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  • Siikamäki, Heli (Helsingin yliopisto, 2015)
    Study I analyzed Finnish travellers health problems abroad during 2010 2012. Information was drawn from a database kept by an assistance organization of insurance companies covering 95% of Finnish cases requiring aid abroad. The study included 50 710 cases. These data were compared to numbers of Finnish travellers from the Official Statistics of Finland to calculate incidences of illness and injury at various destinations. The most common diagnostic categories proved to be infections (60%) and injuries (14%); the most frequent diagnoses were acute gastroenteritis (23%) and respiratory infections (21%). Incidence was high in Africa, southern Europe plus the eastern Mediterranean, and Asia. Pre-travel counselling appears advisable also for visitors to southern Europe. Means for preventing gastrointestinal and respiratory infections are needed. Study II explored the final diagnoses of returning travellers with fever. This retrospective investigation comprised 462 febrile adults returning from malaria-endemic areas admitted to the Helsinki University Central Hospital emergency room during 2005 2009. The most common diagnostic categories were acute diarrhoeal disease (27%), systemic febrile illness (15%), and respiratory illness (15%). One traveller in four had a potentially life-threatening illness; septicemia proved as common as malaria (5% vs. 4%); one in ten had more than one diagnosis. The results suggest that the diagnostic protocol in tertiary hospital should in addition to malaria smears comprise blood cultures, influenza rapid test, and HIV test. Study III analyzed surveillance data on malaria cases reported to the National Infectious Disease Register 1995 2008 totalling 484 cases, and related them to travel statistics and antimalarial drug sales. The number of visits to malaria-endemic areas increased, whereas malaria cases did not, and a decreasing trend appeared in antimalarial drug sales. Infections were mostly acquired in Africa (76%). The most common species was Plasmodium falciparum (61%). Of all cases, 42% proved of foreign origin; in 89%, the infection was contracted in the region of birth, implying that immigrants visiting friends and relatives constitute a risk group with a particular need for pre-travel advice. Study IV analyzed in detail the background information on malaria cases diagnosed in Finland 2003 2011. The data included 265 cases, 54% of whom were born in malaria-endemic countries, and 86% currently lived in non-endemic regions. Of those born in non-endemic regions, 81% had received pre-travel advice, but only 20% of those born in endemic ones. Among travellers infected with P. falciparum, 4% reported regular use of appropriate chemoprophylaxis, yet individual rechecking by interview revealed that none had been fully compliant. These data suggest that, if taken conscientiously, mefloquine, atovaquone/proguanil, and doxycycline are effective as chemoprophylaxis against P. falciparum malaria.
  • Arovuori, Kyösti (Helsingin yliopisto, 2015)
    This dissertation analyses the effectiveness of the Common Agricultural Policy of the European Union. The motivation of this study arises from the fact that there is a lack of empirical research on the effects of policy instruments on the stated policy objectives. In addition, most of the analysis in the literature has focused on the policy objective to secure farmers incomes and, thus, on the efficiency of income redistribution. The policy effectiveness is defined as the ability of agricultural policy to respond to the stated policy objectives, given the general economic and structural conditions under which the policies operate. In this study, an empirical analysis of the effects of implemented policies and policy reforms on the stated policy objectives in the Common Agricultural Policy of the European Union is conducted. The analysis is carried out at the EU15 level and the time period analysed ranges from 1975 to 2007. In the empirical part, an econometric model utilising panel data for the EU15 countries is built. In the model, the development of the defined policy target variables is explained with policy variables and a set of economic and structural control variables. The results show that policy target variables have, in general, developed in the desired direction. The productivity of agriculture has increased, markets have been stable, self-sufficiency ratios have been achieved and the real term food prices have declined. However, farmers incomes have mostly declined. The implemented agricultural policy reforms have improved the policy effectiveness in general. The main contribution of the implemented reforms has been to the use of resources in agriculture. A policy shift from coupled price support to direct payments has released resources from agriculture to be utilised in other sectors. On the other hand, policy reforms have led to increasing price variation. This is a self-explanatory impact in the sense that administrative price setting was reduced and later on abolished in the policy reforms. According to this study, the impact of agricultural policy on the policy objectives is multifunctional. The implemented policy instruments may also have worked in the opposite direction compared to the targets set. While agricultural policies have clearly contributed to increasing agricultural productivity, they have also absorbed resources into the sector which could have been utilised more efficiently in other sectors. The country-level heterogeneity of economic and agricultural structures has increased especially due to the recent enlargements of the EU from EU15 to EU28. The analysis in this study shows that country-level heterogeneity has a significant impact on the development of policy target variables. Although the implemented policy reforms have made a contribution towards the desired direction and improved the effectiveness of the policies, it is a major challenge for future agricultural policies to effectively tackle the different structures.
