the nanoinjection of sediment extracts into newly fertilised rainbow trout (Oncorhynchus mykiss) eggs, followed by the measurement of biomarkers in the developing larvae. No biological effects of the dumped munitions were found. The same stations in the Stockholm archipelago as in the second project were investigated as a positive control. The results of the three projects agreed well, which demonstrated the continuous pollution of the Baltic Sea and the severe pollution conditions and adverse biological effects in central Stockholm. Further investigations are urgently needed to identify which pollutants or other factors are causing the observed biological effects, both in the background areas in the Baltic Sea and in the Stockholm recipient.

BAKGRUND: Per- och polyfluorerade alkylsubstanser, PFAS, är ett samlingsnamn för en stor grupp mycket stabila substanser. Vissa PFAS är klassade som persistenta, bioackumulerbara och toxiska (PBT-ämnen) och har en negativ hälsoeffekt på människor och djur. De är vanligt förekommande i exempelvis brandskum och förhöjda halter finns därför ofta i vattensystem som ligger nära brandövningsområden. Perflourooktansyra (PFOS) är idag den enda PFAS som är förbjuden inom EU och är förmodligen den mest toxiska. Tolerabla gränsvärden finns idag endast för två PFAS; PFOS och PFOA. En av de största källorna för humanexponering av PFOS är fiskkonsumtion, medan PFOA inte brukar vara detekterbar i fisk. SYFTE: Syftet med denna studie var att analysera humanexponering av 11 olika PFAS (PFOS, PFOA, PFHxA, PFHxS, PFHpA, PFNA, PFDA, PFDS, PFBS, PFUnDA och PFOSA) efter konsumtion av abborre (Perca fluviatilis), baserad på analysdata från Stockholms stads miljögiftsövervakning. Då fokus i arbetet framförallt ligger på PFOS så har uppskattat PFOS-intag via denna exponeringsväg jämförts med tillgängliga toxikologiska referensvärden. Likaså syftar arbetet att undersöka hur PFOS- exponeringen via konsumtion av sötvattenfisk förhåller sig till bakgrundsexponering av PFOS i normalbefolkningen. METOD: PFOS-exponeringen via konsumtion av sötvattenfisk beräkandes för tvånivåer av fiskkonsumenter; ”mest sannolik exponeringsnivå” (MSE, där man äteregenfångad sötvattenfisk max 6 ggr per år), och ”hög exponeringssnivå” (HE, där manäter egenfångad sötvattenfisk 1 ggr/vecka). En vidare uppdelning gjordes på”normalkonsumenter” (person som äter normalstora portioner och har en medelvikt) och ”storkonsumenter” (person med låg vikt som äter stora portioner). Beräkningarna grundar sig på tre års analysdata (2015-2017) på abborre från 14 olika vattensystem kring Stockholm stad, vilka kan betraktas som representativa fiskevatten i en urban miljö, utan känd PFOS-påverkan från punktkälla. PFOS-analyser i fisk användes tillsammans med data från litteraturen över fiskkonsumtion och kroppsvikt. Litteraturdata användes även för att beräkna exponeringen i normalbefolkningen viaandra livsmedel och dricksvatten. Även här gjordes beräkningarna för ”mest sannolik exponering” (normalkonsumtion av dricksvatten och normalexponering av PFOS via övriga livsmedel) och ”hög exponering” (storkonsumtion av dricksvatten och högexponering via övriga livsmedel). Därefter jämfördes resultatet med tolerabla dagliga intag (TDI) och PFOS-exponeringen via fiskkonsumtion relaterades till det PFOS-intag som normalbefolkningen blir utsatt för via konsumtion av dricksvatten och andra livsmedel. RESULTAT: I de analyserade proverna av abborrmuskel stod PFOS för ca 90% av PFAS-innehållet, PFOS är den förening som exponeringsberäkningarna gjorts för. Bakgrundsexponeringen för PFOS i normalbefolkningen, via konsumtion av dricksvatten och andra livsmedel (även köpt fisk), beräknades till ca 0,5 ng/kg/dag (vuxna) och 0,7 ng/kg/dag (barn) för den mest sannolika exponeringsnivån (MSE). För den högre exponeringsnivån (HE) beräknades intaget till ca 1,7 ng/kg/dag för både vuxna och barn – alltså väl under EFSAs rådande TDI-värde på 150 ng/kg/dag. Vid jämförelse med dessa bakgrundsnivåer visade sig följande för de 4 exponeringsnivåer som karakteriserats vid konsumtion av egenfångad insjöfisk: MSE – normalkonsument: I denna grupp var PFOS-exponeringen via konsumtionen av egenfångad sötvattenfisk omkring 0,5 ng/kg/dag (vuxna) och 1,1 ng/kg/dag (barn), innebärandes att redan individer som konsumerar egenfångad fisk 6 ggr/år dubblerar sin PFOS-exponering. Det totala intaget ligger dock långt ifrån rådande TDI-värde. MSE – storkonsument: För vuxna och barn beräknades PFOS-exponeringen via sötvattenfisk till ca 1 respektive 2 ng/kg/dag. HE – normalkonsument: För kvinnor blev PFOS-exponeringen 17 ng/kg/dag och för män 18 ng/kg/dag medan resultatet för barn var 37 ng/kg/dag. HE – storkonsument: PFOS-exponeringen från insjöfiskkonsumtion, i detta ”worst-case-scenario”, var 27 och 33 ng/kg/dag för kvinnor respektive män och 69 ng/kg/dag för barn. Hos denna grupp blev exponeringen runt 15 gånger högre än i normalbefolkningens hos vuxna och 40 gånger hos barn. DISKUSSION: Att PFOS stod för den största