Abstract This study investigated the ecophysiology of one of the world's northernmost Atlantic salmon (Salmo salar L.) stocks in the River Teno. The juvenile years of salmon of same genetic background were studied in the wild and in the hatchery conditions. In addition, the maternal size effect on reproduction was studied in wild females. Benefit of body size was not only quantitative but also qualitative in reproduction success of the wild female salmon in the River Teno. Total number of eggs and energy content of eggs were higher in big females than in smaller ones. In the hatchery, under natural day length and water temperature conditions, the growth rate, liver glycogen content and condition factor of the parr was higher than in the wild. The liver glycogen content of the hatchery-reared parr increased throughout the growing season and decreased during winter, whereas that of the wild parr was the lowest in summer, and stayed relative stable from September to May. The observed differences in annual fluctuation in liver glycogen content may reflect the differences in carbohydrate content of feed and in behaviour between the hatchery and wild. Overall, the hatchery-reared juveniles maturated and smoltificated 1–2 years earlier than the wild fish. The mean age of wild precocious males was 3 years and that of wild smolts 4 years. However, there was considerable variation in the age of precocious males (1–6 yr) and smolts (2–8 yr) in the wild. The maturing and smolting juvenile age groups were restricted to two (1–2 yr and 2–3 yr, respectively) in the hatchery. In June, the hypo-osmoregulatory ability of hatchery smolts was developed parallel to the wild smolts. Some differences in physiological parameters between different smolt groups could be observed in the wild and between hatchery-reared and wild smolts indicating that completing of smolting process varies to some extent under different conditions. However, the hatchery-reared smolts showed higher levels of fin damage and body energy stores than the wild smolts. As the hatchery practices should aim at controlling the quantity and quality of the juvenile salmon in production, the environmental conditions governing the physiological development of the juvenile fish should be taken into account. Hatchery practices should be planned so that the seasonal timing of smolting would follow the wild fish as close as possible.

Abstract Long-term monitoring of the wild salmon (Salmo salar L.) stock of the Simojoki river and the stocked hatchery-reared salmon parr and smolts has provided a considerable amount of information on the development of the stock and factors affecting it. Data on the relationships between wild and reared salmon were collected by tagging and trapping both smolts and adult salmon having either a wild or reared background. The tag recapture rate of wild smolts was about twice as high as that of smolts stocked as two-year-olds and slightly greater than for smolts stocked as parr. When survival was measured in relation to the smolt size, the difference between the wild and reared smolts was even greater, and it seemed to be emphasized in years with a low survival rate. The difference observed between the wild and reared salmon in the smolt phase generally disappeared in the adult phase. When adult salmon returned to the river to spawn the difference in the timing of the ascent depended more on the age or sex of the salmon, and less on their origin. Similarly, the survival of adult salmon in the river before or after spawning and later after returning to the sea depended on the sex and age of the fish. The origin of fish affected their behaviour, the reared salmon wandering more than wild adults before settling down into spawning areas. When the yield of wild and reared smolts as returning adults was compared, the wild smolts gave the best results, although the survival from smolt to adult was low in all smolt groups, probably due to the high fishing pressure in the sea. The smolts stocked as parr and those stocked as two-year-old fish were similar in this respect. The former group gave better results if the yield was measured as the number of returning multi-sea-winter adults, while the latter group gave better results if one-sea-winter grilse were also included in the yield. The low yield of adult salmon from stocking and the generally low survival of smolt groups irrespective of their origin emphasises the importance of fishing regulations as a tool in the maintenance or enhancement of naturally reproducing salmon stocks. However, despite the low profitability of stocking, it probably safeguarded the existence of the wild Simojoki salmon stock during its critical phase in the early 1990s. At that time the fishing regulations were not strict enough to prevent the alarming decrease in the salmon stock, and the adult spawners produced by stocking of young salmon may have had a relatively higher value than their number suggests.