  • Li, Jianyong (Helsingin yliopisto, 2015)
    Assessing the numerical relationships among pollen spectra, human influence and climate is a critical prerequisite for obtaining reliable and robust pollen-based quantitative reconstructions for past climatic variations and human-induced environmental changes. Here, modern pollen data from small river catchments in human-influenced mountainous ecosystems of northern China are collected to investigate the pollen/vegetation/land-use relationships. The effects of human influence index (HII) in comparison with key climate variables on influencing modern pollen distribution are evaluated using a continental-scale dataset. The importance of different climate variables accounting for modern pollen distribution is quantified for identifying the determinant variables that can be reconstructed in different bioclimatic regions of China. Furthermore, pollen-based calibration models for HII and climate parameters are developed and applied to a set of fossil pollen data for quantitative human influence and climate reconstructions. In addition, the bias caused by long-term and intensive human influence on vegetation in quantitative pollen-based palaeoclimatic reconstructions is examined, and pollen-based climate reconstructions are used to validate the reliability of climate model simulations. The results reveal that pollen assemblages from surface sediment samples in the Tuoliang and Qipanshan catchments from northern China are dominated by Artemisia, Chenopodiaceae, Pinus and Selaginella sinensis. Pollen types indicative of human activities such as Cerealia-type are common but not abundant. Artemisia and Chenopodiaceae percentages decline with decreasing altitude, whereas Pinus and S. sinensis percentages increase, suggesting that saccate Pinus pollen and S. sinensis spores can be transported further than non-saccate pollen types and that pollen sorting is taking place over water transportation. There is a positive linear relationship between percentages of Cerealia-type pollen and proportions of farmland, whereas correlation between percentages of trees, shrubs and herbs pollen and proportions of woodland, scrubland and grassland respectively is poor. This can be attributed to the differences in pollen productivity, deposition and dispersal. These findings may provide the basis for a modeling approach to reconstruct past land-cover changes at small catchment scale in northern China. The effects of HII on accounting for modern pollen distribution are smaller than annual precipitation (PANN) or annual average temperature (TANN) in most regions of China, with the exception of east-central China that has a significant pollen HII relationship. The quantified importance of six climate variables (PANN; TANN; actual/potential evapotranspiration ratio, Alpha; mean temperature of the warmest month, MTWA; mean temperature of the coldest month, MTCO; annual sum of the growing degree days above 5 °C, GDD5) for the geographical distribution of modern pollen data differs among regions in China. Hydrological variables are more important than temperature-related variables in north-western and -eastern China and the Tibetan Plateau, whilst temperature-related variables are dominant in east-central and southern China. This suggests that the critical climate variables which can be optimally reconstructed from pollen data vary regionally in China. These regional differences are common to other large-scale surface pollen datasets and need to be taken into considerations in pollen-based climate reconstructions. The calibration models for HII, PANN and TANN show good statistical performances and are applied to a fossil pollen sequence from Lake Tianchi in east-central China. The inferred HII increases abruptly at 1100 cal. yr BP and remains high until present-day. Correspondingly, the reconstructed PANN shows a sudden decrease since 1100 cal. yr BP, whereas TANN displays an increase. However, other independent palaeoclimatic records mostly contradict the reconstructed climatic trends for the last 1100 years, suggesting that the Tianchi climate reconstructions for this period are biased by the intensification of human influence. This bias probably occurs in other palaeoclimatic reconstructions using pollen data from other regions where human impact on vegetation is also intensive and long-lasting. The calibration models for the summer monsoon precipitation (Pjja) also exhibit good predictive performances and are applied to a set of fossil pollen data from monsoonal China. The Holocene Pjja reconstructions from northern China are stacked to one record, showing that Pjja increases since 9500 cal. yr BP, reaches the maximum at 7000 4000 cal. yr BP, and falls down to present-day. The reconstructions are compared with Pjja simulations from the Kiel Climate Model (KCM). The model output and reconstructions are different for the early-Holocene for which the model suggests higher Pjja. The simulated Pjja changes for the Holocene summer monsoon maximum (HSMM) are lower than the reconstructed changes. The increasing or decreasing Pjja trends before or after the HSMM are more pronounced in the reconstructions than in the model results. Other palaeoclimatic data show substantial Holocene monsoon precipitation changes. This suggests that the KCM underestimates the magnitude of the summer monsoon precipitation changes. The results obtained in this work highlight the necessity for critically assessing the modern pollen/human-influence/climate relationships at various spatial scales and can be used to improve the correctness and robustness of pollen-based climate and human influence reconstructions in China and other regions of the world with large-scale surface pollen datasets. The pollen-based calibration models and their forthcoming applications can be employed to further quantitatively explore the past environmental changes and climatic dynamics and to validate the accuracy of climate model output.