delen PFAS stämmer väl med förväntat resultat. Studien visar att normal- och storkonsumtion, inom ”den mest sannolika exponeringsnivån”, av sötvattenfisk från de vattensystem i närheten av Stockholm stadsom omfattats av studien, inte utgör någon hälsofara enligt de TDI som gäller idag även om det ger en procentuellt sett stor ökning av PFOS-intaget relativt vad som normalt är att förvänta via intag av livsmedel (även dricksvatten inräknat). Storkonsumenter inomkategorin ”hög exponeringsnivå” utgör ett worst case-scenario, där barn i gruppen kommer upp i ungefär halva TDI. Vid eventuell revidering av TDI så finns risk att både normal- och storkonsumenter, inom ”hög exponeringsnivå”, av sötvattenfisk hamnar påPFOS-intag runt eller över TDI. / BACKGROUND: Per- and polyfluorinated alkyl substances (PFAS) is a group of very stable molecules. Some PFAS are classed as persistant, bioaccumulative and toxic (PBT-substances) with negative impacts on humans and animals. PFAS is a common ingredient in fire foam, and elevated levels are consequently often seen in water systems near fire drill areas. Today, perflourooctanoic acid (PFOS) is the only prohibited PFAS within the EU, and probably the most toxic.Tolerable daily intake (TDI) recommenadtions are only available for two PFAS within the EU; PFOS and PFOA. One of the main exposure routes of PFOS in humans are consumption of fish, whilePFOA usually isn’t detectable in fish. OBJECTIVES: The aim of this study was to analyze human exposure of 11 different PFAS (PFOS, PFOA, PFHxA, PFHxS, PFHpA, PFNA, PFDA, PFDS, PFBS, PFUnDA och PFOSA) after consumption of european perch (Perca fluviatilis) collected from water systems in an urban region of Sweden. The main focus substance of the study was PFOS, where different scenarios of PFOS-exposure after consumtion of perch were compared with toxicological reference values (TDIs) and with background exposure from consumtion of other foods and drinking water. METHODS: The PFOS-exposure from consumption of freshwater fish was calculatedfor two levels of consumers; a “most likely exposure level” (MSE, with a consumptionrate of maximum 6 times/year), and a “high exposure level” (HE, with a consumption rate of at least 1 time/week). The groups were further divided into “normal consumers” (who ate normal portions and had normal body weight) and “large consumers” (who atelarge portions and had a low body weight). The caclulations were based on three years of analysis data (2015-2017) on european perch from 14 different water systems around the city of Stockholm, which can be considered as representative fishing waters in an urban environment, without known PFOS point source. The analyzed PFOS concentrations in fish were used in conjunction with literature data on fish consumption and body weight to assess exposure. Data from the litterature were also used to calculate the exposure in the normal population, after consumption of other foods and drinkingwater. Also here a “most likely exposure level” (average consumtion of drinking water and average consumption rates and PFOS concentrations in other foods) and “high exposure level” (large consumer of drinking water and high PFOS exposure from other foods) were calculated. Thereafter, the results were compared with the tolerable daily intake (TDI), and the PFOS-exposure through consumption of perch was compared to the PFOS-intake following consumtion of other foods and drinking water. RESULTS: PFOS accounted for about 90% of the total PFAS in european perch. PFOS is the compound for which the exposure calculations were made.Background exposure by PFOS in the normal population, through consumtion of drinking water and other foods (including commercial fish), was calculated to 0,5 ng/kg/day (adults) and 0,7 ng/kg/day (children) for the most likely exposure level (MSE). För the higher exposure level (HE) the intake was calculated to 1,7 ng/kg/day for both adults and children, which is far below EFSA ́s current TDI of 150 ng/kg/day. When the background levels are related to the 4 exposure levels characterized after consumption of self-caught freshwater fish the results showed that: MSE – normal consumer: The PFOS exposure from consumption of freshwater fish in this group was 0,5 ng/kg/day (adults) and 1,1 ng/kg/day (children), which means that by consuming self-caught fish 6 times/year the PFOS-exposure will be doubled. Even so, the total intake is far below current TDI. MSE – high consumer: PFOS exposure from freshwater fish were 1 ng/kg/day and 2 ng/kg/day for adults and children respectively. HE – normal consumer: The PFOS exposure for women were 17 ng/kg/day and 18 ng/kg/day for men while the result form children were 37 ng/kg/day. HE – high consumer: The PFOS exposure in this “worst case scenario” was 27 and 33ng/kg/day for women and men respectively and 70 ng/kg/day for children. In this group the exposure were about 15 times higher than the background exposure for adults and 40 times higher for children. CONCLUSIONS: That PFOS accounted for the largest part of PFAS in fish is in accordance with previously published studies. Normal and large consumption offreshwater fish within the “MSE”-group, of freshwater fish, from the water systems near the city of Stockholm which are covered in the study, is not a health risk according to the TDI that apply today. However, it gives a percentually large increase in PFOS intake relative to what is normally expected from food intake (including drinkingwater). High consumers in the “HE”-group constitute a worst case scenario, where children in the group reach a PFOS-exposure of about half of TDI. In the event of a possible revision of TDI also normal consumers in the “HE”-group may close in to, or pass, the TDI-limits.

