The genetics of behaviour and other adaptive traits in nine-spined sticklebacks (Pungitius pungitius)

One of the main goals in current evolutionary biology research is to identify genes behind adaptive phenotypic variations. The advances in genomic technologies have made it possible to identify genetic loci behind these variations, also concerning non-model species. This thesis investigates the genetics of the behaviour and other adaptive traits of the nine-spined stickleback (Pungitius pungitius) through the application of different genetic approaches. Fennoscandian nine-spined stickleback populations express large phenotypical differences especially in behaviour, life –history traits and morphology. However the underlying genetic bases for these phenotypical differences have not been studied in detail. The results of the project will lay the foundation for further genetics studies and provide valuable information for our understanding of the genetics of the adaptive divergence of the nine-spined stickleback. A candidate gene approach was used to develop microsatellite markers situating close to candidate genes for behaviour in the nine-spined stickleback. Altogether 13 markers were developed and these markers were used in the subsequent studies with the anonymous random markers and physiologically important gene markers which are already currently available for nine-spined sticklebacks. It was shown that heterozygosity correlated with behaviour in one of the marine nine-spined stickleback populations but with contrasting effects: correlations with behaviour were negative when using physiological gene markers and positive with random markers. No correlation was found between behavioural markers and behaviour. From the physiological gene markers, a strong correlation was found between osmoregulation-related gene markers and behaviour. These results indicate that both local (physiological) and general (random) effects are important in the shaping of behaviour and that heterozygosity– behaviour correlations are population dependent. In this thesis a second linkage map for nine-spined sticklebacks was constructed. Compared to the earlier nine-spined stickleback linkage map, genomic rearrangements were observed between autosomal (LG7) and sex-determing (LG12) linkage groups. This newly constructed map was used in QTL mapping studies in order to locate genomic regions associated with pelvic structures, behaviour and body size/growth. One major QTL was found for pelvic structures and Pitx1 gene was related to these traits as was predicted from three-spined stickleback studies, but this was in contrast to earlier nine-spined stickleback study. The QTL studies also revealed that behaviour and body size/growth were genetically more complex by having more QTL than pelvic traits. However, in many cases, pelvic structure, body size/growth and behaviour were linked to similar map locations indicating possible pleiotropic effects of genes locating in these QTL regions. Many of the gene related markers resided in the QTL area. In the future, studying these possible candidate genes in depth might reveal the underlying mechanism behind the measured traits.

Lipase and omega-transaminase catalysis in preparation of alcohol and amine enantiomers

Enantiopure intermediates are of high value in drug synthesis. Biocatalysis alone or combined with chemical synthesis provides powerful tools to access enantiopure compounds. In biocatalysis, chemo-, regio- and enantioselectivity of enzymes are combined with their inherent environmentally benign nature. Enzymes can be applied in versatile chemical reactions with non-natural substrates under synthesis conditions. Immobilization of an enzyme is a crucial part of an efficient biocatalytic synthesis method. Successful immobilization enhances the catalytic performance of an enzyme and enables its reuse in successive reactions. This thesis demonstrates the feasibility of biocatalysis in the preparation of enantiopure secondary alcohols and primary amines. Viability and synthetic usability of the studied biocatalytic methods have been addressed throughout this thesis. Candida antarctica lipase B (CAL-B) catalyzed enantioselective O-acylation of racemic secondary alcohols was successfully incorporated with in situ racemization in the dynamic kinetic resolution, affording the (R)-esters in high yields and enantiopurities. Side reactions causing decrease in yield and enantiopurity were suppressed. CAL-B was also utilized in the solvent-free kinetic resolution of racemic primary amines. This method produced the enantiomers as (R)-amides and (S)-amines under ambient conditions. An in-house sol-gel entrapment increased the reusability of CAL-B. Arthrobacter sp. omega-transaminase was entrapped in sol-gel matrices to obtain a reusable catalyst for the preparation enantiopure primary amines in an aqueous medium. The obtained heterogeneous omega-transaminase catalyst enabled the enantiomeric enrichment of the racemic amines to their (S)-enantiomers. The synthetic usability of the sol-gel catalyst was demonstrated in five successive preparative kinetic resolutions.

The role of nuclear and other conventional power plants in the flexible energy system

This thesis reviews the role of nuclear and conventional power plants in the future energy system. The review is done by utilizing freely accesible publications in addition to generating load duration and ramping curves for Nordic energy system. As the aim of the future energy system is to reduce GHG-emissions and avoid further global warming, the need for flexible power generation increases with the increased share of intermittent renewables. The goal of this thesis is to offer extensive understanding of possibilities and restrictions that nuclear power and conventional power plants have regarding flexible and sustainable generation. As a conclusion, nuclear power is the only technology that is able to provide large scale GHG-free power output variations with good ramping values. Most of the currently operating plants are able to take part in load following as the requirement to do so is already required to be included in the plant design. Load duration and ramping curves produced prove that nuclear power is able to cover most of the annual generation variation and ramping needs in the Nordic energy system. From the conventional power generation methods, only biomass combustion can be considered GHG-free because biomass is considered carbon neutral. CFB combusted biomass has good load follow capabilities in good ramping and turndown ratios. All the other conventional power generation technologies generate GHG-emissions and therefore the use of these technologies should be reduced.

