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Papers 1,3,4 and 5 of this thesis are not available in Munin: 1. Cooper EJ, Dullinger S, Semenchuk PR: 'Late snowmelt delays plant development and results in lower reproductive success in the High Arctic', Plant science : an international journal of experimental plant biology (2011), vol.180:157–67, available at http://dx.doi.org/10.1016/j.plantsci.2010.09.005 3. Rumpf SB, Semenchuk PR, Dullinger S, Cooper EJ: 'Idiosyncratic Responses of High Arctic Plants to Changing Snow Regimes' (manuscript) 4. Semenchuk PR, Elberling B, Amtorp C, Winkler J, Rumpf S, Michelsen A, Cooper EJ: 'Deeper snow alters soil nutrient availability, leaf nutrient status and plant growth in high Arctic tundra' (manuscript) 5. Semenchuk PR, Christiansen CT, Morgner E, Grogan P, Elberling B, Cooper EJ: 'Long-term experimentally deepened snow decreases summertime CO2 release rates in distinct High and Low Arctic tundra ecosystems' (manuscript) My thesis ‘The influence of snow cover and cold-season temperatures on growing-season processes’ is treating the ongoing trend and future projections of increasing snow cover thickness during the Arctic winter. More snow means two things: warmer soils during winter and later melt-out in spring. Both are important changes for Arctic terrestrial ecosystems, since warmer soils will increase CO2 emissions from the soil and at the same time increase nutrients for plant growth. Later melt-out means that the plants have even less time to grow and take up C than usual, which means that the Arctic could become a net source of CO2. However, increased nutrients might counteract that effect. We increased snow depth with the help of snow fences to test how Arctic ecosystems might react in a more-snow scenario. The extra snow has increased soil temperatures during the whole, long winter, but also made the winter longer by delaying melt-out. During winter, we measured CO2 emissions from the soil, a natural process mainly done by microorganisms (ecosystem respiration). We found that more CO2 was coming from areas with increased snow, because higher temperatures stimulate microbial activity – even when the soil is frozen. Surprisingly, CO2 emissions during summer were lower in these areas after a few years of increased snow depth. That was probably because the microorganisms used up their substrates or ‘food’ during the warm winters so they didn’t have enough left for summer. This high microbial activity during winter simultaneously produced more nutrients for plants. Therefore, some species grew bigger in areas with deep snow although they had less time to grow because of the delayed melt out. This extra uptake of C could counteract the increased CO2 emissions during winter.

The papers of this thesis are not available in Munin: 1. Aarbakke, O. N. S., Bucklin, A., Halsband, C. and Norrbin, F.: 'Discovery of Pseudocalanus moultoni Frost 1989 in Northeast Atlantic waters based on mitochondrial COI sequence variation', Journal of Plankton Research (2011), vol. 33(10):1487-1495, availble at http://dx.doi.org/10.1093/plankt/fbr057 2. Aarbakke, O. N. S., Weydmann, A. and Fevolden, S-E.: 'Pseudocalanus (Copepoda: Calanoida) species distribution and relative abundance as indicators of changing sea temperature. Submitted to Journal of Plankton Research' (manuscript) 3. Aarbakke, O., Bucklin, A., Halsband, C. and Norrbin, F.: 'Comparative phylogeography and demographic history of five sibling copepod species in the North Atlantic Ocean' (manuscript) The seven species of the genus Pseudocalanus (Copepoda: Calanoida) are difficult to identify because of very small interspecific, and comparatively large intraspecific, divergence of morphologic and morphometric traits. Thus, despite the fact that Pseudocalanus spp. are among the most abundant metazoans in the world, our knowledge of them at the species level is limited. The main objective of this thesis was to lay the groundwork for future studies of Pseudocalanus spp. in the Northeast Atlantic Ocean and Euro-Arctic by determining which species are present. Through a combined DNA barcoding and traditional identification approach, this work reveals the presence of at least four species of Pseudocalanus, one of which was previously not believed to occur in the East Atlantic Ocean. Furthermore, the thesis reveals differences in environmental preferences and distributions of Pseudocalanus elongatus, P. minutus, P. moultoni and P. acuspes in the Northeast Atlantic Ocean and Euro-Arctic, and discusses implications of these differences in relation to rising sea temperatures. Finally, this work examines the phylogeography and demographic history of Pseudocalanus minutus, P. moultoni, P. elongatus, P. acuspes and P. newmani in the North Atlantic Ocean and based on these results suggests an evolutionary hypothesis for the divergence of Pseudocalanus spp. into one oceanic and one coastal clade. Hopefully, this thesis has achieved its main goal of providing a basis from which the biology and ecology of these fascinating copepod species can be further explored.

Papers 1, 2 and 3 of this thesis are not available in Munin: 1. Reiertsen, T.K., Erikstad, K.E., Anker-Nilssen, T., Barrett, R.T., Boulinier, T., Frederiksen, M., González-Solís, J., Gremillet, D., Johns, D., Moe, B., Ponchon, A., Sandvik, H., Skern-Mauritzen, M. and Yoccoz, N.G.: 'Prey density in non-breeding areas affects adult survival of black-legged kittiwakes Rissa tridactyla breeding in the southern Barents Sea' (manuscript). 2. Reiertsen, T.K., Barrett, R.T. and Erikstad, K.E.: 'Kittiwakes on the cliff edge: a demographic analysis of a steeply declining arctic kittiwake population' (manuscript). 3. Erikstad, K.E., Reiertsen, T.K., Barrett, R.T., Vikebø, F. and Sandvik, H.: 'Seabird–fish interactions: the fall and rise of a common guillemot Uria aalge population', Marine Ecology Progress Series (2013), vol.475:267-276. Available at http://dx.doi.org/10.3354/meps10084 Many seabird populations have declined dramatically over recent decades, and studying the demography and population dynamic in seabird populations is important to gaining a better understanding of causes and mechanisms lying behind such changes and to identify targets for conservation and management. Climate and prey availability are known to affect demography and population dynamics of seabirds. Climate is now changing and a warming of the ocean may lead to changed availability of prey species and will affect seabirds through their demographic traits and ultimately their abundance and distribution. The main aim of this thesis was to study the impact of climate and prey abundance on seabird demography and population dynamics, using two seabird populations; the black-legged kittiwake Rissa tridactyla and the Common guillemot Uria aalge breeding on Hornøya in NE Norway whose populations are declining in Norway. The first question addressed in the thesis was how environmental conditions affect the demography and population variability of kittiwakes and guillemots. The non-breeding distribution of the kittiwakes has recently been documented using year-round light-based geolocators, and we investigated the relationship between environmental conditions in these non-breeding areas and the adult survival of kittiwakes breeding on Hornøya. We found for the kittiwakes that Thecosomata, a group of pteropods (also called sea butterflies), in the Grand Banks/Labrador Sea area in winter and the capelin Mallotus villosus stock in Barents Sea in the pre-breeding season together explained as much as 52 % of the yearly variation in adult survival rate. Further we found that the availability of 0-group cod Gadus morhua was a much more important factor affecting demography and determining the population dynamics of the common guillemot in the Barents Sea than previously considered. The second question was which demographic trait drives the population dynamics of the steeply declining kittiwake population on Hornøya. We showed, by using demographic matrix modeling and LTRE analyses, that both the variability in breeding success (and hence the recruitment to the population) and adult survival rate contributed to the steep decline in the kittiwake population on Hornøya. The strong reduction in breeding success (and hence a decline in recruitment to the population) had, however, the highest impact. The final question was whether populations can adapt to environmental changes through micro-evolutionary processes. This study found that adult survival of the two different genetic colour morphs (bridled and non-bridled morph) of the common guillemot responded differently to yearly variability in sea surface temperatures. Colour dimorphism may, in this case serve as a genetic marker in the guillemot population, and can potentially visualize the direction of selection and micro-evolutionary processes driven by climate. The studies in this thesis contribute to the understanding of the population dynamic of two declining seabird species which are both on the Norwegian Red List, and to the directions of future studies and conservation targets of both species.

The papers of this thesis are not available in Munin: 1. Schott T., Hagen S.B., Ims R.A. & Yoccoz N.G.: 'Are population outbreaks in sub-arctic geometrids terminated by larval parasitoids?', Journal of Animal Ecology (2010), vol.79:701-708. Available at http://dx.doi.org/10.1111/j.1365-2656.2010.01673.x 2. Schott T., Kapari L., Hagen S.B., Vindstad O.P.L., Jepsen J.U. & Ims R.A.: 'Predator release from invertebrate generalists does not explain geometrid moth outbreaks at high altitudes', The Canadian Entomologist (2013), vol.145:184-192. Available at http://dx.doi.org/10.4039/tce.2012.109 3. Schott T., Ims R.A., Hagen S.B. & Yoccoz N.G.: 'Sources of variation in larval parasitism of two symmetrically outbreaking birch forest defoliators', Ecological Entomology (2012), vol. 37:471-479. Available at http://dx.doi.org/10.1111/j.1365-2311.2012.01386.x 4. Schott T., Svavarsdottir S., Hagen S.B., Kapari L., Vindstad O.P.L., Biuw M., Jepsen J.U. & Ims R.A.: 'Coping with variation in prey abundance and phenology: Comparative breeding performance of two passerines at the northern limit of their geographic range' (manuscript). The aim of this PhD-project has been to increase our knowledge of trophic interactions along altitudinal gradients in a system of sympatric, cyclically outbreaking geometrid moth species in sub-arctic, coastal birch forest in northern Norway. The project resulted in four scientific publications; two sub-studies focused on the causal effect of natural enemies on the observed complex spatio-temporal dynamics of the moth species (Papers 1 & 2). The other two sub-studies investigated the ecological effects of variable moth larvae densities as a resource for predators in the sub-Arctic ecosystem (Papers 3 & 4). In paper 1, we rejected the hypothesis that larval parasitoids might terminate moth population peaks, one of the most important hypotheses on insect population regulation. We, however, found that larval parasitoids can influence local spatial variation in moth population density. Hence, we concluded that spatial replication of sampling sites is crucial to arrive at non-biased estimates for the temporal effect of larval parasitoids on host populations. The long enduring debate on the regulation potential of parasitoids on insect populations in the scientific literature may be viewed in light of our findings. Similarly, in Paper 2, we tested if the often-observed spatial outbreak patterns of moth populations might be a result of release from invertebrate predators. Along mountain slopes, moth outbreaks in northern Fennoscandia often occur close to the tree line, and we tested if the abundance of ground dwelling invertebrate predators was inversely correlated with altitude, testing the assumption that moth outbreaks may be a result of reduced predation impact at high altitudes. As our analyses could not reveal this predicted pattern, we rejected the hypothesis. Consequently, the first two sub-studies delimitated the number of candidate factors that might shape Fennoscandian moth dynamics by reevaluating the role of larval parasitoids and ground-dwelling invertebrate predators. In paper 3, we reanalyzed the data from Paper 1 from the parasitoid's perspective by determining the dependencies of individual parasitoid species to host densities. For comparison, we also estimated the strength of effects hidden in the spatial and temporal attributes of altitude and year on parasitoid prevalence rates. We found that altitude and year, rather than host density, explained by far most of the extra-binomial variation in the parasitoid prevalence data. Hence, we concluded that unknown factors, which co-vary with altitude and year, dominate the prevalence dynamics of the larval parasitoids in our study and, consequently, act to decouple parasitoid dynamics from the dynamics of their hosts. Moth larvae as a resource for predators were also the focal issue in Paper 4, which focused on the importance of prey availability and forest phenology for breeding parameters in the two passerine birds great tit and pied flycatcher. The study was based on the phenological conflict that spring arrives in mountain slopes first at low altitude, while the bird's main food resource during the breeding season (i.e. moth larvae) often dominates forest habitat at higher altitude. Thus, we tested if passerine birds select breeding habitat according to the onset of spring or the availability of moth larvae and if the two bird species differed in this respect. We found that the phenological timing was the main driving force for both species when selecting breeding habitat lowest in the gradients where spring arrived first. However, also the positive correlations with moth larvae density appeared to be significant in our analyses. The main difference between the two species was found in the timing strategy for egg laying and size of clutches, which might make the great tit more prone to mismatch during springs of lower temperature. In addition, while nest box occupancy decreased with altitude in both species, the same relationship with respect to clutch size was only found for pied flycatchers, but not for great tits. Finally, great tits were more prone to fail during their breeding attempt and we concluded that the pied flycatcher, by laying eggs up to two weeks after great tits, appears to be the better adapted species with respect to breedig at sub-arctic conditions. This PhD-thesis strengthens the view that so-called top-down interactions in the ecological moth system in northern Fennoscandia cannot explain complex moth population dynamics, but that moth larvae can be regarded as a valuable resource for species at other trophic levels. The consistent dominance of altitude and year in all our analyses, on the other hand, suggests that explanations for the complex spatio-temporal moth population dynamics ought to be looked for in mechanisms ruled by environmentally governed processes such as phenology.

The papers of this thesis are not available in Munin: 1. B. N. Fredriksen, L. B. Hølvold, J. Bøgwald and R. A. Dalmo: 'Optimization of formulation variables to increase antigen entrapment in PLGA particles', Polymer-Plastics Technology and Engineering (2012), vol.51(14):1468-1473. Available at http://dx.doi.org/10.1080/03602559.2012.715358 2. L. B. Hølvold, B. N. Fredriksen, J. Bøgwald and R. A. Dalmo: 'Transgene and immune gene expression following intramuscular injection of Atlantic salmon (Salmo salar L.) with DNA-releasing PLGA nano- and microparticles' (manuscript) Adjuvanter benyttes for å styrke/bedre immunresponsen ved vaksinering. PLGA (poly(lactic-co-glycolic)acid) er en biologisk nedbrytbar polymer som innehar adjuvantegenskaper når den benyttes i form av partikler. Som vaksinebærere kan partikler beskytte mot nedbryting, stimulere inflammatoriske immunresponser og initiere adaptive immunresponser. Partikler er også vist å gi økt opptak i ulike celletyper. Avhandlingen tar for seg PLGA partikler som bærere/adjuvanter for DNA vaksinering, en vaksineform som har vist potensiale for bekjempelse av virussykdommer blant ulike fiskearter. PLGA partikler med plasmid ble injisert intramuskulært i atlantisk laks. Rent plasmid førte til høyest uttrykk av transgen. Bruk av partikler ga sterke inflammatoriske responser, også sammenlignet med en oljeadjuvant. Partikler med plasmid induserte også antivirale responser. Resultatene tyder på at inkapsulering i PLGA partikler kan bidra til å styrke immunresponser som kan være gunstige ved DNA vaksinering.