  • Müller, Susann (Helsingin yliopisto, 2015)
    Biogeochemical processes in sea ice and the ice-water interface depend on abiotic processes and biological activity. Abiotic processes in sea ice are controlled by the crystallization process of freezing water and the associated formation of saline brine. Also the heat budget of sea ice and the resulting changes in abiotic properties such as porosity and salinity need to be taken into account. The dissolved fraction of sea ice brines contain ions and dissolved organic matter (DOM). Ions are rejected from the ice by diffusion and gravity drainage whereas dissolved organic matter with highly complex and diverse chemical composition can react in many ways with other molecules and surfaces. Hence, the present work compares the behavior of different fractions of DOM to the ones of salts during initial sea ice formation. Controlled tank studies were combined with natural sea ice sampling to exclude the disadvantages of both systems such as the effects of small-scale experiments, artificial additions in tank experiments and the unknown history of natural samples. The studies were conducted with brackish sea ice from the Baltic Sea with its high nutrient and DOM concentrations, but also with oceanic sea ice from the North Sea and the Arctic Ocean. This allows a general conclusion about the behavior of solutes during the formation of sea ice. The present studies indicate that the major seawater ions are significantly fractionated due to differential diffusion and coupled diffuse-convective salt transport through the brine channel network. Ions with a lower diffusivity than Cl¯, in this study SO42¯, Ca+ and Mg2+, remained longer in the brine channel network and got therefore enriched in sea ice relative to Cl¯. K+, on the other hand, diffused faster than Cl¯ and was depleted in sea ice in this study. The behavior of DOM in sea ice was more complex compared to ions because of the complex structure of DOM and the effect of secondary processes on DOM, such as biological production and degradation in sea ice. The quantification of DOM is challenging since only certain fractions such as chromophoric DOM can be measured instead of estimating the total concentration of DOM. Nevertheless, the present studies on DOM in sea ice from Baltic and North Sea water indicated enrichment of DOM compared to sea water ions. The magnitude of this enrichment was higher than expected from diffusion and convection following the transport of salts. The enrichment varied among DOM fractions with highest enrichment of amino-acid like DOM and lowest enrichment of humic-like substances. The results therefore suggest that additional processes, such as selective drainage that depends on the chemical properties of the DOM molecules, affect the enrichment of DOM in sea ice. The optical properties of sea ice were used to develop a bio-optical model. The model estimates the primary production in Baltic Sea ice based on the absorption by particles and chromophoric DOM and the quantum yield for C fixation calculated from photosynthesis-irradiance curves. The results were compared to in situ primary production measurements. The combination of modelled primary production estimates and the measurements of optical properties and primary production in different types of Baltic Sea ice gave a good overview over bio-optical properties in Baltic Sea ice and can be used as a tool to improve different parameters of ecosystem models.
  • Franti, Lasse (Helsingin yliopisto, 2015)
    The gauge/gravity duality connects the dynamics of gravity theories in the bulk with the dynamics of field theories on the boundary. In this thesis we introduce two thermalization scenarios and investigate them using a suitable holographic description. We will first study the thermalization of equal-time correlators and entanglement entropy in a hyperscaling violating AdS-Lifshitz-Vaidya metric. This work verifies the agreement between numerical procedures and preceding analytical predictions and generalises the previous studies of thermalization in this kind of situations. In the latter part we will use the duality to describe the quark-qluon plasma created in heavy ion collisions. The anisotropic plasma is modelled by introducing anisotropies into the source on the gravity side and letting them evolve according to the equations of motion. The boundary dynamics is extracted by finding the boundary stress-energy tensor. The results agree with the conventional models. The situations considered here are rather simple but this work demonstrates the applicability of holography in the anisotropic case.