Competition is defined as the demand of two or more organisms for the same extrinsic resources in excess of supply. The distribution, movements, behaviour and food of trout and shiners in Paul and Pinantan lakes were studied in order to determine the item's and mechanisms of interspecific competition between them. Data from recent years were compared with data for years when trout alone inhabited the lake. No interspecific aggression was observed. The possibility that the two species were competing for space was discounted. Stomach contents of shiners in Pinantan Lake revealed a marked qualitative diurnal food cycle. In Paul Lake, shiners have drastically reduced the Gammarus population relative to its pre-shiner abundance. This overgrazing was caused by the concentration of large numbers of shiners over the shoals where Gammarus are also present in their highest concentrations and the ability of shiners to pursue food deeper into the weeds and to graze an area more thoroughly than trout. In Pinantan Lake shiners have apparently-reduced the density of Daphnia to a point where trout are unable to feed on them as rapidly as in pre-shiner years. The ability of both species to utilize many types of food tends to reduce the intensity of competition. The study demonstrates how false implications may arise from a delayed appraisal of competition. If observations had not been made on Paul Lake until after competition had been observed the importance of Gammarus as an item of competition would have probably been overlooked and the whole competitive relationship misconstrued. Included among the basic mechanisms of competition is the consumption by one or more organisms of something in short supply before it reaches a potential habitat where it would become available to another organism or group. Environmental factors and behaviour were shown to be important influences in the dynamics of competition. The physical and biological environment and the distribution and behaviour of competitors may be in states of continual flux. Hence natural competitive relationships can be considerably more complicated and variable than situations described by the most elastic of theoretical models. / Science, Faculty of / Zoology, Department of / Graduate

La smoltification est un évènement métamorphique chez le saumon qui initie la migration de dévalaison et pré-adapte le juvénile à l’entrée en mer. Cette thèse a pour objectif d’étudier les régulations endocrines et environnementales de la smoltification chez le saumon Atlantique de la longue rivière Loire-Allier, population qui est en danger. Nous montrons la présence et la divergence fonctionnelle de deux paralogues de la sous-unité ß (tshßa & tshßb) de la thyrotropine (TSH) chez le saumon Atlantique et observons un pic d’expression de tshßb dans l’hypophyse à la smoltification, pic concomitant à l’initiation de la migration de dévalaison. Ce résultat est le premier à mettre en relation l’expression hypophysaire de TSH avec la smoltification et le comportement migratoire de dévalaison. L’exposition expérimentale à une photopériode constante de jours courts ou à une température augmentée n’affecte pas nettement le pic de tshßb ni l’initiation de la migration de dévalaison, ce qui met en évidence l’importance de contrôles endogènes. Cette étude apporte de nouvelles connaissances fondamentales sur le cycle de vie du saumon Atlantique avec la découverte de nouveaux acteurs dans le processus de smoltification et avec des implications dans le domaine de la conservation. / Smoltification is a metamorphic event in salmon, which initiates downstream migration and pre-adapts juvenile for seawater entry. The PhD aimed at investigating endocrine and environmental regulation of smoltification in the endangered long-river Loire-Allier Atlantic salmon. We report the presence and functional divergence of thyrotropin ß-subunit paralogs (tshßa & tshßb) in Atlantic salmon and showed a peak pituitary expression of tshßb at smoltification which was concomitant with the initiation of downstream migration. This is the first time pituitary TSH expression is related to smoltification and downstream migratory behavior. Experimental exposure to constant short-day photoperiod or to increased temperature did not markedly affect the peak of tshßb nor the initiation of downstream migration, highlighting the importance of endogenous controls. This study brings new insights to the life cycle of Atlantic salmon with the discovery of novel components of the smoltification process, and with implications for conservation.

Diss. (sammanfattning). Umeå : Sveriges lantbruksuniv. / Härtill 6 uppsatser.