All but paper 5 of this thesis are not available in Munin: 1. Aars J, Marques T A, Buckland ST, Andersen M, Belikov S, Boltunov A & Wiig Ø.: 'Estimating the Barents Sea polar bear subpopulation size', Marine Mammal Science (2009), vol. 25(1):35-52. Available at http://dx.doi.org/10.1111/j.1748-7692.2008.00228.x 2. Andersen M, Derocher AE, Wiig Ø & Aars J.: 'Movements of two Svalbard polar bears recorded using geographical positioning system satellite transmitters', Polar Biology (2008), vol. 31:501-507. Available at http://dx.doi.org/10.1007/s00300-008-0428-x 3. Freitas C, Kovacs KM, Andersen M, Aars J, Sandven S, Mauritzen M, Pavlova O & Lydersen C.: 'Importance of fast ice and glacier fronts for female polar bears and their cubs during spring in Svalbard, Norway', Marine Ecology Progress Series (2012), 447:289-304. Available at http://dx.doi.org/10.3354/meps09516 4. Andersen M, Derocher AE, Wiig Ø & Aars J.: 'Polar bear (Ursus maritimus) maternity den distribution in Svalbard, Norway', Polar Biology (2012), vol.35:499-508. Available at http://dx.doi.org/10.1007/s00300-011-1094-y 6. Andersen M and Aars J.: 'Short-term behavioural responses of polar bears to disturbance by snowmobiles', Polar Biology (2008), vol.31:501-507. Available at http://dx.doi.org/10.1007/s00300-007-0376-x 7. Derocher AE, Wiig Ø & Andersen M.: 'Diet composition of polar bears in Svalbard and the western Barents Sea', Polar Biology (2002), vol.25:448-452. Available at http://dx.doi.org/10.1007/s00300-002-0364-0 8. Andersen M, Lie E, Derocher AE, Belikov SE, Bernhoft A, Boltunov AN, Garner GW, Skaare JU & Wiig Ø.: 'Geographic variation of PCB congeners in polar bears (Ursus maritimus), from Svalbard to the Chukchi Sea', Polar Biology (2001), vol. 24:231-238. Available at http://dx.doi.org/10.1007/s003000000201 9. Andersen M, Gwynn JP, Dowdall M, Lydersen C & Kovacs KM.: 'Radiocaesium in marine mammals from Svalbard, the Barents Sea and the North Greenland Sea', Science of the Total Environment (2006), vol. 363:87-94. Available at http://dx.doi.org/10.1016/j.scitotenv.2005.06.019 Polar bears in the Barents Sea population have been protected from hunting in Russia since 1956 and following the signing of the international Polar Bear Agreement in 1973 in Norway. This thesis seeks to summarise current knowledge on key population biology issues four decades after the Norwegian protection and almost six after the Russian. Further, it discusses threats that have developed in the decades following protection against human harvesting. It concludes with a discussion of the effect of multiple stressors on the population, and some thoughts on future research, monitoring and management. Polar bears in Svalbard and the Barents Sea area have been studied during the last 40 years with the aim of gaining knowledge regarding population biology and to evaluate potential sources of impact on the population from anthropogenic activity and changes to their habitat. The initial threat to polar bears in the region was unquestionably overharvest. Polar bear numbers were reduced quite drastically and hunting was clearly not sustainable. After the harvesting was stopped, the population grew in size to an estimated 2650 (1900-3600) in 2004. We believe that population recovery led to a wider distribution of maternity denning in the Svalbard Archipelago, compared to the period just after the protection of the population in 1973. However, during recent decades, the population has faced challenges from a variety of new anthropogenic impacts. The population has been exposed to a range of pollutants and an increasing level of human presence and activity within their range. Contaminants are bioaccumulated through the trophic levels in the marine food web, culminating in this top predator that consumes primarily ringed, bearded and harp seals. Females with small cubs use the land-fast sea ice for hunting, and are vulnerable to human disturbance. Changes in sea ice conditions also affect polar bears in the region, and reduced access to denning areas on the eastern islands of Svalbard is currently a concern. A decrease in spring land-fast ice close to important denning areas could negatively affect the survival of cubs. Research and monitoring provides advice to management bodies both locally and globally. Information on the presence of toxic compounds in High Arctic systems has resulted in progress in recent decades in having better control of harmful substances and in some cases international bans on their production and use. This has resulted in declining contaminant burdens in polar bears. Unfortunately, new harmful substances are finding their way to the Arctic, while others, such as radionuclides, are stored locally (within Russian Territories) in large quantities, representing potential sources of pollution. The protection of important habitats locally with restrictions on motorized traffic may help reduce negative impacts from human activity on polar bears in the region. The fate of polar bears with regard to climate change is uncertain, but significant negative effects have been documented and these impacts are expected to increase in the coming decades. Relevant research and monitoring of polar bears is essential for future management of the species. The arctic environment should be managed in such a way that the combined effects of stressors on polar bear populations are minimized.

Papers 1, 2 and 3 of this thesis are not available in Munin: 1. Øistein Harsem, Arne Eide and Knut Heen: 'Factors influencing future oil and gas prospects in the Arctic', Energy Policy (2011), vol. 39(12):8037–8045. Available at http://dx.doi.org/10.1016/j.enpol.2011.09.058 2. Øistein Harsem, Knut Heen, Joao Rodrigues and Terje Vassdal: 'Oil exploration and sea ice projections in the Arctic' (manuscript). 3. Dag Harald Claes and Øistein Harsem: 'The interdependence of European-Russian energy relations' (manuscript). The aim of this thesis is to analyze what drives Arctic oil and gas activity, and the political effects of the oil and gas industry in the high north. Based on existing political economy theories, four different articles explore these questions by using different methodological approaches. The main findings are that while the potential for a large increase in Arctic oil and gas output is significant, only certain Arctic provinces may become more attractive. An Arctic oil and gas bonanza is unlikely. Further, the Arctic energy resources are to a large extent embedded in the Russian-European energy relationship, which is best characterized by interdependence. Natural gas can be used as a political instrument under specific conditions, but future developments may increase diversification of natural gas and decrease the potential of using natural gas for political purposes.

The papers of this thesis are not available in Munin: 1. Silje Ramsvatn and Torstein Pedersen: 'Ontogenetic niche changes in haddock Melanogrammus aeglefinus reflected by stable isotope signatures, δ13C and δ15N' Marine Ecology Progress Series (2012), vol. 451:175–185, available at http://dx.doi.org/10.3354/meps09604 2. Torstein Pedersen, Silje Ramsvatn, Einar M. Nilssen, Marianne Nilsen, Lyne Morissette, Trond Ivarjord, Geir Systad, Ina Kolsum and Håvard Fause: 'Running headline: D iversity and Mass Flows Patterns in Fjord Ecosystems Species diversity affects marine ecosystem structure and mass flows' (manuscript). 3. Silje Ramsvatn, Torstein Pedersen, Marianne Nilsen and Einar M. Nilssen: 'Can stable isotopes be used to parameterize a marine food-web model in Ecopath?' (manuscript) Marine økosystemer er svært komplekse, og derfor bruker vi modeller for å kunne forenkle og systematisere kompliserte sammenhenger. Slike økologiske modeller brukes i utstrakt grad innen forskning og forvalting. Hva de ulike artene av fisk og krepsdyr spiser er svært viktig å vite for å kunne forstå hvordan artene påvirker hverandre, og dette kan vi finne ut med en liten vevsprøve for å måle stabile isotoper. Dette prosjektet undersøkte om denne metoden gav like god informasjon som analyse av mageinnholdet.

It is evident that, both due to the large water volumes they hold and that they stretches from coast to inland, it is to be expected that the northern Norwegian fjords plays an important role in a climatic scientific context. Also generally northern regions are considered as a choke point in the regional as well as the global climate. In my opinion it is therefore surprising to find thorough physical oceanographical examinations of these areas are scarce or missing. Since large data sets are compiled in the present thesis, it is my hope that it will contribute and trigger further investigations and interpretations. Due to the large spread of data in time and space, it is clear that my physical oceanographical interpretations are far from comprehensive. The periodic collection of hydrophysical data along the northern Norwegian coast (from Malangen to Porsangerfjord) started in the beginning of the 20th century and some of the oldest stations are still in operation by the “Havmiljødata” sampling programme orga nized by BFE/UIT. One of the the main aims of the current PhD project was to organize and systematize the datasets and analyze for long term trends and also aspects of shorter period variations. The complete database from 1920–2012 has been digitalized and pre sented to the general public and scholars on the website http://purl.org/hmd. The results of the research demonstrates some significant differences in the hydrophysical characteristics between the investigated fjords. There are also hydrographic variations between Inner fjords and Outer fjords. The hydrographic features of the fjords of northern Norway notably differ from the Arctic and Southern ones in many ways. The peak surface temperatures in the northern Norwegian fjords were recorded in July–August whereas the highest bottom temperatures were measured in November–December. In terms of temperature and salinity variations, Balsfjord demonstrates the highest range of temperature and salinity among the fjords, followed by Altafjord. Malangen shows a narrower distribution in both parameters whilst Porsangefjord is considerably colder (has the lowest temperatures) and also operates at the narrowest salinity range. According to the computations Altafjord has the least water exchange, followed by Malan gen and Balsfjord, whereas Porsangerfjord has the most heat and water advected into the fjord. The outer and coastal stations are affected by Atlantic Water (AW), however the in ner stations in Porsangerfjord and Altafjord and other locations are not. The overall heat content and the temperature anomalies for 5m at stations Outer Malangen (open fjord), Inner Balsfjord (closed fjord), Refsbotn (coastal station) and Outer Porsangefjord (north ern open fjord) have been calculated. They demonstrate no significant heating or cooling trends during the period of 1920–2012. Main warm periods during the winter season were 1929–1930, 1990–1992 and 2000 in all the stations ( Malangen, Balsfjorden, Refsbotn Skipsholmen and Outer Porsangerfjor den ). Also Outer Malangen and Inner Balsfjorden were warmer in 1938. Malangen and Balsfjorden experienced another warm period during 2005 followed by a constant cooling trend until 2012 whereas in Altafjorden and Porsangerfjorden the warm period was in 2007 before it started cooling. The correlation between interannual variability of temperature in Inner Balsfjorden and NAO index has been discovered. The changes of the NAO index parameters cause the subsequent temperature shift in the fjord that may occur after a considerable time.

The papers of this thesis are not available in Munin: 1. Hai P. Nguyen, Roger B. Larsen, Hong H. Hoang: '“Trash Fish” in a Small Scale Fishery: a Case Study of Nha Trang Based Trawl Fishery in Vietnam', Asian Fisheries Science (2011), vol.24(3):387-396, available at http://www.asianfisheriessociety.org/publication/abstract.php?id=82 2. Hai P. Nguyen, Roger B. Larsen: 'Effect of codend mesh size increases on the size selectivity of commercial species in a small mesh bottom trawl fishery', Journal of Applied Ichthyology (2013), vol.29(4):762-768, available at http://dx.doi.org/10.1111/jai.12177 3. Hai P. Nguyen: 'Size selectivity of alternative codends in a mixed bottom pair trawl fishery' (manuscript) 4. Hai P. Nguyen: 'Willingness of fishermen to use a bycatch reduction device (BRD) and their attitudes toward an indiscriminate fishery' (manuscript) Trawling is an important fishing method used by the inshore fishermen of Nha Trang city, Khanh Hoa province, Vietnam. The inshore trawl fishery in Nha Trang city is known to have very low selectivity, which may not be a responsible fishing practice. The trawl nets of the fishery retain large numbers of juvenile individuals, which has negative effects on the marine resources. The goal of this thesis is to provide initial suggestions to improve the selectivity of the inshore trawl fishery of Nha Trang city in order to reduce the catch of trash fish and juveniles of economically important species. To fulfil this goal, I performed the following steps: (1) studied the current status of juvenile and trash fish caught by the fishery in order to gather information on the composition and sizes of untargeted species ; (2) studied the selectivity of conventional codends with different diamond mesh sizes and with a square mesh panel; (3) aimed to determine the willingness of fishermen to use a bycatch reduction device (BRD) to reduce the catch of juvenile individuals. The status of juvenile specimens caught by the fishery was determined by: (1) direct access to trash fish samples from the trawlers of the fishery; and (2) catch notes made by the wives of fishermen who were responsible for selling the catch at landing points. The selectivity of the codends tested was studied during sea trials on chartered trawlers from the fishery where the cover codend method was used. I also interviewed the captains of the trawlers to collect information about their willingness to test future versions of a BRD. We found that trash fish accounted for reasonable part of the catch by trawlers from the fishery and played an important role in the incomes of fishermen, especially fishermen from small trawler fleets (i.e., trawlers with an engine power of less than 33 Hp). The trash fish comprised several economically important species (i.e., Cephalopoda spp., Portunidae spp., Penaeidae spp., Trichiurus lepturus, Saurida tumbil, and Epinephelus spp.). Most of the economically important species in the trash fish category measured less than 13 cm in total length (TL), except for Trichiurus lepturus where TL could reach 15 cm. This project also showed that the current mesh size of the diamond mesh at the codend had very poor selectivity performance. An increase in the mesh sizes used by the fishery to the legal mesh size would help to increase the selectivity for lizardfish, shrimp, squid, and threadfin bream. In addition, increasing the current codend mesh size to match the legal mesh size would help to reduce the catch of trash fish significantly. However, the legal mesh size is not sufficient to totally protect juveniles caught by the gears. With the current situation, the legal mesh size would be accepted to compromise between the protection of juveniles and the economic loss of fishermen. In the future, mesh sizes which is larger than current legal mesh sizes are suggested. The square mesh panel also helped to reduce the catch of trash fish and increased the selectivity for squid and lizardfish. Fishermen from the small trawler fleet refused to use the BRD to release the juvenile individuals. Fishermen from the larger trawler fleets may use the BRD provided there is no reduction in their catch of small squid. Information learnt from interviews with fishermen showed that the fish stocks exploited by the studied trawl fishery are declining sharply and it is necessary to deliver suitable measures to protect the fish stocks. Improving selectivity of the trawls used in the fishery is one of the measures.