Electrical and dielectric properties of uncoated and coated wood-free paper for electrophotography

The print substrate influences the print result in dry toner electrophotography, which is a widely used digital printing method. The influence of the substrate can be seen more easily in color printing, as that is a more complex process compared to monochrome printing. However, the print quality is also affected by the print substrate in grayscale printing. It is thus in the interests of both substrate producers and printing equipment manufacturers to understand the substrate properties that influence the quality of printed images in more detail. In dry toner electrophotography, the image is printed by transferring charged toner particles to the print substrate in the toner transfer nip, utilizing an electric field, in addition to the forces linked to the contact between toner particles and substrate in the nip. The toner transfer and the resulting image quality are thus influenced by the surface texture and the electrical and dielectric properties of the print substrate. In the investigation of the electrical and dielectric properties of the papers and the effects of substrate roughness, in addition to commercial papers, controlled sample sets were made on pilot paper machines and coating machines to exclude uncontrolled variables from the experiments. The electrical and dielectric properties of the papers investigated were electrical resistivity and conductivity, charge acceptance, charge decay, and the dielectric permittivity and losses at different frequencies, including the effect of temperature. The objective was to gain an understanding of how the electrical and dielectric properties are affected by normal variables in papermaking, including basis weight, material density, filler content, ion and moisture contents, and coating. In addition, the dependency of substrate resistivity on the electric field applied was investigated. Local discharging did not inhibit transfer with the paper roughness levels that are normal in electrophotographic color printing. The potential decay of paper revealed that the charge decay cannot be accurately described with a single exponential function, since in charge decay there are overlapping mechanisms of conduction and depolarization of paper. The resistivity of the paper depends on the NaCl content and exponentially on moisture content although it is also strongly dependent on the electric field applied. This dependency is influenced by the thickness, density, and filler contents of the paper. Furthermore, the Poole-Frenkel model can be applied to the resistivity of uncoated paper. The real part of the dielectric constant ε’ increases with NaCl content and relative humidity, but when these materials cannot polarize freely, the increase cannot be explained by summing the effects of their dielectric constants. Dependencies between the dielectric constant and dielectric loss factor and NaCl content, temperature, and frequency show that in the presence of a sufficient amount of moisture and NaCl, new structures with a relaxation time of the order of 10-3 s are formed in paper. The ε’ of coated papers is influenced by the addition of pigments and other coating additives with polarizable groups and due to the increase in density. The charging potential decreases and the electrical conductivity, potential decay rate, and dielectric constant of paper increase with increasing temperature. The dependencies are exponential and the temperature dependencies and their activation energies are altered by the ion content. The results have been utilized in manufacturing substrates for electrophotographic color printing.

Nordic alcohol policy in Europe: the adaptation of Finland’s, Sweden’s and Norway’s alcohol policies to a new policy framework, 1994-2013

The purpose of this thesis is to study how and to which extent Finland, Sweden and Norway have adapted their alcohol policies to the framework imposed to them by the EU and the European Economic Area (EEA) since the mid-1990s. This is done by studying the underlying mechanisms that have influenced the formation of alcohol policy in the Nordic countries in that period. As a part of this analysis main differences in alcohol policies and alcohol consumption between the three countries are assessed and the phenomenon of cross-border trade with alcohol is discussed. The study examines also the development of Finnish, Norwegian and Swedish alcohol policies between 1994 and 2012 and compares the Nordic alcohol policies with other alcohol policies in Europe as the situation was in 2012. The time frame of the study spans from the mid-1990s to the end of 2013 and is divided into three phases. Studying the role of the Europeanisation process on the formation of alcohol policies has a key role in the analysis. Besides alcohol policies, the analyses comprise the development of alcohol consumption and cross-border trade with alcohol. In addition, a quantitative scale constructed to measure the strictness of alcohol policies is utilised in the analyses. The results from the scale are used to substantiate the qualitative analysis and to test whether the stereotypical view of a strict Nordic alcohol policy is still true. The results from the study clearly corroborate earlier findings on the significance of Europeanisation and the Single Market for the development of alcohol policies in the Nordic countries. Free movement of goods and unhindered competition have challenged the principle of disinterest and enabled private profit seeking in alcohol trade. The Single Market has also contributed to the increase in availability of alcohol and made it more difficult for the Nordic EU member states to maintain restrictive alcohol policies. All in all, alcohol policies in the Nordic countries are more liberal in 2013 than they were in 1994. Norway, being outside the EU has, however, managed to maintain a stricter alcohol policy than Finland and Sweden. Norway has also been spared from several EU directives that have affected Finland and Sweden, the most remarkable being the abolishment of the travellers’ import quotas for alcohol within the EU. Due to its position as a non-EU country Norway has been able to maintain high alcohol taxes without being subjected to a ”race to the bottom” regarding alcohol taxes the same way as Finland and Sweden. Finland distinguishes as the country that has liberalised its alcohol policy most during the study period. The changes in alcohol policies were not only induced by Europeanisation and the Single Market, but also by autonomous decision-making and political processes in the individual countries. Furthermore, the study shows that alcohol policy measures are implemented more widely in Europe than before and that there is a slow process of convergence going on regarding alcohol policy in Europe. Despite this, alcohol policies in the Nordic countries are still by far the strictest in all of Europe. From a Europeanisation perspective, the Nordic countries were clearly on the receiving end during the first two study phases (1994–2007), having more to adjust to rules from the EU and the Single Market than having success in uploading and shaping alcohol policy on the European and international field. During the third and final study phase (2008–2013), however, the Nordic countries have increasingly succeeded in contributing to shape the alcohol policy arena in the EU and also more widely through the WHOs global alcohol strategy. The restrictive Nordic policy tradition on which the current alcohol policies in Finland, Sweden and Norway were built on has still quite a solid evidence base. Although the basis of the restrictive alcohol policy has crumbled somewhat during the past twenty years and the policies have become less effective, nothing prevents it from being the base for alcohol policy in the Nordic countries even in the long term. In the future, all that is needed for an effective and successful alcohol policy is a solid evidence base, enough political will and support from the general public.