Winter has been generally considered as a bottleneck period for salmonid populations, but recent studies show it might be more context related. The purpose of this study was to examine how surface ice changes spatial distribution of juvenile one-year-old brown trout in a small boreal stream. I hypothesized that the presence of surface ice will allow a more even distribution of trout over the entire width of the stream while in the absence of ice, trout will be more heavily associated with near-edge habitats. I also hypothesized trout will be more evenly dispersed at night over the width of the stream even in the absence of surface ice. My results show a strong positive correlation between icecover and spatial distribution. In the presence of surface ice trout use the whole width of the stream, while in the absence of ice the middle regions of the stream were almost completely devoid of fish. My results also show there was no difference in the spatial distribution between night and day in the presence of ice cover, but in the absence of ice cover trout were more tightly associated with the stream edge during day whereas at night they were more evenly dispersed over the entire width of thestream. / Vinter har i allmänhet ansetts vara en flaskhals-period för laxfiskar, men nya studier visar att det kan vara mer sammanhangsrelaterat. Syftet med denna studie var att undersöka hur istäcket påverkar distribution av unga ett-åriga öring längs vattendragets bredd i en liten boreal ström. Jag hypotiserade att närvaron av istäcket kommer att möjliggöra en jämnare fördelning av öring över strömmens bredd. Min andra hypotes var att öring kommer att sprida sig mer jämnt på natten över strömmens bredd även vid frånvaro av istäcke. Mina resultat visar en stark positiv korrelation mellan istäckningsgrad och rumslig distribution. Vid närvaro av istäcke använde öring mer effektivt habitater över hela bredden av vattendraget, medans i frånvaro var de habitater närmare mitten av vattendraget nästan helt tomma av fisk. Mina resultat visar också att det inte fanns någon skillnad i rumslig fördelning mellan natt och dag i närvaro av istäcke men i frånvaro av istäcke var öring mer tätt associerad med strömkant habitater under dagen och på natten var de mer jämnt spridda över bredden av strömmen.

Les co-produits sont les parties non utilisées et récupérables lors des opérations de transformation du poisson, comme les têtes, les peaux, les os ou la pulpe. L’industrie de transformation du saumon Atlantique d’élevage (Salmo salar) génère environ 50% de co-produits qui constituent une source de protéines de haute qualité. Ce travail de thèse s’inscrit dans le cadre du projet Pesk&Co, qui réunit quatre partenaires industriels (Meralliance-Thai Union, Yslab, SPF-DIANA, AGH-Socofag) et un partenaire académique (LEMAR UMR 6539, UBO) dans le but d’extraire et de caractériser des ingrédients à haute valeur ajoutée issus des co-produits de saumon Atlantique d’élevage (Salmo salar). Le premier objectif de la thèse visait à développer un procédé d’extraction du collagène à partir des peaux de saumon par la mise en place d'un procédé non conventionnel au LEMAR, puis développé au stade pilote et enfin industriel. Le collagène obtenu a été caractérisé par différentes méthodes analytiques (SEC-FPLC, SDS-PAGE, FTIR, rhéologie, microscopie). Ensuite, des essais de réticulation du collagène par voie enzymatique avec une transglutaminase microbienne ont conduit à l’obtention d’un hydrogel. Le deuxième objectif de ce travail a porté sur l’hydrolyse en conditions contrôlées des têtes de saumon pour générer des peptides fonctionnels pour l’aquaculture. Deux protocoles d’hydrolyse enzymatique ont été développés et transposés à l’échelle pilote. Les hydrolysats ont été incorporés dans des aliments aquacoles afin d’être testés sur des larves de bar (Dicentrarchus labrax). Les deux ingrédients développés au cours de ce travail ont pour vocation future d'être commercialisés, et différents marchés et applications sont visés. / By-products are the not used parts and recoverable in the fish processing operations, such as heads, skins, bones or pulp. The processing industry of farmed Atlantic salmon (Salmo salar) generates about 50% of co-products, which are a source of high quality protein. This work is part of Pesk&Co project, which gather four industrial partners (Meralliance-Thai Union Yslab, SPF-DIANA, AGH-SOCOFAG) and one academic partner (LEMAR UMR 6539, UBO) in order to extract and characterize high value ingredients from farmed Atlantic salmon by-products (Salmo salar). The first aim of the thesis was to develop a method for extracting collagen from salmon skins by the setting up of a non-conventional process at LEMAR and finally developed it at pilot and industrial scale. The collagen obtained was characterized by different analytical methods (FPLC-SEC, SDS-PAGE, FTIR, rheology, microscopy). Then, enzymatic cross-linking assays of collagen with a microbial transglutaminase led to obtain a collagen hydrogel. The second objective of this work was focused on the hydrolysis under controlled conditions of salmon heads to generate functional peptides for aquaculture. Two enzymatic hydrolysis protocols have been developed and transferred at pilot scale. The hydrolysates were incorporated into diets to be tested on bass larvae (Dicentrarchus labrax). Both ingredients developed during this work have for future use to be commercialized and different markets and applications are targeted.