The papers of this thesis are not available in Munin: 1. Pham Thi Thanh Thuy, Ola Flaaten and Nguyen Thi Kim Anh: 'Remuneration Systems and Economic Performance: Theory and Vietnamese Small-scale Purse Seine Fisheries' Marine Resource Economics (2013), vol.28(1):19-41. Available at http://dx.doi.org/10.5950/0738-1360-28.1.19 2. Pham Thi Thanh Thuy and Ola Flaaten: 'Middlemen: Good for Resources and Fishermen?' (manuscript) 3. Pham Thi Thanh Thuy and Ola Flaaten: 'The Backward-Bending Supply Curve in Fisheries – Revisited' (manuscript) In Vietnam, as in many other countries, the fisheries sector plays an important role in securing food, providing livelihood, and promoting economic development. Vietnamese fisheries are characterized by open access, and the number of fishing vessels has increased tremendously over the past years while the harvest growth has slowed down. This seems to indicate that the fisheries have reached the limit on the growth, and that overfishing in near-shore waters may be imminent. Overfishing has become an important issue for governments and organizations in all coastal states in recent years. Sustainability is now stated as an overriding policy objective. The concept of sustainable development makes clear that environmental, economic and social goals must be met simultaneously. Hence, in the management process, both environmental well-being and human well-being must be taken into consideration. The central topic of this thesis is how and to what extent open-access fisheries may be consistent with or opposed to the goals of sustainable development. It is obvious that open-access fisheries may contradict the goal of environmental well-being. However, it is more unclear what an open-access regime does mean for the goal of human well-being. In the literature, it is generally assumed that the economic rent will disappear because of the common property problem. This thesis questions this assumption and takes a closer look at the processes of rent creation and distribution, and how these may affect environmental sustainability. The main concerns of the three papers are the following: Paper 1 examines if rent can be generated and distributed in open-access fisheries by looking at the way fishermen formulate remuneration contracts. This issue is also discussed in paper 2, but in light of how the fishermen link with downstream markets when fish is traded internationally. Here, the effects of different market structures are highlighted. Both papers 2 and 3 deal with the linkage between the fishermen and the resources, which is an important issue from the point of view of environmental sustainability The analytical framework applied in this thesis is presented in the model of the fishery system. This model emphasizes the relationship between the human system and the environment, and the complex interplay between the various components of the fishery system (resources, fishermen, community, market, and state). The theoretical models employed vary from general economic theories applicable to many different fields (principal-agent theory, expected utility theory, theories of market structures) to theories relating particularly to fisheries (the biological overfishing model, the backward bending supply curve in open-access fisheries. Various methodological techniques are also included (propensity score matching, autoregressive model, weight and constrained least square). In all three papers, theoretical discussions and empirical findings are combined. The data is from Vietnam – mainly related to the inshore anchovy purse seine fishery. In addition, data from the offshore yellow fin tuna longline fishery is used in order to compare how different the results are between offshore and inshore fisheries. This thesis shows that rent creation and distribution can be achieved in open-access fisheries, and that the existence of market power by middlemen to some extent can limit the overfishing problem. Hence, market power is not necessarily a failure. It also turns out that economic data might be applied to measure the problem of overfishing and the health situation of the resource if biological data is lacking. A more interdisciplinary approach, including behavior models and a holistic concept of the fishery system, could allow for an improvement of today’s fisheries management. This thesis has particularly pointed out the importance of taking the fishermen’s decision-making behavior into account. This is an issue which so far has not received much attention in the literature on fisheries management.

The papers of this thesis are not available in Munin: 1. Ofstad, L.H., Pedersen, T., Angus, C.H., and Steingrund, P.: 'Age and growth of anglerfish (Lophius piscatorius) in Faroese waters', Fisheries Research (2013), vol.139:51–60. Available at http://dx.doi.org/10.1016/j.fishres.2012.05.011 2. Ofstad, L.H., Pedersen, T., and Steingrund, P.: 'Maturation, reproduction and early life history of anglerfish Lophius piscatorius in Faroese waters' (manuscript) 3. Ofstad, L.H., Steingrund, P., and Pedersen, T.: 'Seasonal offshore-onshore migration and distribution of anglerfish Lophius piscatorius in Faroese waters' (manuscript) 4. Ofstad, L.H., Steingrund, P., and Pedersen, T.: 'Feeding ecology of anglerfish Lophius piscatorius in Faroese waters' (manuscript) Anglerfish Lophius piscatorius has during the last two decades become a highly exploited resource and is now one of the five most economically important demersal fish species in the Faroese fishery. Despite this, there is at present little information available on the population status of anglerfish in Faroese waters. The main aim of this study is to increase the biological knowledge of anglerfish and to perform an assessment of the stock in order to evaluate the sustainability of the anglerfish fishery and to investigate trophic interactions between anglerfish and other commercially important demersal fish species. Anglerfish illicia, the first fin ray, were used for age reading. The age determinations were validated and concluded to be useful in an age-based assessment because the growth rates were consistent with length frequency and mark-recapture analyses. Anglerfish have a very high growth rate during their first year of life, compared with the older ages. Males mature at a lower length and younger age than females. The L50 for males and females was 58 cm and 84 cm, respectively, corresponding to an age of around four years for males and seven years for females. Observations of spawning males and females, egg-ribbons and pelagic anglerfish larvae revealed that anglerfish have a prolonged spawning season, which peaks from February to April. In addition, observations showed that there are at least two local spawning areas, one southwest of the Faroe Plateau and one in the Faroe Bank area. Large anglerfish tagged with data storage tags and conventional tags showed a seasonal offshore-onshore migration. They migrated to shallow depths in summer to feed and to deeper waters in winter, presumably to spawn. Anglerfish performed vertical movements up into the water column, and were most active in these vertical movements during the winter time and during the nights, and light intensity seemed to trigger the migrations. Anglerfish distribution was temperature dependent, since anglerfish generally preferred temperatures warmer than 4ºC, usually between 6.5–11ºC. This may explain why anglerfish were distributed deeper west of the Faroe Bank area compared to the Faroe Plateau, which is surrounded by cold water deeper than approximately 500 m. Anglerfish in Faroese waters preyed on a variety of food items. They were generally eating fish and the most important prey species in terms of numbers were Norway pout and blue whiting, whereas cod and haddock contributed most in weight. Annual consumption by anglerfish on commercially important demersal species, such as cod, haddock and saithe, was estimated to correspond to 33, 19 and 2% of the landings and to 75, 20 and 2% of biomass losses due to natural mortality, respectively. Even with this high cod consumption by anglerfish, cod cannibalism and fishing mortality still have greater impact on the cod stock dynamics than predation by anglerfish. The annual biomass loss of anglerfish due to cannibalism was estimated to nearly 15% of the annual loss due to natural mortality. Anglerfish in Faroese waters may be regarded as a separate stock because all life stages are found in the area, local spawning area, seasonal offshore-onshore migration on the Faroe Plateau, together with insignificant emigration/immigration (< 5%). An age-based stock assessment of anglerfish in Faroese waters for the period 1999–2011 showed that the stock biomass ranged between 9000 and 19000 t with a peak in 2004–2005 and the fishing mortality for age 3–8 varied between 0.2 and 0.5 year-1. The yield per recruit curve indicated that the stock was slightly growth overfished. Decreasing the fishing mortality could potentially increase the yield per recruit by around 10% leading to a 50% higher equilibrium biomass of anglerfish. No clear stock-recruitment relationship was found. On the other hand, anglerfish year-class strength was correlated with environmental variables, such as sandeel abundance. Hence, the rather short time series of anglerfish year-class strength provides little or no evidence of recruitment overfishing. It is recommended that stock assessments should be performed on an annual basis, allowing managers to react timely when there are signs of overfishing or recruitment failure due to natural or fishery causes.

The papers of this thesis are not available in Munin: 1. Ingrid Bay-Larsen: 'The conservationists’ concerns: on national administration response to integrated use and protection planning', Local Environment (2010), vol.15(4):357-371. Available at http://dx.doi.org/10.1080/13549831003677688> 2. Ingrid Bay-Larsen: 'The Premises and Promises of Trolls in Norwegian Biodiversity Preservation', Environmental Management (2012), vol. 49(5):942-953 . Available at http://dx.doi.org/10.1007/s00267-012-9837-2 3. Ingrid Bay-Larsen: 'The constitution of power in Norway’s protected areas: on shore and in the sea', Local Environment (2012), vol. 17(3):331–347. Available at http://dx.doi.org/10.1080/13549839.2012.665862 Loss of biodiversity and fragmentation of habitats are two of the world’s persistent environmental problems. This is due to the emerging complexities, uncertainties and questions of societal scale in both defining and resolving the problems. One of the challenges connected to resolving these problems pertains to the divide between the local level, which carries the costs of restrictions and regulations of use and resource extraction, and the national and global levels, where benefits are realized in terms of multiple ecosystem services (also for future generations). Another salient challenge pertains to the divide between the aggregate level of scientific expertise about the environment and the concepts and capacities of stakeholders and their resource use. These are well-known divides that typically emerge in analyses of coupled social-ecological systems. Resolving environmental problems therefore takes place in the intersection between multiple scales, actors, agencies and competencies. The local and national/international levels are likely to differ in their perceptions of environmental problems and in their perceived needs and strategies for environmental protection. Deliberative environmental governance is increasingly a focus of the international environmental agenda, and a wide range of political programs and efforts have been launched to implement and increase the efficacy and legitimacy of environmental protection. The shift from environmental governing to governance is placing more emphasis on sectors, stakeholders and knowledge across multiple scales in environmental and other policy areas. This has added the dimension of vertical divides to persistent environmental problems in addition to the horizontal ones described above. The integration of actors, agencies and competencies across both the horizontal and vertical divides influences decision-making processes and the role and position of the environmental administration. Compared with environmental governing, deliberative environmental governance places less importance on legal instruments and formal rules, new roles for civil society, as well as trust in other institutions' expert knowledge. These shifts have implications for how the environmental administration maintains power and legitimacy for their decisions and programs. This thesis asks how Norwegian environmental administration responds to such a shift from governing to governance, in particular with respect to the empowerment of local user interests and the implementation of expert knowledge in the protection of areas and biodiversity. It also addresses how the administration utilizes the potential for combining national and local ambitions and perceptions of environmental policy. The overall research question is examined in three published articles, each answering one sub-question. Findings from these articles show that: 1) there are differing responses at the regional and national administrative levels to challenges of accountability and competence resulting from the decentralization of power; 2) the environmental administration struggles to address the inherent limitations in expert knowledge applied in environmental planning, particularly with respect to the complexities of environmental problems and how to handle scientific uncertainties; and 3) the contesters of environmental protection apply various power resources to challenge the comprehension of environmental problems upon which environmental policy rests. This thesis clearly shows how the ideals embedded in deliberative environmental governance are quickly faced with political, juridical and scientific complexities in the administrative reality. A paradox evolves as diverging strategies at administrative levels, and the lack of strategies to cope with the limitations of science, may weaken their power and position. Instead of protecting their role and the mandate of environmental programs, a further marginalizing of environmental protection may be the consequence. This thesis suggests that administrative strategies and guidelines also need to be amended in order to solve persistent environmental problems. Lessons learned from this thesis include the need to harmonize strategies across administrative levels and to develop guidelines to differentiate between users that are potential allies for nature protection (as embedded in international environmental conventions), versus the users that are capable of marginalizing the environmental administration and their legitimacy and efficiency. Environmental administration at both levels also needs to develop strategies to manage the limitations of scientific expertise (complexities and uncertainties) so that these are not used strategically to contest environmental programs. Finally, this thesis shows how the environmental administration not only deals with the protection of areas and nature, but also with the protection of discourses that regulate the administrative power, the boundaries between science and policy and the comprehension of coupled social-ecological systems.

Papers 1 and 3 of this thesis are not available in Munin: 1. Vo Van Can: 'Effects of economic and non-economic factors on domestic tourism demand – A general-to-specific approach' (manuscript). 3. Vo Van Can: 'Destination loyalty as a consequence of satisfaction and switching barriers' (manuscript). This dissertation has three simultaneous purposes: (1) to investigate the effects of both economic and non-economic factors on the domestic tourist flow to Khanh Hoa province in the long term and the short term; (2) to examine how the characteristics of domestic tourists and the attributes of travel modes influence tourists’ choice of travel mode to Nha Trang, Khanh Hoa; and (3) to investigate how satisfaction and switching barriers influence destination loyalty. To achieve the first purpose, the dissertation has used the general-to-specific approach; for the second purpose, the multinomial probit model has been applied, and structural equation modelling has been used to accomplish the third purpose. For the first purpose, the findings reveal that weather variables have a significant effect on the tourism demand in both the short term and the long term. Furthermore, a positive effect from the lagged dependent variable suggests that word-of-mouth recommendation to potential tourists has good effects for the Khanh Hoa tourism industry. Also, the demand from Vietnamese tourists seems to be highly sensitive to tourism costs. Meanwhile, in the study on travel mode choice, travel time and cost, mode quality, and income are key elements in explaining the tourists’ behaviour. Tourists with a lower income tend to be more sensitive to change in the per-kilometre cost; high-income tourists are much more likely to choose a plane or train rather than a coach. Finally, in the study on tourists’ loyalty behaviour, the findings indicate that both satisfaction and switching barriers have significant effects on this behaviour. Of the multi-components of switching barriers, interpersonal relationships and switching costs have positive impacts, while lack of attractiveness of other destinations has a negative effect on destination loyalty. Furthermore, all three dimensions of tourist satisfaction and overall satisfaction have positive effects on destination loyalty.