The work in this thesis deals with the ecology and evolution of adaptive individual variation. Ecologists have long used niche theory to describe the ecology of a species as a whole, treating conspecific individuals as ecological equivalent. During recent years, research about individual variation in diet and morphology has gained interest in adaptive radiations and ecological speciation. Such variation among individual niche use may have important conservation implications as well as ecological and evolutionary implications. However, up to date we know very little about the extension of this phenomenon in natural populations and the mechanisms behind it. The results in this thesis show that the extension of individual diet specialization is widely spread throughout the animal kingdom. The variation in diet is mainly correlated to morphological variation but not always. Furthermore, this variation in diet and morphology among individuals could be both genetically determined and environmentally induced and it mainly comes from trade-offs in foraging efficiency between different prey types. The results from a number of studies of perch also show that individual perch differ in morphology and diet depending on habitat, where littoral perch has a deeper body compared to pelagic perch. This difference in morphology corresponds to functional expectations and is related to foraging efficiency trade-offs between foraging in the littoral and pelagic zone of a lake. The variation in morphology in perch is mainly due to phenotypic plasticity but there are also small genetic differences between the littoral and pelagic perch. Two separate studies show that both predation and competition may be important mechanism for the variation in morphology and diet in perch. In conclusion, the results in this thesis show that individual variation in diet and habitat choice is a common phenomenon with lots of ecological and evolutionary implications. However, there are many mechanisms involved in this phenomenon on which we are just about to start learning more about, and only further research in this area will give us the full insight.

This thesis was written within the framework of a biomanipulation project where young-of-the-year (YOY) pikeperch (Sander lucioperca) were stocked to a Baltic Sea bay to improve water quality through a top-down trophic cascade. The aim of my doctorial studies was however focused on a broader ecological question, namely predation (the main driving force in a biomanipulation). Hence, this thesis consists of four papers where we study the interactions between predator and prey using fish and zooplankton and how these interactions can be measured. In paper I we evaluated the performance of different diet analysis methods by individual based modelling and found that when having a nutritional gain perspective, mass based methods described diets best. Paper II investigated how the explorative, foraging and anti-predator behaviour of the YOY pikeperch used for stocking were affected by their rearing environment (pond vs. tank rearing). The more complex and varied environment in the semi-natural ponds seemed to promote a more flexible and active behaviour, better equipping young fish for survival in the wild. For paper III we studied the diel vertical migration in the six copepodite stages of the zooplankton Acartia spp. and Eurytemora affinis in relation to fish biomass, phytoplankton abundance and temperature. Both species migrated and in addition showed increased migration range with size within species, indicating evasion from visual predators. Paper IV addressed the movement of littoral Eurasian perch (Perca fluviatilis) via stable isotope signatures (13C and 15N) and body condition. We found clear indications of sedentarity and intra-habitat dietary differences. Interactions between predators and prey are complex and affected by both physiological and environmental characteristics as well as behavioural traits. The results in this thesis suggest that different species and even different life stages pursue different strategies to survive. / At the time of doctoral defense, the following papers were unpublished and had a status as follows: Paper 1: Submitted.  Paper 2: In press.  Paper 4: Submitted.