Vissa vattenkraftverk drivs med korttidsreglering som möjliggör snabb anpassning av elproduktion, vilket sker genom hastiga förändringar i vattenflöde. Flera olika typer av organismer påverkas negativt av korttidsreglering, till exempel fiskar. En möjlig konsekvens är strandning, vilket innebär att fiskar hamnar i torrlagt substrat eller annat ogynnsamt habitat. Risken för strandning påverkas av flera faktorer. I denna studie undersöktes hur strandningsrisk hos öringyngel (Salmo trutta) påverkas av tillgång till skydd och ljusförhållande, där prediktionerna var att strandningsrisken skulle vara större vid tillgång till skydd och i mörker. Dessa två faktorer testades i en 2x2 design. Det fanns en tendens till en större andel strandade yngel vid tillgång till skydd (48 ± 8,2 %) jämfört med när det inte fanns skydd (35,7 ± 23,7 %) och en större andel strandade vid ljusa förhållanden (50,2 ± 16 %) än i mörker (33,4 ± 17,3 %). Påverkan av tillgång till skydd på strandningsrisk var inte annorlunda vid de olika ljusförhållandena. Utifrån dessa resultat är rekommendationen för att minska strandningsrisken hos öringyngel att flödesreglering genomförs i mörker och att bottenstrukturen beaktas, vilket måste göras i varje enskilt fall. / Some hydropower stations use hydropeaking to be able to swiftly adjust electricity production, which is done through rapid alterations in water flow. Several types of organisms are negatively affected by hydropeaking, including fishes. One possible consequence of hydropeaking is stranding, which occurs when fish are separated from the water body and end up in dewatered substrate. The risk of stranding is influenced by several factors. The aim of this study was to investigate how stranding risk in brown trout fry (Salmo trutta) is affected by shelter availability and light conditions. The predictions were that stranding risk would be higher in conditions with available shelters and in darkness. The two factors were tested in a 2x2-design. There was a tendency for stranding rate to be higher when shelters were available (48 ± 8.2 %) compared to when there were none (35.7 ± 23.7 %), and stranding rate was higher in light (50.2 ± 16 %) than in darkness (33.4 ± 17.3 %). The effect of shelter availability on stranding risk did not differ between the two light treatments. Based on these results, in order to reduce stranding risk in brown trout fry the recommendation is that flow regulation should be carried out in darkness and that the bottom structure should be considered, which must be done on a case-by-case basis.