The papers of this thesis are not available in Munin: 1. Willebrand, T., Hörnell-Willebrand, M. and Asmyhr, L.: 'Willow grouse bag size is more sensitive to variation in hunter effort than to variation in willow grouse density', Oikos (2011), vol.120, no.11:1667-1673. Available at http://dx.doi.org/10.1111/j.1600-0706.2011.19204.x 2. Asmyhr, L., Willebrand, T. and Hörnell-Willebrand, M.: 'General experience rather than of local knowledge is important for grouse hunters bag size', (forthcomming manuscript in Human Dimension of Wildlife). 3. Asmyhr, L., Willebrand, T. and Hörnell-Willebrand, M.: 'Successful adult willow grouse are exposed to increased harvest risk', Journal of Wildlife Management (2012), vol.76, no.5:940-943. Available at http://dx.doi.org/10.1002/jwmg.340 4. Asmyhr, L., Willebrand, T. and Hörnell-Willebrand, M.: 'Previous catch per unit effort (CPUE) increases rate of subsequent return and CPUE of willow grouse hunters' (manuscript). An important aspect of ecological research is to provide guidelines and tools to wildlife managers. The ecological feature of density dependence which shapes population responses to harvest are important for wildlife management, but monitoring practices and exploiter dynamics to the changing resources and management regimes should also be included. Bag records are often collected by wildlife managers as part of their management regime, in which monitoring of the harvest is crucial. However, bag records are also commonly used as an index for population abundance and reproductive success. Bag records are valuable for their accessibility at a low cost and are used by researchers to parameterize harvest models and in studying population dynamics. However, seldom are bag records utilised as an index for abundance and reproduction, properly evaluated against abundance estimates based on count data. Moreover, which factors influence hunters’ effort and success (and hence bag size), are not well explored for many game species. In this thesis I present my aims, by means of long-term monitoring records and more short-term experimental data, are (1) to evaluate bag records of willow grouse (Lagopus lagopus) as indices for willow grouse abundance and breeding success and (2) to provide advancement in understanding of the underlying mechanisms determining the harvest rates; the hunters and their behavioural responses to willow grouse and (3) to derive implications for rational management of willow grouse populations. I found the bag size and harvest rate to be more dependent on hunter effort than on willow grouse density and breeding success. The vulnerability of grouse to harvest was different for young and adult birds; young were most vulnerable. Among adults vulnerability depended on whether they had bred successfully or not; reproductively successful adults were most vulnerable. The sex ratio of the adult grouse in the harvest tended to be male-biased, in accordance with what has been found earlier, and independent of willow grouse density and breeding success. Hunter’s daily bag size was at best weakly density dependent. The number of grouse encounters, which was independent of willow grouse density, and gender of the hunter were the two strongest determinants for daily bag size. Hunters general experience in willow grouse hunting seems highly variable and of great importance for their success in bagging grouse. Still, hunters with several years of local knowledge from the hunting area were equally efficient in bagging grouse as hunters with equal experience, but from other hunting areas. Hunters considered less experienced in grouse hunting managed to increase their Catch Per Unit Effort (CPUE) in an area from one year to the next. Less than one third of the hunters return to the same area from one year to the next and the return rate was dependent of their previous CPUE, contrary to what hunters express is important for their satisfaction. In conclusion, I urge caution in relying on bag size and harvest age ratios as indexes for willow grouse abundance and breeding success. The harvest rate was mostly determined by hunter effort and implies a potential for the regulation of hunter effort to maintain sustainable harvest rates. The finding that hunters appear to be selective towards key demographic components of willow grouse populations, such as adults with high reproductive success, suggests that this aspect of hunters’ behaviour should be taken into account when deciding on sustainable harvest levels for willow grouse populations.

The papers of this thesis are not available in Munin: 1. Aubert, A.B., Svensen, C., Hessen, D., Tamelander, T.: 'CNP stoichiometry of a lipid-synthesising calanoid copepod, Calanus finmarchicus, from winter to spring bloom in a sub-Arctic sound', Journal of Marine Systems (2012), Volumes 111–112, pp.19–28. Available at http://dx.doi.org/10.1016/j.jmarsys.2012.09.004 2. Aubert, A.B., Tamelander, T.: 'Stoichiometric approach to elemental limitation of pelagic producers and grazers during the Arctic summer' (manuscript). 3. Tamelander, T., Aubert, A. B., Wexels Riser, C.: 'Export stoichiometry and contribution of copepod faecal pellets to vertical flux of particulate carbon, nitrogen and phosphorus', Marine Ecology Progress Series (2012), vol.459, pp.17-28. Available at http://dx.doi.org/10.3354/meps09733 Calanoid copepods, Calanus finmarchicus, C. glacialis and C. hyperboreus, are key species in Arctic and sub-Arctic marine ecosystems. They represent important prey for a number of species such as juvenile herring and cod because of their high but seasonally variable calorific value: they synthesize and store lipids as part of their survival strategy. Limitations in food quantity and quality can have strong impact on the copepods’ lipid synthesis and development, with implications for their predators and thus for the lipid-driven Arctic marine food web. Ecological stoichiometry, i.e. the study of the relative abundance of elements (most commonly carbon, nitrogen, and phosphorus) in biological entities, is a powerful tool to study potential limitation of aquatic organisms, particularly in primary producers and in omnivorous and herbivorous zooplankton grazers. However, studies that apply a stoichiometric approach to the high-latitude marine regions are extremely rare. The main aim of this thesis is to reveal the elemental requirements and potential limitation of high-latitude calanoid copepods by investigating their C:N:P composition in combination with information on other biochemical compounds. The impact of copepods on nutrient recycling and C:N:P export through fecal pellets is also investigated. The study was conducted in a sub-Arctic north-Norwegian fjord, and in the western part of the European Arctic Ocean, focusing on the period of intense primary production during spring. Somatic stoichiometric ratios in copepods were calculated by subtracting C in the lipid sac from the total body C. Due to the strong influence of lipid storage on total body stoichiometry, high-latitude calanoid copepods, and by implication lipid-synthesizing zooplankton in general, should be treated through a two-compartment stoichiometric approach, with lipid storage and somatic tissues separated. Both season and development stage influence the somatic elemental ratios. The somatic C:P ratios of high-latitude calanoid copepods are somewhat lower than those reported in marine copepods from other areas, suggesting that they have higher growth rates than temperate and tropical copepods. Seston C:N:P stoichiometry is used as a proxy for food quality in the calculation of copepod elemental limitation. The seston C:P and N:P ratios are among the first to be reported for the Arctic Ocean and indicate a higher relative proportion of nutrients (N and P) compared to published seston values from other marine regions. Seston C:N:P ratios varied according to bloom stage, which therefore also impacts the potential C, N or P limitation of copepods. Unless they have high assimilation efficiency for C, calanoid copepods are shown to be most likely subject to C limitation. However, variable requirements during ontogeny make specific stages, such as copepodite IV, and females more sensitive to nutrient limitation (N or P). This study is the first to present data on stoichiometric ratios including P for copepod fecal pellets in the Arctic, and shows their high potential not only to vertically export P at depth but also to provide a source of P in the upper layers during the productive season if degraded. The Arctic marine regions are experiencing the greatest climate change, and the potential future stoichiometric changes in the Arctic and how climate change affects copepod-seston interaction are discussed.

The papers of this thesis are not available in Munin: 1. Sone, I., Olsen, R.L., Dahl, R. & Heia, K.: 'Visible/near-infrared spectroscopy detects autolytic changes during storage of Atlantic salmon (Salmo salar L.)', Journal of Food Science (2011), vol.76, no.3:203-S209. Available at http://dx.doi.org/10.1111/j.1750-3841.2011.02062.x 2. Sone, I., Olsen, R.L., Sivertsen, A.H., Eilertsen, G. & Heia, K.: 'Classification of fresh Atlantic salmon (Salmo salar L.) fillets stored under different atmospheres by hyperspectral imaging', Journal of Food Engineering (2011), vol.109, no.3:482-489. Available at http://dx.doi.org/10.1016/j.jfoodeng.2011.11.001 3. Sone, I., Olsen, R.L., & Heia, K.: 'Spectral changes of Atlantic salmon (Salmo salar L.) muscle during cold storage as affected by the oxidation state of haem', Journal of Agricultural and Food Chemistry (2012), vol.60, no.38:9719–9726. Available at http://dx.doi.org/10.1021/jf302505y Spectral effects of microbial growth, packaging atmosphere and heme oxidation were investigated with fresh salmon. The presence of microorganisms and their growth have little effect on the development of spectral features in fresh salmon during storage prior to extensive spoilage. Important spectral changes occur at 606 and 636 nm in the spectra of fresh salmon during storage. Heme oxidation is the primary source of spectral changes occurring at 636 nm in fresh salmon fillets during air storage. The origin of the shoulder peak appearing at 606 nm is absorption due to water in the salmon muscle. The spectral variations at 606 nm depend on the dominant oxidation state of heme in the muscle and the change in the visibility of the water shoulder peak in the spectrum.

The papers of this thesis are not available in Munin: 1. Elverland, E. & Alm, T.: 'High resolution macrofossil analysis of Late Weichselian Arctic lacustrine sediments on Andøya, northern Norway' (manuscript) 2. Elverland, E., Bjerke, J.W. & Alm, T.: 'Is one core enough? A study of the intrasite macrofossil variability of a Late Weichselian lacustrine record on Andøya, North Norway' (manuscript) 3. Alm, T. & Elverland, E.: 'A Late Weichselian Alle alle colony on Andøya, northern Norway : a contribution to the history of an important Arctic environment' (manuscript) 4. Parducci, L., Jørgensen, T., Tollefsrud. M.M., Elverland, E., Alm, T., Fontana, S.L., Bennett, K.D., Haile, J., Matetovici, I., Suyama, Y., Edwards, M.E., Andersen, K., Rasmussen, M., Boessenkool, S., Coissac, E., Brochmann, C., Taberlet, T., Houmark-Nielsen, M., Krog-Larsen, N.,Orlando, L., Gilbert, M.T.P., Kjær, K.H., Alsos, I.G. & Willerslev: 'Glacial Survival of Boreal trees in Northern Scandinavia', Science (2012) vol.335 no.6072:1083-1086. Available at http://dx.doi.org/10.1126/science.1216043 This thesis shows that Late Weichselian vegetation on Andøya may have been more diverse than previous studies have revealed. The thorough investigation of macrofossils in four parallel cores provides more reliable evidence for interpreting changes in vegetation cover than reconstructions based on a single core. The botanical macrofossils retrieved in this study largely support previous studies at the northern tip of Andøya. Arctic plant communities, probably dominated by Papaver and several species of Poaceae and Brassicaceae characterized the area. Climatic ameliorations occurred at c. 22 000 – 20 100, 20 100 – 19 500, 19 500 – 19 200, 18 800 – 18 100, 17 500 – 16 800 and 15 100 – 14 500 cal. yr BP, and during these ameliorations, the vegetation may have been more diverse than recorded both in the pollen- and macrofossil material. Macrofossils (bones) of little auk (Alle alle), coupled with other evidence, suggest a longlasting presence of sea birds in the area. Manuring by birds made a considerable impact on the local terrestrial environment, and during the ameliorations in particular, these favorable local habitats may have supported species not found in the present-day Arctic, e.g. Urtica dioica. Sedimentary ancient DNA (sedaDNA) provides evidence that during the ameliorations, Andøya may have hosted small enclaves of boreal conifer trees. Their presence on Andøya has yet to be detected by macrofossil- or pollen analyses, but the DNA evidence provides an important contribution to the debate concerning glacial survival of boreal trees within Scandinavia.

The papers of this thesis are not available in Munin: 1. Hjelset, A.M., Sundet, J.H. and Nilssen, E.M.: 'Size at sexual maturity in the female red king crab (Paralithodes camtschaticus) in a newly settled population in the Barents Sea, Norway', Journal of Northwest Atlantic Fishery Science (2009), 41:173-182. Available at http://dx.doi.org/10.2960/J.v41.m633 2. Hjelset, A.M., Nilssen, E.M. and Sundet, J.H.: 'Reduced size composition and fecundity related to fishery and invasion history in the introduced red king crab (Paralithodes camtschaticus) in Norwegian waters', Fisheries Research (2012), 121–122:73-80. Available at http://dx.doi.org/10.1016/j.fishres.2012.01.010 3. Hjelset, A.M.: 'Red king crab (Paralithodes camtschaticus) recruitment potential in Norwegian fjords' (manuscript) The red king crab (Paralithodes camtschaticus) was deliberately introduced to the Barents Sea by Russian scientists during the 1960s, with the purpose to improve coastal fishery and thus improve the local economy. Since the red king crab was caught for the first time in the Varangerfjorden area, the stock has increased in abundance and expanded westward inhabiting coastal areas along the coast of Finnmark from the Russian border in east to the area around Hammerfest in west. Today the population of red king crab supports a valuable fishery in the Barents Sea, representing an ex-vessel value of 150 million NOK in 2011. Several challenges are related to management of the red king crab, since it is an introduced species. The Norwegian management regime today has two goals, one is to keep a long term harvest within a geographical limited area and the second is to limit further spread of the crab. Basic knowledge is needed to meet both objectives, particularly knowledge on life-history traits. The main study area in this thesis has been three large fjords in Finnmark in northern Norway. There is a historical westward spread of the crab along the Norwegian coast, and it has been present in Varangerfjorden, Tanafjorden and Laksefjorden for about 35, 25 and 15 years respectively. Crabs have been sampled annually during scientific autumn cruises from 1994 until 2011. Size at maturity is a key life-history parameter in fishery management and should be monitored routinely. For female red king crabs this means the recording of presence or absence of eggs. In paper I the size at maturity was estimated in female crabs from each of the three fjords which constitute the main study area in this thesis. No temporal differences were seen and the size at maturity varied from 108 to 111 mm carapace length in the three fjords and was found to be generally higher than in native areas. Knowledge about the species individual fecundity gives information on size and number of offspring produced. Fecundity is closely related to life-history traits such as size at maturity, life-span and egg size. In this study both spatial and temporal patterns in fecundity and a reduction in individual egg weight was found. A main finding in paper II was a reduction in the individual fecundity during a part of the study period. An average female produced about 30 % less eggs in 2007 compared to 2002. Fecundity is the life-history parameters that respond most rapidly to changes in environment or other factors, such as fishery. The abundance estimates of red king crab females have increased in our area during the study period, but in the last two years, a decrease was observed in Tanafjorden and Laksefjorden. The stock size composition and the individual fecundity parameters established in paper II has been used in paper III to obtain a reproductive potential of the crab stock for the period 1995 to 2011. The male-only fishery and introduction of a female quota has had influence on the size composition and fecundity, and thus on the potential egg production in the stock. The study also showed that an observed reduction in size of large males was correlated to a size reduction of large females with a one year lag.