Towards sustainable transport: A comparison of demographic and behavioural characteristics of Finnish and international car sharing users

SStrong evidence suggests that the climate is changing and that these changes are largely caused by human activities. A consensus exists among researchers that human activity is causing global warming and that actions to mitigate global warming need to be taken swiftly. The transportation sector, which relies heavily on fossil fuel burning and primarily oil, is one of the big contributors to air pollution problems at local, regional and global levels. It is the fastest growing source of greenhouse gas emissions and is estimated to be responsible for nearly a quarter of global energyrelated carbon dioxide emissions. Car sharing is a mobility solution encouraging its users to decrease private car usage in favour of communal transit and environmental goals. The idea of car sharing originates from the aspiration to decrease personal car ownership and to reduce vehicle distance travelled. This thesis seeks to complement the understanding of Finnish car sharing users and their usage through better categorization. Through better categorization and segmentation of Finnish car sharing users the thesis seeks to provide information for improved marketing insight. Research is done on the demographic and behavioural characteristics of Finnish car sharing users and they are compared with international findings about the characteristics of International car sharing users. The main research problem is Are Finnish car sharing users similar to international ones? A theoretical research framework on the determinants of individual car sharing usage is built based on international research about demographic and behaviouristic characteristics. After this a quantitative survey is performed to the customers of a Finnish car sharing organization. The data analysed in the thesis consist out of 532 answers received from the car sharing organizations customers. The data is analysed with descriptive and other exploratory methods, which create an understanding of Finnish car sharing users. At the end of the analysis the demographic and behavioural characteristics of Finnish car sharing users are compared with international ones. The research findings of the thesis indicate that the demographic and behavioural characteristics of Finnish car sharing usage largely follow those of their international counterparts. Thanks to the thesis results the car sharing organization is able to better target their customers through improved marketing insight.

Towards sustainable transport: A comparison of demographic and behavioural characteristics of Finnish and international car sharing users

Strong evidence suggests that the climate is changing and that these changes are largely caused by human activities. A consensus exists among researchers that human activity is causing global warming and that actions to mitigate global warming need to be taken swiftly. The transportation sector, which relies heavily on fossil fuel burning and primarily oil, is one of the big contributors to air pollution problems at local, regional and global levels. It is the fastest growing source of greenhouse gas emissions and is estimated to be responsible for nearly a quarter of global energyrelated carbon dioxide emissions. Car sharing is a mobility solution encouraging its users to decrease private car usage in favour of communal transit and environmental goals. The idea of car sharing originates from the aspiration to decrease personal car ownership and to reduce vehicle distance travelled. This thesis seeks to complement the understanding of Finnish car sharing users and their usage through better categorization. Through better categorization and segmentation of Finnish car sharing users the thesis seeks to provide information for improved marketing insight. Research is done on the demographic and behavioural characteristics of Finnish car sharing users and they are compared with international findings about the characteristics of International car sharing users. The main research problem is Are Finnish car sharing users similar to international ones? A theoretical research framework on the determinants of individual car sharing usage is built based on international research about demographic and behaviouristic characteristics. After this a quantitative survey is performed to the customers of a Finnish car sharing organization. The data analysed in the thesis consist out of 532 answers received from the car sharing organizations customers. The data is analysed with descriptive and other exploratory methods, which create an understanding of Finnish car sharing users. At the end of the analysis the demographic and behavioural characteristics of Finnish car sharing users are compared with international ones. The research findings of the thesis indicate that the demographic and behavioural characteristics of Finnish car sharing usage largely follow those of their international counterparts. Thanks to the thesis results the car sharing organization is able to better target their customers through improved marketing insight.