Salmonid fish such as brown trout (Salmo trutta) are known to adapt to local environments in large water systems such as in Lake Vänern. Since brown trout returns to spawn at the location of birth, they become reproductively isolated from populations spawning elsewhere. Thus, genetically-based physiological and ecological differences may develop between them. There is evidence that two of the stocks of Lake Vänern brown trout, Gullspångälven brown trout (G-trout) and Klarälven brown trout (K-trout) differ, with G-trout growing faster and becoming bigger than K-trout, but there is no study identifying why this is the case. The purpose of this study was therefore to investigate whether there were differences in three variables: metabolic rate (gill-beat frequency, referred to as ventilation rate), foraging rate (number of captured prey per unit time) and aggressive behavior (number of displays) between the two stocks of brown trout. By conducting laboratory experiments on G-trout and K-trout in artificial streams with a constant supply of food through drift, behavioral observations were made to quantify these variables. The results showed that G-trout had significantly higher gill-beat frequency and foraging rate than did K-trout. In terms of aggression there was no significant difference observed between the two stocks, though there was a trend for G-trout to show more display behavior. The differences in metabolic rate and foraging rate are consistent with the different growth rates reported for the two stocks. The differences between the stocks suggest that they have developed local adaptations and have maintained these after a number of years of being raised in hatcheries. The observed differences should be taken into account when it comes to management and restoration of populations adapted to local environments. / Laxfisk som till exempel öringen (Salmo trutta) är kända att anpassa sig till lokala miljöer i stora vattensystem, såsom i sjön Vänern. Eftersom öringen återvänder och leker där de är födda blir de reproduktivt isolerade från populationer som leker på annan plats. Därmed kan ärftliga fysiologiska och ekologiska skillnader utvecklas dem emellan. Där finns bevis på att Gullspångsöringen tillväxer snabbare och blir större än Klarälvsöringen men det finns ingen riktig vetskap om varför. Syftet med denna studie var därmed att undersöka om där fanns skillnader i tre variabler: metabolisk hastighet (gälslagsfrekvens), furagering (antal tagna byten) och aggressivt beteende (antal display uppvisanden) mellan öringstammarna. Med separata försök av G-öring och K-öring i ett försöksakvarium med en konstgjord vattenström och konstant tillförsel av mat genom drift, gjordes beteendeobservervationer av dessa variabler. Resultaten visade på att G-öring hade en signifikant högre gälslagsfrekvens och furageringshastighet än K-öring. Vad gällde displaybeteende fanns ingen signifikantskillnad mellan stammarna, dock observerades en trend att G-öringen uppvisade mer display. Dessa resultat tyder på att metabolisk hastighet och furagering kan ha en roll i tillväxttakten. Skillnaderna mellan stammarna kan tyda på att de har utvecklat lokala anpassningar och har kvar dessa efter ett antal års fiskodling. De observerade skillnaderna är viktiga att ta med i beräkning när det kommer till förvaltning och restauration av populationer anpassade till lokala miljöer.

The average global temperature is expected to rise by 3-5 °C at the end of the century, as a consequence of global warming. Negative effects are expected on poikilothermic animals, including fish, with changes in their physiology including metabolism. Metabolism has earlier been proven to have an association with propensities to migrate, which might be affected by raised temperatures as a consequence of global warming. The purpose of this study was to investigate if raised temperatures during incubation of eggs has an effect on metabolism in a partly migrating species of fish. The hypotheses for this study were: (i) ventilation rate (VR) will decrease as a consequence of warmer incubation temperatures, and (ii) offspring of anadromous parents are expected to have a higher VR compared to offspring of residential parents. VR, as a proxy for metabolic rate, was measured on young-of-the-year brown trout (Salmo trutta) during trials in respiration chambers. Fertilized eggs from four different crossings of parents (anadromous x anadromous, residential x residential, anadromous male x residential female, residential male x anadromous female) underwent incubation in normal and warm (normal + 3 °C) water temperatures respectively. A Two-way ANOVA showed a significant interaction between incubation temperature and crossing on VR. Warmer incubation temperatures in 3 out of 4 crossings resulted in a higher VR, except for individuals with two anadromous parents. Individuals with an anadromous father had a higher VR than individuals with a residential father. Evidence that partly supports both hypotheses was therefore found. Metabolism might have a strong genetic component, and other factors that might have an effect on metabolism and VR are epigenetics, stress, egg size etcetera. Studies of this kind will be of importance in getting a greater understanding of the effects of global warming on migrating species of fish, and on poikilothermic vertebrates in general.