Papers 1 and 4 of this thesis are not available in Munin: 1. Anna Siwertsson, Rune Knudsen and Per-Arne Amundsen: 'Temporal stability in gill raker numbers of subarctic European whitefish populations', Advances in Limnology (2012), vol. 63:229-240. Available at http://www.schweizerbart.de/papers/adv_limnology/detail/63/77932/Temporal_stability_ingill_raker_numbers_of_subarct 4. Anna Siwertsson, Rune Knudsen, Colin Adams and Per-Arne Amundsen: 'Replicated morphological divergence supports incipient ecological morph formation in European whitefish (Coregonus lavaretus)', (manuscript) How new species are formed is still a major question in evolutionary biology. In ecological speciation, natural selection drives the evolution of slight distinctions between individuals into extensive differences between species. In this thesis, early stages of an ecologically driven speciation process are addressed using ecological and morphological data from a large number of European whitefish (Coregonus lavaretus) populations in northern Fennoscandia. Northern Fennoscandia is an area scattered with lakes created during the glacial retreat (~10-12 kyr BP). Many of these lakes are inhabited by one or up to three different morphs of whitefish. As many other temperate freshwater fishes, whitefish typically diverge into pelagic and littoral resource specialists, the densely rakered (DR) and the large sparsely rakered (LSR) morph, respectively. Parallel within-lake divergence is the most probable origin of these sympatric morphs. In addition, a third whitefish morph (the small sparsely rakered; SSR), specializing on profundal resources, was recently described in a few lakes in the area. The aims of this thesis were to explore and document the phenotypic diversity of whitefish in northern Fennoscandia, and to investigate some of the factors limiting the level of diversity reached within each lake. Further, the ability of LSR whitefish morphs to utilize profundal resources was explored, as this is a probable origin of profundal specialist SSR morphs. Based on the distribution of gill raker number, a temporally stable morphological trait in the studied populations, I have documented a continuum of increasing diversity in northern Fennoscandian whitefish. This whitefish diversity comprises mono (only LSR), di- (LSR and DR), and tri-morphic (LSR, DR and SSR) populations, in addition to intermediate stages between them. Within three different lakes with dimorphic whitefish populations, some individuals of the LSR morph were specialized to exploit profundal resources. This could indicate an incipient evolution towards a profundal specialist morph (SSR) in some dimorphic lakes. Ecological opportunity (lake size and productivity) is likely a prerequisite for whitefish to diverge, although limitations related to the colonization history also were present. In conclusion, there is astonishing parallel divergence in ecology, morphology and genetics along a continuum of increasing diversity in whitefish. Thus, ecologically-based natural selection is likely driving the divergence, promoting reproductive isolation and incipient ecological speciation in northern Fennoscandian whitefish populations.

Megaprosjekter er store investeringsprosjekter med betydelig innflytelse på samfunn, miljø og økonomi. Til tross for grundige utredninger og beregninger i forkant, presterer slike prosjekter sjelden i overenstemmelse med planene: de blir gjerne både forsinket og mye dyrere enn estimatene tilsa. Hva er det som skjer i megaprosjekter som gjør at planer om suksess blir til fortellinger om fiasko? Dette spørsmålet har fått stor oppmerksomhet både i offentlige institusjoner, i media og i faglitteraturen. De skisserte løsningene for å endre på den klassiske utviklingen har ofte dreiet seg om å forbedre bruken av styringsverktøy, med sterkere krav til nøyaktighet i beregninger og gjennomsiktighet i beslutningsprosessene. Denne avhandlingen problematiserer slike resepter, og argumenterer for at fokuset på mer nøyaktighet i megaprosjekter samtidig overser hvilken innvirkning dette kravet i seg selv har på hvordan prosjektene presterer. Studien flytter på denne måten fokuset fra bruken av styringsverktøy til verktøyene selv. Hva er deres rolle i megaprosjektenes dynamikk? Avhandlingen tar for seg utbyggingen av Snøhvit LNG. Prosjektet ble gjennomført i 2002-2007, mer enn 20 år etter at funnene ble gjort i Hammerfestbassenget. Den lange tiden skyldtes at var et nytt og usikkert prosjekt, og det var derfor vanskelig å finne en utbyggingsløsning som ble ansett både som gjennomførbar og lønnsom. Til slutt lyktes arbeidet med dette og Snøhvit ble realisert. Men til tross for omfattende forberedelser sluttet prosjektet seg likevel raskt til rekken av megaprosjekter som ikke presterer som forutsatt, med både forsinkelser og store kostnadsoverskridelser. Det ble anvendt en rekke styringsredskaper for å manøvrere i prosjektets usikre og komplekse omgivelser. Disse verktøyene baserer seg på økonomiske prinsipper: Oppfatningen er at tallene vil gi nøytrale svar. Avhandlingen anvender et STS-perspektiv og studerer bruken av to slike styringsredskaper i prosjektet. Først handler det om hvordan Snøhvit ble konstruert som et lønnsomt prosjekt. Det sentrale verktøyet for å vurdere dette er nettonåverdimodellen. Deretter handler det om hvordan kostnadsoverskridelsene ble håndtert. Det er inntjent verdistyring som hjelper til med dette. Ved å bruke disse instrumentene, kunne prosjektet tilpasses kalkylenes logikk. Dette kalles økonomisering. Ved å oversette usikre momenter til tall, blir kompleks informasjon mulig å sammenligne, håndtere og ta stilling til. Uten denne forenklingen er det vanskelig å ta beslutninger. Det er imidlertid den samme forenklingen som skaper problemer i gjennomføringen fordi de ikke favner kompleksiteten i utbyggingen. Studien gir nye innspill til megaprosjektforskningen. Det er ikke bare menneskelige aktører som spiller en rolle; hvordan prosjektverktøyene virker sier også noe om sammenhengen og dynamikken i et stort prosjekt. Jo mer nøyaktig omgivelsene tallfestes, jo mer sannsynlig er det at ting vil utvikle seg annerledes. Det kan således synes som at det nødvendige valget for megaprosjekter er et kostnadsoverskridende prosjekt eller ingen prosjekt over hode. En aksept for dette innebærer at framtidige forsknings- og samfunnsspørsmål bør stilles på en måte som er mer relevant for dynamikken i megaprosjekter. Hvilken type kunnskap er nødvendig for styring og evaluering av prosjekter, og hvilken type kunnskap koster mer enn den gir fordeler?

Paper 3 of this thesis is not available in Munin: 3. Ulvevadet, B. and Hausner, V. H.: 'Incentives and regulations to reconcile conservation and development : thirty years of governance of the Sami pastoral ecosystem in Finnmark, Norway', Journal of Environmental Management(2011), 92:2794-2802. Available at http://dx.doi.org/10.1016/j.jenvman.2011.06.026 Denne avhandlingen er basert på en kvalitativ studie fra forvaltningen av reindriften i Norge. Målet med forvaltningen av reindriften i Norge er å oppnå en økologisk, økonomisk, og kulturell bærekraftig reindrift. Disse målene har blitt forsøkt oppnådd gjennom et governance system fordi myndighetene har erkjent at det å inkludere ressursbrukere i forvaltningsmessige beslutningsprosesser skaper en mer rettmessig forvaltningspolitikk. For å øke medvirkningen for ressursbrukere har norske myndigheter iverksatt to ulike forvaltningsmodeller i den samiske reindriften. På den ene siden er det innført et korporativt system hvor utvalgte reineiere representerer reindriften i de årlige forhandlingene med myndighetene. I disse forhandlingene blir det økonomiske grunnlaget for reindriften lagt gjennom ulike subsidieordninger. På den andre siden er det iverksatt et medforvaltningssystem hvor reineiere er representert i styrer på lokalt, regionalt og nasjonalt nivå. Staten har også delegert en del forvaltningsgjøremål til disse styrene. Medforvaltningssystemet utgjør fire nivåer; de to øverste nivåene består av styremedlemmer (både reineiere og andre) som er politisk valgt og dermed tar politiske beslutninger. De to lavere nivåene, på det lokale plan, har kun reineiere som styremedlemmer. Det har vært vanskelig for myndighetene å få et så stort og komplekst forvaltningssystem til å fungere slik det var planlagt og det har vært utfordringer på flere hold. Det er mange grupper som skal være med i forvaltningen, både reindriftssamer og representanter fra andre næringer som har interesser i de samme områdene som reineierne bruker. I denne prosessen har noen grupper fått mer makt enn andre og dette har satt sitt preg på forvaltningssystemet. I styrer, for eksempel, så ser man at noen grupper ender opp i mindretall og på den måten alltid ”taper” sine saker. Reindriftens næringsorganisasjon, Norges Reindriftssamers landsforbund (NRL), har fått en relativt sterk posisjon i forvaltningen av reindriftsnæringen. NRL har både monopol på forhandlingsrett med staten i det korporative systemet og i tillegg har NRL en lovfestet rett til å foreslå kandidater til medforvaltningsstyrene, som blir formelt oppnevnt av Fylkestinget, Sametinget og Landbruks- og matdepartementet (LMD). I loven står det at NRLs forlag til kandidater bør følges så langt dette er mulig. Naturlig nok så foreslår NRL alltid egne medlemmer. På den måten så er NRL (sentralt) med på å utforme subsidiesystemet for reindriftsnæringen og NRL medlemmer er med på å forme og implementere politikk gjennom medforvaltningsstyrene. For NRL medlemmer er dette positivt, for en del ikke-medlemmer så er ikke dette like populært. Subsidiesystemet har vært forhandlet frem mellom LMD og NRL i over 30 år. Målet med subsidiesystemet har vært å oppnå en økologisk, økonomisk og kulturell bærekraftig reindrift. For Finnmarks vedkommende, som sliter med et for høyt reintall, så har subsidiesystemet ment å kompensere økonomisk for en reduksjon i reintallet. Mye av reindriftspolitikken dreier seg om Finnmark, men 75% av reindriften i Norge drives også i Finnmak. I store deler av Finnmark har subsidiesystemet ikke fungert etter intensjonen. Intensjonen med subsidiesystemet er at reineierne blir belønnet for å slakte mange rein. På denne måten kan reintallet reduseres samtidig som at reineierne kompenseres for dette tapet gjennom å motta subsidier. Det motsatte har skjedd. Reineierne øker reintallet, fordi flere rein betyr høyere inntekt fra både salg av kjøtt og subsidieordningen. Grunnen til at dette skjer er at subsidiene er knyttet til inntekt fra kjøttproduksjon. Høyere inntekt fra kjøttproduksjon gir mer subsidier. Målet om høyere økonomisk inntjening (økonomisk bærekraft) søkes oppnådd gjennom subsidieordningen. Målet om en økologisk bærekraftig reindrift søkes oppnådd gjennom både subsidieordningen og lovverket. Lovverket ble revidert i 1996 og erstattet med ny lov i 2007. I dag, 5 år etter at loven trådte i kraft, er situasjonen like vanskelig. The goals for the management of reindeer husbandry in Norway are multifaceted and include ecological, economic and cultural sustainability. To achieve sustainability, a corporative system and a co-management system have been developed to establish a power-sharing agreement between the state at the central level and resource users at the local level. Together, these two systems are referred to as the governance system. This governance system is a rather comprehensive and complex multi-level system in which many different institutions participate at various administrative levels. Power is difficult to share in ways that please everyone involved. For various reasons, some institutions are granted more power than others; thus, some interests are more strongly represented in the governance system. Represented by the Ministry of Agriculture and Food (MAF), the state has, for example, shared more power with the Sami Reindeer Herders’ Association of Norway (NRL) than other institutions by giving the NRL the sole right to negotiate on behalf of Sami pastoralists. The state/MAF has also shared power with the NRL in the co-management system. Specifically, the NRL has been granted a statutory right to nominate members to the co-management boards. In practice, this power has enabled these two agents to play a central role in most efforts to achieve ecological and economic sustainability. With the recent law revision in 2007, more authority has been devolved to local boards. These boards are now responsible for creating their own rules for using pastures, allocating pasture areas, achieving a sustainable number of reindeer in each district and settling numerous other issues that are important for reindeer husbandry. However, the creation of these rules is more challenging for the two largest reindeer husbandry areas in Finnmark, as many Sami pastoralists in Finnmark struggle with an excessive number of reindeer, low weights and degraded pastures (Bråthen 1997; Tverraa, 1997; Hausner et al., 2012). The pastoralists are thus tasked with resolving these difficult issues amongst themselves. Because natural resources are affected by various shifting social and ecological conditions, adaptive management systems that address change and uncertainty are important. In practice, rapid adaptation to shifting conditions is usually difficult to achieve because the management of natural resources is firmly regulated by laws and formal rules. Although the governance system of reindeer husbandry has experienced some changes over the years, these changes have been relatively incremental. Only slight modifications have occurred in the last 20 years, perhaps because important stakeholders from the industry itself, notably the NRL, have shown little interest in making changes. The NRL has yet to propose any major changes to the reindeer husbandry incentive system, which, in practice, is the primary policy instrument for achieving the important goals of ecological and economic sustainability. In addition, the efforts of the co-management boards at the regional level – the Area Board – have been characterized by internal power struggles rather than attempts to determine optimal solutions for critical problems. The state/MAF has been well aware of these difficulties in the governance system, especially in the co-management system, and has recently proposed the abolishment of the Area Board system. I conclude that the lack of capacity in the governance system to help achieving the sustainability goals is due to the asymmetric relationship between the corporative system and the comanagement system. In practice, this has resulted in weak regulations and the use of incentive-based mechanisms as the dominant policy instruments. I therefore argue that a revision of the governance system is necessary if devolution is to work as intended.