It is increasingly recognized that individuals of the same species differ from each other and influence and respond to their environment in unique ways. This thesis deals with size variation among individuals that not only are of the same species but also of similar age. Such variation may develop even when individuals are born in the same environment, i.e. within a cohort. I have studied the sources and consequences of variation within and among cohorts from egg through early ontogeny using young-of-the-year (YOY) perch (Perca fluviatilis) as study organism. In agreement with predictions based on model results only taking exploitative interactions among individuals into account, I found that the broader the initial size distributions were, the more did the degree of size variation among individuals decrease over time. Still, with initially small size variation among individuals, in several experiments also size divergence was observed. Furthermore, size variation among individuals increased more under high compared to at low densities. Increased size variation over time may be explained by size-dependent diet shifts allowing for initially larger individuals to make an early diet shift when the first resource becomes limiting. However, as size divergence also was observed in situations with only shared resources available, it can be concluded that diet shifts are not a prerequisite for size divergence in young animal cohorts. Hence, I also suggest that mechanisms not related to competition for limiting resources, such as genetic variation, stochasticity and behavioural traits must be taken into account, especially when initial size differences are small. The importance of considering size variation among individuals within cohorts was demonstrated in a study of winter mortality in YOY perch cohorts. A large individual size in autumn was shown to increase overwinter survival within cohorts. However, late summer growth rather than average body size reached in autumn explained variation in overwinter survival between cohorts. Higher accumulation to lipid reserves and accordingly lower mortality over winter was observed in years with high growth rates late in the season. In another study I showed that apparent patterns of density-dependent growth can emerge among larval fish, but rather than a result of density-dependent resource limitation this was due to variation in size-selective predation pressure. Individuals in the right end of the size distributions grew in to a high predation pressure from cannibalistic perch when cannibal density was high, coinciding with high larval perch densities. Finally, as substantial size variation among individuals can develop within cohorts, also intra-cohort cannibalism can occur. Using a physiologically structured population model it was shown that the development of size bimodality within cohorts as a result of intra-cohort cannibalism is critically dependent on long hatching periods, high victim densities and density-dependent feedbacks on shared resources. / Det faktum att individer som tillhör samma art skiljer sig från varandra och påverkar och påverkas av sin omgivande miljö på ett unikt sätt tillskrivs allt större betydelse inom ekologin. Den här avhandlingen handlar framför allt om storleksvariation mellan individer som förutom att tillhöra samma art dessutom tillhör samma årsklass. Sådan storleksvariation kan till och med utvecklas mellan individer som föds och växer upp i samma miljö (inom en kohort). Jag har studerat orsaker bakom och konsekvenser av variation inom och mellan kohorter. Som studieorganism har jag använt mig av årsyngel av abborre (Perca fluviatilis). I överensstämmelse med förutsägelser baserade på en modell som enbart tar hänsyn till konkurrens om en gemensam resurs visade det sig såväl i ett dammexperiment som i en naturlig sjö att ju bredare den initiala storleksfördelningen var desto mer minskade graden av variation i kroppsstorlek mellan individer över tid. Å andra sidan, när den initiala variationen var relativt liten observerades i flera oberoende experiment även storleksdivergens över tid mellan individer. Variationen i storlek ökade särskilt i miljöer med höga tätheter av konsumenter (abborrar). För att förstå de bakomliggande mekanismerna av sådana täthetseffekter måste man ta hänsyn till den återkoppling som sker mellan antalet konsumenter och mängden resurser. Ökad storleksvariation över tid skulle kunna förklaras med storleksberoende dietskiften som tillåter individer med en initial storleksfördel att genomgå ett tidigt dietskifte samtidigt som tillgången av den första resursen begränsar övriga individers tillväxt. Eftersom storleksdivergens även observerades i situationer där enbart en delad resurs var tillgänglig kan man dock dra slutsatsen att dietskiften inte är en förutsättning för storleksdivergens inom kohorter. Jag föreslår därför också att mekanismer som inte är relaterade till konkurrens om en begränsad resurs, såsom inneboende variation mellan individer och variation i beteendemönster bör beaktas för att förklara uppkomsten av storleksvariation, speciellt i de fall då den initiala variationen är liten. De resultat som presenteras visar på betydelsen av att ta hänsyn till storleksvariation mellan individer. Storlek efter den första tillväxtsäsongen var viktig för att förklara vilka individer inom en kohort av årsyngel som överlever sin första vinter. För att förklara variation i vinteröverlevnad mellan kohorter (mellan