The papers of this thesis are not available in Munin: 1. Falk-Petersen, J, Bøhn, T. and Sandlund, O.T.: 'On the numerous concepts in invasion biology', Biological Invasions (2006), 8: 1409-1424. Available at http://dx.doi.org/10.1007/s10530-005-0710-6 2. Armstrong, C.W. and Falk-Petersen, J.: Habitat-fisheries interactions : a missing link?', ICES Journal of Marine Science (2008), 65: 817-821. Available at http://dx.doi.org/10.1093/icesjms/fsn092 3. Falk-Petersen, J., Renaud, P. and Anisimova, N.: 'Establishment and ecosystem effects of the alien invasive red king crab in the Barents Sea', ICES Journal of Marine Science (2011), 68 (3): 479-488. Available at http://dx.doi.org/10.1093/icesjms/fsq192 4. Falk-Petersen, J. and Armstrong, C.: 'To have one’s cake and eat it : managing the alien invasive red king crab stock' (submitted manuscript). 5. Falk-Petersen, J.: 'Option lost or opportunity gained? Perceptions on the Barents Sea red king crab invasion', (submitted manuscript). The red king crab (Paralithodes camtschaticus) is a highly valuable resource that also represents a potential threat since it is an alien invasive species in the Barents Sea ecosystem. This thesis explores the use of different interdisciplinary frameworks to analyse how ecological, social and economic concerns could be accounted for in deciding on how the king crab should be managed in the Norwegian part of the Barents Sea. Clarification of terminology used in invasion biology and systematisation of ecological information in order to establish the ecological knowledge base for management forms the basis for further analysis. Bio-economic modelling and discourse analysis are used to explore how uncertainty and the range of services the ecosystem provides can be incorporated in management. Finally the use of ecosystem service frameworks to integrate ecosystem and social science research in ecosystem based management is discussed. The ecological review revealed numerous knowledge gaps in our understanding of how the king crab interacts with native biota. It also identified negative impacts on benthic ecosystems that provide supporting services, and on provisioning services through predation on eggs of commercial fish. Bio-economic analysis illustrated the need to identify the correct relationship between crab stock size and ecosystem damage. It also showed that optimal harvest of king crab cannot be reconciled with the Barents Sea management goal of securing the ecosystem structure. The discourse analysis showed that people recognise both positive and negative impacts of the crab on supporting, provisioning and cultural services. In addition the impacts on ecosystem services we do not have the knowledge to value today, or option values, were important in forming people’s perception on how the crab should be managed. This thesis demonstrates that while the natural sciences have a clear role to play in establishing the ecological knowledge base, uncertainty and the need to account for stakeholder concerns calls for integration of social science research in the management process. Ecosystem service frameworks can be useful tools for identification, integration and evaluation of such information.

Papers 1-4 of this thesis are not available in Munin: 1. B.N. Fredriksen, K. Sævareid, L. McAuley, M.E. Lane, J. Bøgwald and R.A. Dalmo.: 'Early immune responses in Atlantic salmon (Salmo salar L) after immunization with PLGA nanoparticles loaded with a model antigen and β-glucan', Vaccine (2011) 29(46):8338-8349. Available at http://dx.doi.org/10.1016/j.vaccine.2011.08.087 2. B.N. Fredriksen and J. Grip.: 'PLGA/PLA micro- and nanoparticle formulations serve as antigen depots and induce elevated humoral responses after immunization of Atlantic salmon (Salmo salar L)', Vaccine (2012) 30(3):656-667. Available at http://dx.doi.org/10.1016/j.vaccine.2011.10.105 3. H.M. Munang‟andu, B. N. Fredriksen, S. Mutoloki, B. Brudeseth, T.Y. Kuo, I. S. Marjara, R.A. Dalmo and Ø. Evensen.: 'Comparison of vaccine efficacy for different antigen delivery systems for infectious pancreatic necrosis virus vaccines in Atlantic salmon (Salmo salar L)' (submitted manuscript). 4. B.N. Fredriksen, L.B. Hølvold, J.Bøgwald and R.A. Dalmo.: 'Optimization of formulation variables to increase antigen entrapment in PLGA particles' (submitted manuscript). Vaccines are regarded as the safest and most cost-effective strategy to prevent infectious diseases. For some diseases, vaccine improvements are required as protection levels are still inadequate. The key to solving this challenge might lie in the development of more efficacious vaccine delivery systems and adjuvants. Poly (lactide-co-glycolide) (PLGA) is a biodegradable polymer which has an extensive safety record in biological systems and possesses immunological adjuvant properties as injectable particles. In the present work, micro- and nanoparticles of PLGA and PLA were explored as a vaccine delivery system in Atlantic salmon (Salmo salar). The overall objectives were to investigate their adjuvant abilities in provoking innate and adaptive immune responses, forming antigen depots and inducing protective immunity in a challenge test with infectious pancreatic necrosis virus (IPNV). Formulation parameters in preparation of polymeric particles were systematically optimized (paper IV) to achieve stable PLGA particle products containing co-entrapped model antigens and β-glucan (paper I and II), or virus particles of infectious pancreatic necrosis virus (IPNV) (paper III). Post immunization potency of nanoparticles (300-400 nm) was demonstrated by their ability to induce early innate responses (day 2, 4 and 8) at transcription levels equal to or higher than the oil-adjuvanted formulation (paper I). Temporal differences in expression levels of innate markers were observed, suggesting rapid systemic distribution of particles (paper I). By tracing of isotope labelled proteins, nanoparticles (˂ 1000 nm) were found to localize antigens in the head kidney while micro-sized (~ 8 µm) particles generally retained antigen at the injection site. Irrespective of size, particles made of polymers with high molecular weight (MW) generally had superior depot capabilities compared to their low MW counterparts (paper II). Adaptive immune responses to immunization were assessed by QPCR and ELISA. T cell markers were not differentially expressed at the selected early time points (paper I), but at day 60 and 75 antibody responses were found to be elevated (paper II and III). In a dose-response study, micro- but not nanoparticles were demonstrated to be equally potent compared to the oil-adjuvanted control group with regard to induction of antibody responses. Long-term antibody responses induced by particles were generally less robust and therefore declined towards the end of the experimental period (120 days), while responses induced by the oil-adjuvanted formulation progressively increased. Following immunization, antibody responses were not related to polymer qualities or the ability of particles to depot or distribute antigens. Scoring of side effects demonstrated excellent safety profiles for the particle formulations (paper II and discussed in paper V). In paper III, vaccine efficacy was tested in a cohabitation challenge with IPN. Survival rates for the nanoparticle vaccinated groups were comparable to the non-vaccinated control fish and demonstrated that their ability to induce protection against IPN was inferior to the oil-adjuvanted vaccines. Virus re-isolation from head kidney and blood during the challenge period did however demonstrate some level of protection as the nanoparticle vaccinated groups were able to delay the IPNV infection. In the presented studies, the principal adjuvant properties of PLGA particles in Atlantic salmon have been demonstrated to include their capacity to induce strong innate responses and provide antigen depots for long-term delivery of antigens. In addition, indication of particle presence in lymphoid organs was an interesting finding that could suggest a certain targeting effect to phagocytic cells. To achieve a better understanding of how PLGA particles may be used to direct immune responses in salmon, more detailed studies on particle qualities-cell interactions/responses are required.

The papers of this thesis are not available in Munin: 1. Bernd Ketelsen, Rainer Schwacke, Kirsten Krause and Karsten Fischer: 'Transcriptional activation by Cytokinin Response Factor 5 is governed by an acidic Cterminus containing two conserved domains' (manuscript) 2. Bernd Ketelsen, Stian Olsen, Kirsten Krause and Karsten Fischer: 'Cytokinin responsive factor 5 (CRF5) is involved in root development, hormonal crosstalk and sugar metabolism in Arabidopsis thaliana' (manuscript) 3. Bernd Ketelsen, Stian Olsen, Karsten Fischer and Kirsten Krause: 'Correlation between 16S/18S rDNA ratio and chloroplast copy numbers in cotyledons of Arabidopsis thaliana : use for assessment of the impact of cytokinin response factor 5 on chloroplast development' (manuscript) 4. Rainer Schwacke, Karsten Fischer, Bernd Ketelsen, Karin Krupinska and Kirsten Krause: 'Comparative survey of plastid and mitochondrial targeting properties of transcription factors in Arabidopsis and rice', Molecular Genetics and Genomics (2007) vol. 277:631-46. Available at http://dx.doi.org/10.1007/s00438-007-0214-4 Acclimation responses to environmental growth conditions in plants involve complex and fine-tuned signalling networks. Environmental signals, biotic and abiotic, are received and mediated, and lead eventually to transcriptional regulation. The plant hormones are vastly involved in these processes as intercellular mediators whereas the reception of such a hormone at the site of action involves intracellular protein signalling cascades. This thesis investigates the function of the transcription factor CRF5 of Arabidopsis thaliana which was so far known to be integrated in the course of cytokinin signalling and cotyledon development. AtCRF5 belongs to the ERF subfamily of the AP2/EREBP transcription family and is accordingly carrying an AP2 DNA binding domain which was shown to enable members of this family to bind to the cis-regulatory elements Ethylene Responsive Element (ERE) and the Drought Responsive Element (DRE). By yeast transactivation assays it was possible to show that AtCRF5 acts not only as an activator of transcription but also that the responsible transactivation domain is located among the 98 final amino acids. A detailed sequence analysis of this so far uncharacterized region and a phylogenetic analysis revealed two conserved sites which could be found throughout the plant kingdom. These sites group the CRF subfamily into four clusters of which only three can be found in Arabidopsis. Promoter interaction studies revealed that AtCRF5 is able to interact with the cis-regulatory DRE element of RD29A and to induce gene expression. The regulation of gene expression of components of the cytokinin signalling pathway could be confirmed by expression analysis of transgenic Arabidopsis lines although it could not be proven if this is a direct or indirect effect. Also the crosstalk between cytokinins and other hormones might be facilitated through AtCRF5. Phenotypic studies on transgenic Arabidopsis seedling constitutively overexpressing AtCRF5 showed pleiotropic effects like shorter roots, smaller shoots, and fewer chloroplasts per cell as well as a hexose/sucrose ratio shift towards the hexoses. Surprisingly, the shoot and the root phenotype could be reversed by adding sucrose to the growth medium pointing towards an effect of AtCRF5 on sugar metabolism. Summarized, it could be shown in this thesis that AtCRF5, as a component of the cytokinin signalling pathway, is able to connect this pathway with the regulatory action of AP2/EREBP transcription factors. AtCRF5 is involved in hormonal crosstalk and sugar metabolism in Arabidopsis thaliana.

Papers 1, 3, 4 and 5 are not available in Munin: 1. Mitschke J, Fuss J, Blum T, Hoglund A, Reski R, Kohlbacher O and Rensing SA.: 'Prediction of dual protein targeting to plant organelles', New Phytologist (2009), 183(1): 224-235. Available at http://dx.doi.org/10.1111/j.1469-8137.2009.02832.x 3. Herrmann U, Fuss J, Krupinska K and Krause K.: 'Moonlighting in plastids : translation elongation factor EF-Tu is a component of chloroplast transcriptionally active chromosomes' (manuscript I submitted to Current Genetics). 4. Fuss J, Krause K.: 'Co-Localization and interaction of ribonucleoproteins in chloroplasts of Arabidopsis thaliana' (manuscript II). 5. Fuss J, Liegmann O, Krause K and Rensing SA.: 'Can Arabidopsis thaliana read the messages from Physcomitrella patens and vice versa? : an analysis on the conservation of targeting signals' (manuscript III submitted to New Phytologist). 6. Fuss J, Krause K.: 'The DNA binding domain of a Whirly protein from Arabidopsis thaliana is engaged in protein translocation across the plastid envelope membrane' (manuscript IV). The endosymbiosis-derived organelles within a plant cell, plastids and mitochondria, have to be equipped with a certain set of proteins to be fully functional. This set of proteins is encoded by different genomes: the organellar genomes and the nuclear genomes. This setup poses some interesting challenges for the regulation of gene expression and protein transport. On the one hand, the targeting signals that transport proteins to the organelles have to be highly specific and on the other hand, the communication between the DNA containing compartments to coordinate their gene expression has to be transmitted somehow, not only from the nucleus but also from the organelles back to the nucleus. In this thesis, two prediction programs are introduced. One of them can predict dual targeted proteins to both organelles (ATP, ambiguous targeting predictor) and the other one is species- specific for Physcomitrella patens (GTP_Pp; green targeting predictor – P. patens-specific). The first predictor can help to gain a more complete picture of the proteins potentially present in the organelles. With the help of that predictor, we predicted that the amount of proteins with dual targeting signals is higher than anticipated and that we to date know only a minor part of actually dual targeted proteins. The second predictor can help to answer the question on the evolutionary consistency of targeting signals within the plant kingdom and the importance of having species-specific approaches in analyzing protein targeting. We actually observed a surprisingly big difference in composition and recognition of mitochondrial and dual-targeting protein signals, which led to the conclusion, that species-specific approaches always should be considered as the optimal option for both, in silico and in vivo experiments. The second part of this thesis focuses the mechanisms of communication between nucleus and the organelles, especially the plastid possibly mediated by dual targeting. We chose several plastid RNA-/DNA-binding proteins to analyze their sub-plastidic localization and their potential additional nuclear localization. Those candidates were AtWHY1 (Arabidopsis thaliana Whirly1), four members of the AtcpRNP (chloroplast ribonucleoprotein) family and AtEF-Tu (elongation factor thermo-unstable). The analyzed members of the AtcpRNP family reflect their described multiple functions within the plastid also in a multiple localization pattern. Furthermore, we were able to show interactions of different members of the AtcpRNP family by yeast-two-hybrid interaction assays. The localization pattern of AtEF-Tu was very similar to the one observed for the AtcpRNPs, which indicated, together with a confirmed localization within the transcriptionally active chromosome, a multiple function for AtEF-Tu. The sub-plastidic localization data suggest overlapping networks of activity for the proteins by observed co-localizations. This was also shown with respect to several marker proteins for plastid functions. For AtEF-Tu and the AtcpRNPs, we also showed experimentally that a second localization in the nucleus is possible for the mature protein, which makes them interesting candidates for a possible mediation of plastid signals to the nucleus next to AtWHY1. For AtWHY1, we were able to show an effect of the DNA binding domain on the known localization pattern which seemed to reflect an aberration in transport processes through the envelope. This offers a potential regulatory mechanism that needs to be explored in detail in the future.

Papers 1,3,4 and 5 of this thesis are not available in Munin: 1. Wold A, Darnis G, Søreide JE, Leu E, Philippe B, Fortier L, Poulin M, Kattner G, Graeve M and Falk-Petersen S: 'Life strategy and diet of Calanus glacialis during the winterspring transition in Amundsen Gulf, southeastern Beaufort Sea' Polar Biology (2011) 34:1929-1946. Available at http://dx.doi.org/10.1007/s00300-011-1062-6 3. Wold A, Leu E, Walkusz W and Falk-Petersen S: 'Lipids in copepodite stages of Calanus glacialis', Polar Biology (2007) 30:655-658. Available at http://dx.doi.org/10.1007/s00300-006-0233-3 4. Wold A, Jæger I, Hop H, Geir Wing Gabrielsen GW and Falk-Petersen S: 'Arctic seabird food chains explored by fatty acid composition and stable isotopes in Kongsfjorden, Svalbard', Polar Biology (2011) 34:1147-1155. Available at http://dx.doi.org/10.1007/s00300-011-0975-4 5. Falk-Petersen S, Haug T, Hop H, Nilssen KT and Wold A: 'Transfer of lipids from plankton to blubber of harp and hooded seals off East Greenland', Deep-Sea Research II (2009) 56:2080-2086. Available at http://dx.doi.org/10.1016/j.dsr2.2008.11.020 The Arctic marine ecosystem is characterized by strong seasonality and a short but productive period of primary production in the spring when the ice breaks up. Calanus glacialis is one of the key links between primary producers and higher trophic levels, and it is adapted to the strong seasonality by life cycle traits such as; seasonal vertical migration, diapausing, reproduction in the spring and extensive energy storage. The timing of these events will have consequences for the success of C. glacialis and also for the energy available at higher trophic levels. The aim of this study was to increase the knowledge about these key life history traits with focus on the role of lipid accumulation, seasonal vertical migration and the transfer of energy from primary producers through Calanus, to higher trophic levels, such as seabirds and seals. Lipid accumulation and seasonal vertical migration of C. glacialis were studied in the Amundsen Gulf, in the Canadian Arctic (Paper I) and in Svalbard waters (Paper II and III). The transfer of energy to higher trophic levels was studied by comparing the fatty acid composition of predators to that of their prey and by using stable isotope analysis to calculate their trophic level. We analyzed five seabird species from Kongsfjorden; Little auk (Alle alle), Brünnich’s guillemot (Uria lomvia), Black-legged kittiwake (Rissa tridactyla), Northern fulmar (Fulmarus glacialis) and Glaucous gull (Larus hyperboreus) (Paper IV) as well as harp seals (Phoca groenlandica) and hooded seals (Cystophora cristata) from the West Ice (Paper V). The timing of the ascent of C. glacialis during spring, the match of reproduction with the primary production as well as the length of the primary production period determines the development of offspring , the overwintering stage the following winter, and hence the length of the life cycle. The shorter the period of primary production, the more important is the match between reproduction and bloom. Ice algae may fuel early maturation of the females and prolong the reproductive period. In the Amundsen Gulf, C. glacialis ascended to the surface water in time to feed on the ice algae bloom, but since CIII was the main overwintering stage, the ice algae fuelled development of CIII which could develop to females that possibly could spawn during the summer. The life history trait of accumulating energy in form of lipids is an adaptation of Calanus to the large seasonal and inter-annual fluctuations in food availability, and the lipids represent an effective transfer of energy up the food chain. High levels of the Calanus markers 20:1n9 and 22:1n11 in the seabirds Little auk, Black-legged kittiwake and Northern fulmar from Kongsfjorden, Svalbard, indicate that these seabirds are part of the Calanus based food web (Paper IV). Harp seals and hooded seals (Paper V) showed considerably overlap in diet, but the fatty acid composition of harp seals originated from a diatom–Calanus based food web, while the fatty acids of hooded seals originated from a food web based on dinoflagellates and the prymnosiophyte Phaeocystis pouchetii. To conclude, the diatom–Calanus food chain is an important driver for the high latitude pelagic arctic food web. The match between timing of primary production, either as ice algae or phytoplankton bloom, and the timing of reproduction of C. glacialis is crucial for the accumulation of lipids in Calanus may have consequences for the energy available for higher trophic levels.

The papers of this thesis are not available in Munin: 1. Glorvigen, P., Bjørnstad O.N., Andreassen, H.P. and Ims, R.A.: 'Settlement in empty versus occupied habitats : an experimental study on bank voles', Population Ecology (2012), vol. 54:55-63. Available at http://dx.doi.org/10.1007/s10144-011-0295-0 2. Glorvigen, P., Gundersen, G., Andreassen, H.P. and Ims, R.A.: 'The role of colonization in patchy population dynamics of a cyclic vole species' (manuscript). 3. Glorvigen, P., Andreassen, H.P. and Ims, R.A.: 'Relative importance of habitat geometry, habitat quality, population size and environmental stochasticity on occupancy dynamics in a riparian mainland-island root vole metapopulation', (manuscript). 4. Andreassen, H.P., Glorvigen, P., Rémy, A. and Ims, R. A.: 'New views on how population-intrinsic and community-extrinsic processes interact during the vole population cycles' (manuscript) The multiannual population cycles in small rodents have triggered an intense debate between proponents of population-intrinsic and community-extrinsic factors as the cycle generating mechanisms. A well-known challenge is the apparent absence of vole individuals during the low phase of the population cycle and the fact that most studies addressing vole and lemming behaviour and demography are conducted at high to moderate population densities. Overall this thesis aim to (1) present new insight for the least studied phase(s)of the vole population cycles: specifically the role of colonisation processes in the transition from small isolated populations in the low-phase to the spatially extensively distributed populations in the peak phase of the cycle, and (2) provide an updated view on how population-intrinsic and community-extrinsic processes may interact during the vole population cycles. The first experiment documented that colonisation is a beneficial strategy in female bank voles (Myodes glareolus) compared to immigration. The second experiment demonstrated that the first born cohort of root voles (Microtus oeconomus) emigrating in early summer have large capacity for colonising spatially scattered habitat patches – a capacity that explains the fast recolonistation of empty habitat space following crashes in cyclic populations. These experimental results also gave evidence for presence of social fences around high quality habitats at low population density. Hence, in the increase phase voles seem to actively search for vacant habitats and settle in them conditional on habitat quality and population density. An observational study of a riparian mainland-island root vole metapopulation confirmed that such patchy populations with cyclic dynamics may be subject to extensive occupancy dynamics. However, owing to high capacities for dispersal and habitat tracking, voles rapidly colonised the high quality islands across the entire metapopulation landscape that previously had gone extinct due to demographic (small population size) and environmental (high water levels) stochasticity. In total, the experimental and observational studies show that spatially scattered (sub)populations that have survived the crash phase are able to provide colonists to remote habitat patches embedded in hostile matrix and thus pave the way to a new increase and peak phase. The updated view on the causes of population cycles in voles emphasises the important role of dispersal (colonisation) and sociality in all phases of the cycle. It is not claimed that intrinsic mechanisms, either alone or in interactions with extrinsic factors, are necessary for generating multiannual cycles. Rather that the interaction between extrinsic and intrinsic factors likely contribute to shape rodent population dynamics, and that different innate propensities for sociality and dispersal among different rodent species may be related to interspecific variation in topology of the cycles (i.e. cycle shape and amplitude) and degree of spatial population synchrony.

Paper 3 and 4 of this thesis are not available in Munin: 3. Seuthe L, Töpper B, Reigstad M, Thyrhaug R and Vaquer-Sunyer R: 'Microbial communities and processes in ice-covered Arctic waters of the northwestern Fram Strait (75 – 80ºN) during the vernal pre-bloom phase', Aquatic Microbial Ecology (2011) 64:253-266. Available at http://dx.doi.org/10.3354/ame01525 4. Svensen C, Seuthe L, Vasilyeva Y, Pasternak A and Hansen E: 'Zooplankton communities across Fram Strait in autumn : are small copepods and protozooplankton important? (in press in Progress in Oceanography). This thesis investigates the structure and function of planktonic food webs at two sites between Greenland and the Svalbard Archipelago, covering a coastal ecosystem influenced by Atlantic water masses (Kongsfjorden, 78º N) and a more oceanic system off the East Greenlandic shelf, influenced by the outflow of Arctic water and sea ice from the Arctic Ocean (northwest Fram Strait, 75 - 80º N). In Kongsfjorden, a seasonal study was conducted with sampling at six occasions between March and December 2006. Logistical constrains prohibited a similar extensive seasonal investigation in the ice-covered waters of northwest Fram Strait. Sampling fell into the onset (April-May 2008) and end (September 2006/2007) of the productive season. All four studies investigated the stocks of pico- to micro-sized autotrophs and heterotrophs, i.e. heterotrophic bacteria, proto- and metazooplankton. Production rates of autotrophs and heterotrophic bacteria were measured in Kongsfjorden. Ratios of heterotrophic and autotrophic biomass (H:A) and of specific bacterial and primary production (µBac:µPhyto) allowed to asses the overall structure and function of the investigated planktonic food webs and to compare them to published data from other Arctic regions. The emerging picture provides three distinct food web modes, where mode A is characterized by H:A < 1 and µBac:µPhyto > 1, new production, and large-celled phytoplankton. Mode B and C are characterized by dominance of heterotrophs (H:A > 1), more regenerated production, and small-celled phytoplankton. The difference between mode B and C is a difference in the specific production ratio, with µBac:µPhyto smaller and larger 1, respectively. According to this scheme, Arctic plankton communities appear to prevail under mode B and C most of the year. In Kongsfjorden, e.g. five of the six months sampled fell under mode B and C, with mode C being typical for light-limited winter communities. From own and literature data, it is suggested that the food web mode controlling physical factors are the amount of incident photosynthetically active radiation (PAR), ice cover, nutrient concentration, and water column stability. Due to an approximately latitudinal change in these physical settings, the relative prevalence of the three food web modes changes from the marginal seas to the central Arctic Ocean, with mode A most likely being absent from the highest latitude waters. In general, Arctic plankton communities appear to sustain on average a 3-fold higher H:A biomass ratio for a given phytoplankton stock than the world’s coastal oceans, which may mainly be due to extensive import of long-lived copepods from sub-Arctic European seas through advection. It is argued that the large stock of heterotrophs plays a crucial role in structuring Arctic plankton communities, with the potential to prohibit phytoplankton bloom formation (mode A) through extensive grazing.

Papers 1 and 2 of this thesis are not available in Munin: 1. Jensen, L.K., Carroll, JL., Pedersen, G., Hylland, K., Dahle, S. and Bakke, T.: 'A multigeneration Calanus finmarchicus culturing system for use in long-term oil exposure experiments', Journal of Experimental Marine Biology and Ecology (2006) 333: 71-78. Available at http://dx.doi.org/10.1016/j.jembe.2005.12.001 2. Jensen, L.K. and Carroll, JL.: 'Effects of the volatile petroleum component xylene on Arctic algae and zooplankton' (submitted manuscript to Marine Environmental Research). Summary Petroleum exploitation is emerging in the Arctic. In the Norwegian Arctic the southern Barents Sea is opened for development of oil and gas fields. This same area holds some of the world's largest and well managed commercial fish stocks such as Herring (Clupea harengus), Capelin (Mallotus villosus), Haddock (Melanogrammus aeglefinus), Atlantic Cod (Gadus morhua). For the interest of the fishing industry, as well as for maintaining a healthy ecosystem, petroleum related Environmental Risk Assessments (ERA) for the Barents Sea must be based on sound scientific knowledge of the special features of the Arctic ecosystem. In colder areas the physical behavior of oil is different compared to warmer areas. The lower temperature and lack of sun light during the Arctic winter slows down the natural physical weathering process of oil. In ERA in temperate areas the effects of exposure to the most volatile fraction of the oil are neglected due to short exposure time. However, in the Arctic the exposure to biota of this volatile fraction may be prolonged due to lower evaporation rates and the volatile fraction may be an important contributor to the overall adverse effect of an oil spill. Likewise, the exposure to the heavier oil fractions may be prolonged and knowledge on the long-term effects of exposure is essential. As an adaptation to the Arctic environment species found here have longer life spans, larger body sizes and higher lipid contents compared to temperate equivalent species. Thus Arctic species may accumulate contaminants over longer time and reach higher life time body levels. Meanwhile higher lipid content implies a higher affinity of lipophilic contaminants such as polycyclic aromatic hydrocarbons (PAHs). So, Arctic species may be exposed to oil compounds for a longer time because of the prolonged phycical presence and may, due to special adaptive features, be more efficient at accumulating oil contaminants. Today the knowledge of effects of oil contaminants on Arctic biota is limited, but growing. This thesis addresses a few of the research challenges in the field and provides knowledge on some effects of petroleum exposure to two copepod species; the sub-Arctic Calanus finmarchicus and the Arctic sibling species C. glacialis. First, Calanus was cultured for a generations under sub-Arctic conditions (Paper I). This work establishes that Calanus is suitable for ecological relevant long-term experiments. In an oil spill scenario, Arctic copepods and algae may be exposed to the more volatile fraction of the oil. Paper II provides data showing that the Arctic diatom species, Fragilariopsis oceanica, was the most sensitive species to exposure to the volatile hydrocarbon xylene. Further, the smaller, less lipid rich copepod species C. finmarchicus, was more sensitive compared to the true Arctic C. glacialis. Extrapolating these results to a spill situation, C. finmarchicus and C. glacialis populations would be affected by xylene exposure through both direct exposure (mortality) and indirectly through a decline in prey organisms (algae). Sub-lethal effects after long-term exposures to low concentrations of oil compounds may also be detected in C. finmarchicus and C. glacialis. Feeding was suppressed in C. finmarchicus after exposure to 7.0 µg PAHs l-1 for 11 days (Paper III). No significant effect on feeding was detected in C. glacialis exposed to 10.4 µg PAHs l-1 for 11 days. However, the hatching success of eggs laid by females exposed to 10.4 µg PAHs l-1 for two days was significantly reduced (Paper III). The long term effects of lipophilic PAHs in biota depend on the bioaccumulation capacity and internal concentrations of the various PAHs. Accumulation of lipophilic compounds is particularly important for the lipid rich Arctic species. Paper IV examines the uptake kinetics of two PAHs in C. finmarchicus and show that the accumulation of the lighter and less lipophilic compound, phenanthrene, is faster that the uptake of the heavier and more lipophilic benzo[a]pyrene (B[a]P). Equilibrium was reached for phenanthrene within 192 h exposure while no equilibrium was detected for B[a]P. The bioconcentration factor (BCF) was higher for B[a]P, reflecting the higher affinity for lipids of B[a]P compared to phenanthrene (Paper IV). So the heavier, more lipophilic PAH accumulates slower but to a higher concentration in C. finmarchicus. This thesis shows that not only acute but also long-term exposures may affect Arctic species. In addition, exposure duration may determine which compounds are accumulated by biota. Both indirect effects, such as decrease of food items, and direct effects, such as mortality, reduced feeding and reproduction failure may have implications for population development. These findings should be taken into consideration when developing ERA for the Barents Sea area.