år och sjöar) var däremot hög tillväxt sent på säsongen (oberoende av medelstorlek på hösten) avgörande. Högre ackumulering av fettreserver och lägre mortalitet inom kohorter av årsyngel under vintern observerades under år med hög tillväxt under den senare delen av tillväxtsäsongen. I en annan studie visade jag att mönster som tyder på täthetsberoende tillväxt kan uppkomma hos fiskyngel men att detta fenomen snarare än täthetsberoende resursbegränsning förklaras av variation i storleksberoende predationstryck. De större individerna inom kohorterna växte in i ett högt predationstryck från kannibalistiska abborrar när tätheten av kannibaler var hög, något som samkorrelerade med höga tätheter av yngel. Slutligen, då en hög grad av storleksvariation mellan individer kan utvecklas inom kohorter är även kannibalism mellan individer inom en kohort möjligt. Genom att använda mig av en så kallad fysiologiskt strukturerad populationsmodell kunde jag visa att divergerande tillväxtkurvor mellan kannibaler och deras byten (vilket resulterar i storleksbimodalitet) som ett resultat av kannibalism inom kohorter är beroende av den tid det tar för ynglen att kläcka ut, antalet bytesfiskar per kannibal samt den återkoppling som finns mellan konsumenterna och deras gemensamma resurs (djurplankton).

The seasonal and diel feeding habits of the goosander, Mergus merganser, the red-breasted merganser, M. serrator, gulls (Larus canus, L. argentatus and L. fuscusj and terns, Sterna hirundo/paradisaea were studied at 64V05'N. Birds' activity patterns were influenced by the nocturnal spawning of the river lamprey (Lampetra fluviatilis). Food selection and food consumption by hand-raised mergansers together with aquarium studies of the diel activity patterns of their most important prey supplemented the field data. River lamprey dominated the diet of the goosander by weight and the sculpin Cottus gobio by number. The fish consumption of the goosanders was found to be about 12% of the available river lamprey biomass and about 17% of the sculpin biomass during the breeding season. In experimental situations the river lamprey was a low- pritority species compared with salmon , Salmo salar, brown trout, 53. trutta, and minnow, Phoxinus phoxinus, when presented to satiated birds of both Mergus species. When hungry, however, the birds caught the available prey irrespective of species, but they selected larger prey when two size classes were present. Experimental results were compared with field data on availability, consumption and the escape behaviour of the various fish species. The rivers emptying in the Bothnian Bay are regarded as important feeding areas for birds breeding along the coast. In early spring they get access to abundant and reliable food resources, e.g the river lamprey. Later on the three-spined stickleback, Gasterosteus aculeatus, supply both Mergus species with food in the river and also along the coast. From the middle of June potential food supply available to birds decreases in the rivers and becomes more abundant in the coastal area. / <p>S. 1-41: sammanfattning, s. 43-227: 7 uppsatser</p> / digitalisering@umu

Genetically structured populations arise when gene flow between groups of individuals is hindered by geographical, behavioural or temporal barriers. The identification of such groups is important for understanding evolution and has large implications for conservation concern. The field of population subdivision has received a lot of interest throughout the years and gained empirical support from a number of species. However, very little is known about population structure at small spatial scales, especially in a highly mobile species such as fish. The main object for my thesis was to further investigate population differentiation, explicitly at small spatial scales in the Eurasian perch. My results show that in this species, genetic differentiation occurs, even at very small spatial scales, both within lakes and in the Baltic Sea. Additionally, the differentiation can be stable over years and thus have a large impact in the evolution of adaptation to different environments. I also found barriers to gene flow that overlapped with the largest change in spring temperature, suggesting a temporal difference in spawning. Morphological differences were found at these small scales as well which indicates that a difference in food resources might be an underlying cause of change. My thesis work shows that the aquatic environment might not be as homogenous as widely thought and that there could be barriers or adaptations to different environments that hinder the fish from genetic panmixia. Slight patterns of isolation by distance (IBD) were found in the Baltic Sea, implying that the distance (i.e. currents) effect the level of differentiation via drifting of larvae and/or small fish. However, I have also theoretically investigated the IBD model of and seen that it is no longer correct when differences in population sizes are introduced. The pattern of IBD can mean high levels of gene flow or no gene flow at all, solely dependent on population size differences and fluctuations. My thesis has resulted in new and important findings regarding the existence and cause of genetic differentiation at very small spatial scales and thus added new knowledge into the field of evolution and speciation. In addition, my results also give insights into the contemporary state of the Eurasian perch and future evolutionary potential.