The papers of this thesis are not available in Munin: 1. Degerlund M and Eilertsen HC: 'Main species characteristics of phytoplankton spring blooms in NE Atlantic and Arctic waters(68–80°N)', Estuaries and Coasts (2010) 33: 242-269. Available at http://dx.doi.org/10.1007/s12237-009-9167-7 2. Eilertsen HC and Degerlund M: 'Phytoplankton and light during the northern high-latitude winter', Journal of Plankton Research (2010) 32: 899-912. Available at http://dx.doi.org/10.1093/plankt/fbq017 3. Degerlund M, Huseby S, Zingone A, Sarno D and Landfald B: 'Functional diversity in cryptic species of Chaetoceros socialis Lauder (Bacillariophyceae)' (submitted paper to Journal of Plankton Research). 4. Huseby S, Degerlund M, Zingone A and Hansen, E: 'Metabolite fingerprinting and physiology of the cryptic diatom Chaetoceros socialis Lauder' (manuscript) The choice of species concept when delineating phytoplankton species is decisive for our knowledge of species diversity and distribution, but will also affect our understanding of the functioning of marine ecosystems. This is particularly true for so-called cryptic species, which may show high degree of genetic and physiological heterogeneity although being morphologically similar. A precise species delimitation procedure appreciating the value of genotypic as well as phenotypic traits is therefore important. In this thesis, species concepts and functional aspects in abundant cold-water diatoms were investigated using biogeographic (species presence and abundance), taxonomic (morphological and molecular data), and functional (experimental physiological and metabolomics data) tools. A compilation of multiannual phytoplankton species abundance data gathered during the spring bloom period from north-east Atlantic and Arctic waters (68-80oN) revealed high similarities in associations of the most abundant species. Spring associations were dominated by the diatom Chaetoceros socialis and the prymnesiophyte Phaeocystis pouchetii. A corresponding investigation of the winter period in this area could not identify associations of actively growing, autotrophic phytoplankton species. Change in species composition and abundance in time could not be assessed due to lack of consistency and regularity in sampling. A high-resolution time series of phytoplankton distribution is therefore needed in this area. A case study of the so-called cosmopolitan diatom C. socialis was performed using strains from north-east Atlantic/Arctic and from Mediterranean waters. By applying a phylo-phenetic species concept the two geographical populations were found to be pseudo-cryptic. They were genetically distinct, but only slight morphological differences were observed, in this case in resting spore morphology. Furthermore, when cultivated at 2.5, 8 and 13oC, the two groups of strains were functionally different in terms of growth rates (doublings day-1), photosynthetic efficiency (maximum quantum yield), and metabolic profiles. Our results do not support the assumed cosmopolitan distribution of C. socialis and therefore the present taxonomy of this species will need revision. A taxonomic update of the pseudo-cryptic diatom formerly known as Skeletonema costatum indicated that the species present in northern Norwegian and Barents Sea waters is identical to S. marinoi. Due to reports of high genetic, metabolic, and physiological diversity within this species, a functional approach would be valuable for understanding the ecology of S. marinoi in this area. Integrated approaches combining taxonomic and functional variables are recommended for future taxonomic work on phytoplankton species. Similarly, phytoplankton ecologists are urged to acknowledge the high degree of hidden taxonomic and functional diversity present in several phytoplankton species. Functional diversity studies are suggested to be a useful spot for integration and eventual modelling phenotypic, genotypic and ecological species data. Ultimately, such an approach should be beneficial also at the ecosystem level.

The papers of this thesis are not available in Munin: 1. Natasha A. Hynes, Clemens Furnes, Børge Nilsen Fredriksen, Tori Winther, Jarl Bøgwald, Atle N. Larsen, Roy A. Dalmo: 'Immune response of Atlantic salmon to recombinant flagellin', Vaccine (2011) vol. 29, no 44, pp 7678–7687. Available at http://dx.doi.org/10.1016/j.vaccine.2011.07.138 2. Natasha A. Hynes, Tiago S. Hori, Roy A. Dalmo and Matthew L. Rise: 'Functional genomics analysis of the immunological responses of Atlantic salmon (Salmo salar) spleen to injection with recombinant flagellin from Vibrio anguillarum' (manuscript) Flagellin is the principal structural protein in the locomotive organ flagellum present on flagellated bacteria, and is known to be an important evolutionarily conserved PAMP. Flagellin has been shown to bind to the PRR TLR5 which induces innate immune system responses and signaling pathways. In mammals, much focus has been placed on this protein in vaccine studies for its possible function as an adjuvant. Flagellin (FlaD from Vibrio anguillarum) was recombinantly produced in two forms, full-length (FDL) and a truncated form (FDS) with portions of the N- and C-termini removed to prevent polymerization. Both forms of flagellin were tested alone and in combination with an antigen in a dose response study to determine the most effective dosage to produce a strong immune response. A polyclonal antibody for FDS was produced and showed good specificity in immunoblot testing. Cell culture was used to compare the NF-κB response after stimulation with FDL versus FDS and resulted in a significantly larger response in cells stimulated with FDL. QPCR mRNA gene expression results showed a strong innate immune response with a number of genes known to be induced downstream by the TLR signaling pathway up- regulated including important TLR5S and inflammatory cytokines and chemokines (TNFα, IL-6, IL-8, IL-1β). Microarray analysis of salmon injected with flagellin caused reproducible gene up-regulation of inflammatory cytokines, chemokines and receptors (IL-8, TNFRSF11B, IL-1R), antimicrobial peptides (hepcidin, cathelicidin), immune genes (C/EBP, thioredoxin, C-type lectin receptor B), complement genes (complement component C7 and C7-1), peptidases (MMP-9) and genes involved in the Ras/MAPK pathway (Ras-related proteins). Down-regulation of interesting immune genes also occurred including myxovirus resistance 1, clusterin-1 and LPS-induced TNF-α homolog. Some genes exhibited early, delayed or extended response over the sampling time and flagellin seemed to be the key component to eliciting a response. Flagellin was shown to induce innate immune responses in Atlantic salmon.

The papers of this thesis are not available in Munin: 1. Elina Halttunen, Audun H. Rikardsen, Jan G. Davidsen, Eva B. Thorstad and J. Brian Dempson: 'Survival, migration speed and swimming depth of Atlantic Salmon Kelts during sea entry and fjord migration', Reviews: Methods and Technologies in Fish Biology and Fisheries (2009) Volume 9, Part I, Part 1, 35-49. Available at http://dx.doi.org/10.1007/978-1-4020-9640-2_3 2. Elina Halttunena, Audun H. Rikardsen, Eva B. Thorstad, Tor F. Næsje, Jenny L.A. Jensen and Øystein Aas: 'Impact of catch-and-release practices on behavior and mortality of Atlantic salmon (Salmo salar L.) kelts', Fisheries Research (2010) vol 105, 141–147. Available at http://dx.doi.org/10.1016/j.fishres.2010.03.017 3. Elina Halttunen, Jenny Lovisa Alexandra Jensen, Tor Fredrik Næsje, Jan Grimsrud Davidsen, Eva Bonsak Thorstad, Cedar Marget Chittenden, Sandra Hamel, Raul Primicerio and Audun Håvard Rikardsen: 'State-dependent migratory behaviour of post-spawned Atlantic salmon (Salmo salar)' (unpublished paper). 4. Elina Halttunen, Audun Håvard Rikardsen, Ola Ugedal, Eva Bonsak Thorstad, Jenny Lovisa Alexandra Jensen and Tor Fredrik Næsje: 'The significance of repeat spawning for the production of Atlantic salmon (Salmo salar)' (unpublished paper). Summary Atlantic salmon (Salmo salar L.) repeat-spawners might play an important role in population persistence by increasing both total recruitment and long-term stability of a population. This is because repeat spawners return at larger sizes and with greater fecundity than first time spawners, and the large majority of repeat-spawners are females. Nevertheless, little knowledge exists of this potentially significant population sub-unit. We conducted a four-year field study in the River Alta, Northern Norway, using acoustic and radio telemetry combined with scale sample analysis and mark recapture, with the aim of 1) assessing the survival and migration patterns after spawning, 2) assessing the post and repeat-spawner mortality in the fisheries, and 3) quantifying the returning rate of post spawning females. These results were further used in combination with historical catch-records from both sea and in-river fisheries to 4) model the repeat spawning rate in the female population and the significance of repeat spawning for egg production during fluctuating maiden runs. We found that 1) survival after spawning and early marine migration was high (63-80% and 96%, respectively), and that the post-spawners (kelts) in best condition waited longest in the river before migrating. 2) The fisheries mortality was low (4%) when exiting the river and fjord, and moderate (11 %) when re-entering the fjord and river upon return. 3) Female survival until spawning was on average 32 %, and 4) according to model simulations, on average 20% of the female population consisted of repeat-spawners, which might have an important stabilizing function reducing fluctuations in a population by contributing an average of 27% (2%-59%) of all the eggs spawned in the river. We conclude that the post-spawners have high survival both in the river and the sea despite their weakened condition after spawning and overwintering in the river, and that repeat-spawners might contribute significantly to Atlantic salmon production. Their numeric and genetic contribution might be especially important during declining and fluctuating populations as a buffer against poor maiden recruitment years. Therefore post- and repeat-spawners are worthy of special conservational attention and management measures.

This dissertation is about the changing relationship between science and society and the attempt to organize knowledge production and advisory processes in new ways: How can the relations between science and society be organized in a way that ensures sound advice as well as democratic ideals such as transparency and inclusion? Is it possible to relax the boundary between science and society without making it too blurry? How can arenas be built where scientists and stakeholders can cooperate effectively in knowledge production? How can lay people and their knowledge be included in advisory processes? To include stakeholders and at the same provide sound scientific advice, new solutions that require more openness in scientific processes are called for. In order to explore such solutions, the dissertation turn to the fisheries sector, where the top-down, traditional divide between science and other forms of knowledge is obvious: scientists are the experts with relevant and reliable knowledge for fisheries management. In recent decades, however, fisher stakeholders are identified as having valuable knowledge based on their experiences. Some also argue that fishers are experience based experts. But how to include them in a meaningful way, and how they are experts, are issues that are still open for debate. In Norway, the Institute of Marine Research runs a project called the Norwegian Reference Fleet, where fishers are invited to participate in knowledge production for advice. The Reference Fleet project provides a relevant framework to investigate empirically how to solve dilemmas related to the organization of knowledge production and advisory processes in more open, yet effective ways. The thesis makes a detailed account of how fishers are included by describing the knowledge chains in which they participate, and what happens to the knowledge as it is made to circulate and used for advice in fisheries management issues. Based on ethnographic methods, the Reference Fleets’ knowledge production has been followed wherever it has taken place: in laboratories and at sea; and in formal and informal settings. Theoretically, this thesis is supported by Science and Technology Studies, and Actor Network Theory in particular. The thesis aims to contribute to the on-going discourses on the 'lay expert' and how to open up science for stakeholders. It demonstrates that it is possible to include stakeholders in knowledge production for advice, and that lay people can become experts. However, it also argues that this hinges on participation at appropriate stages in the process, and that lay people, like scientists, needs access to both laboratories and authorization mechanisms in order to be included or be part of the expertise.

The papers of this thesis are not available in Munin. 1. Tandberg, A.H.S. and Vader, W.: 'A redescription of Metopa species (Amphipoda, Stenothoidae) based on the type material. 1. Zoological Museum, Copenhagen (ZMUC)', Zootaxa (2009) 2093:1-36. Available at http://www.mapress.com/zootaxa/ 2. Tandberg, A.H.S.: 'A redescription of Metopa species (Amphipoda, Stenothoidae) based on the type material. 2. The United States National Museum of Natural History (NMNH)', Zootaxa (2009) 2309:43-68. Available at http://www.mapress.com/zootaxa/ 3. Tandberg, A.H.S.: 'A redescription of Metopa species (Amphipoda, Stenothoidae) based on the type material. 2. The Natural History Museum Oslo (NHM)', Zootaxa (2010) 2465:1-94. Available at http://www.mapress.com/zootaxa/ 4. Tandberg, A.H.S., Schander, C. and Pleijel, F.: 'First record of the association between the amphipod Metopa alderii and the bivalve Musculus', Marine Biodiversity Records (2010) 3, e5. Available at http://dx.doi.org/10.1017/S1755267209991102 5. Tandberg, A.H.S., Vader, W. and Berge, J.: 'Metopa glacialis : an analysis of its ecology and its association with Musculus discors', Polar Biology (2010) 33:1407-1418. Available at http://dx.doi.org/10.1007/s00300-010-0833-9 6. Tandberg, A.H.S. and Vader, W.: 'Phylogenetic analysis of the stenothoid genera Metopa and Stenula (Crustacea, Amphipoda, Stenothoidae)' (manuscript) The Amphipod-genus Metopa is a relatively large genus, with currently 55 accepted species. This PhD thesis consists of redescriptions of 23 of these species based on typespecimens or specimens collected and identified by the original authors; this has been done to create consistent descriptions that could form a basis for a phylogenetic analysis of Metopa. For some species Scanning Electron Microscopy has been used to look for characters that have been overlooked by the original authors. The redescriptions are presented in 3 papers (Papers 1-3), with a phylogenetic analysis of 37 Metopa species (mostly based on personal examinations, but 6 based on literature-information) together with 6 Stenula species (partly based on literature, but most from examinations performed by Dr. Traudl Krapp-Schickel) presented in a manuscript (Paper 6). The result of the phylogenetic analysis is two clear clades (A and B), both including species from both original genera. This result could suggest a thorough change in stenothoid systematics, with the possible synonymization of Stenula into the older Metopa and erection of one or more new genera to include the species of clade A, but we have argued against doing this before further investigations have been undertaken. The stability of the resulting phylogeny is discussed, and in our opinion further studies are necessary to strengthen the conclusions reached here, before we formally make the necessarily very extensive nomenclatorial changes. However, the indications that Metopa and Stenula are paraphyletic are strong. Two papers (Papers 4-5) examine the ecology of Metopa species associated with mollusks. Paper 4 is a short note about findings of Metopa alderii inside the mussel Musculus spp., whereas Paper 5 is a more thorough discussion on the possible life history strategies for Metopa glacialis in its well known association with the mussel Musculus discors based on data collected by Svalbard. We show the presence of more than one broods in several examined mussels, and from the brood sizes and cohort compositions we suggest two possible scenarios: 3 or more seasons of single broods and parental care over more than 1 year after hatching, or multiple broods in one season and extended parental care. The introductory synthesis to the papers gives a short discussion of the papers, in addition to a historical introduction to both taxonomy and the amphipod-genus Metopa. A complete list of taxa that at some time have been included in Metopa is presented, this includes synonymies and references to the auctors and different authors who have moved the different taxa. There is also a table with biogeographical information about the genus, and the ecology of Metopa is discussed. A short discussion on molecular methods and the possible ways to ensure more data for a better phylogenetic analysis is also included.