  • Lindgren, Maija (Helsingin yliopisto, 2015)
    Psychosis is usually preceded by a prodromal period. This phase is characterized by psychotic-like symptoms, attenuated positive symptoms not severe enough to reach a psychotic level. For example, a person may hear voices that are not real. The objective of this study was to investigate whether it is possible and useful to screen for psychosis risk in an unselected clinical adolescent population seeking help for psychiatric symptoms. By finding which symptoms predict transition to a severe psychiatric illness, these risk symptoms can be identified early, enabling effective intervention. This study collected data on adolescent psychiatric patients aged 15 18 years in Helsinki during the years of 2003 2004 and 2007 2008. The participants were screened using the Prodromal Questionnaire (PQ) for prepsychotic symptoms, completed by 731 adolescents. The Structured Interview for Prodromal Syndromes (SIPS) was administered to 174 adolescents to ascertain their psychosis risk status, and broad cognitive testing was done. The participants were followed via patient files and the national hospital discharge register for 3 9 years. A third of the adolescents were identified as psychosis risk patients, but psychosis incidence during follow-up was low, and psychosis risk was not specifically predictive of psychosis. Hospital admissions for psychotic disorder were predicted by the depersonalization symptom intensity of the questionnaire and the positive symptom intensity of the interview. In addition, psychosis risk status predicted psychiatric hospitalizations overall during the following years. Visuospatial performance was poorer among the adolescents with a psychosis risk compared to other patients. Particularly poorer verbal performance was associated with stronger negative symptoms among adolescent patients, regardless of the psychosis risk status. Psychosis risk was associated with suicidal ideation among the adolescent psychiatric patients, but did not predict an increased risk of severe, hospital-treated self-harm during follow-up. The best predictor of intentional self-harm was emotional inexpressivity. Psychotic-like symptoms are common in general adolescent psychiatric services, but the development of psychosis is rare, and predicting psychosis with psychotic-like symptoms is not possible in the clinical environment. However, identifying and treating psychotic-like symptoms is important, as not only are they often distracting experiences in themselves, they can also be associated with cognitive deficits and suicidality, predict hospitalizations, and thus indicate a more serious disorder.
  • Poikolainen, Janne (Nuorisotutkimusverkosto/Nuorisotutkimusseura, 2015)
    The study introduces a historical perspective to the discussion on fandom by examining the emergence of popular music fan culture in Finland from the 1950s to early 1970s. The analysis focuses on the ways the forms and meanings of music fandom, as well as the images attributed to fans, developed in the interaction between the music industry, publicity and audience. The source material consists mainly of written reminiscences on popular music and fandom, and music magazines from the research period. The material also includes e.g. fan letters and statistics. The historical context of the analysis is comprised of the substantial changes in youth brought about by the post-war social change. In the study, these changes are referred to as the modernization of youth. The study examines the technological, social and cultural changes linked to the change in youth that facilitated the emergence of the fan culture. Secondly, the study identifies the socio-cultural needs, created by modernization, to which music fandom as a phenomenon responded. In terms of content, the analysis focuses on three dimensions of fan culture. The first dimension comprises the musical and material settings of fandom, such as recordings, concerts and music magazines. The second consists of media discourses concerning the fan phenomenon. Here the study also questions and disassembles the gendered stereotypes constructed within the discourses. The third dimension comprises the socio-cultural meanings of fandom, particularly in respect of identity work taking place in the forms of identification and social distinction. Scrutiny of these dimensions also highlights the links between the fan phenomenon and the constituent phenomena of modernizing youth: for example, the mediatization, Anglo-Americanization and sexualization of youth culture, as well as the weakening of the traditional identity models. The study shows that the emergence of fan culture was a process where the media contents and ideas concerning fandom interacted in multi-dimensional ways between the various actors. The music industry, media publicity and fan audience formed the macro-level of this interactive network. The contents of fandom formed within this framework assumed their practical meaning in the daily lives of young people. These everyday meanings of fandom were concretized in the form of various consumption and production practices, through which the macro-level interrelationships were again redefined.
  • Creutz, Katja (Hakapaino, 2015)
    This thesis explores the law of state responsibility in international law from a critical standpoint. The main argument is that there are no longer justifications for holding state responsibility as the foremost responsibility system in international law. The importance of state responsibility has diminished; state responsibility has moved from a paradigm to the periphery in the international legal order. The thesis advances on two fronts to prove the veracity of the argument. First, it evaluates the law of state responsibility as adopted in the International Law Commission s 2001 Articles on State Responsibility in order to pinpoint problems in the general state responsibility rules. Second, the thesis explores how alternative constructions of responsibility, that is international liability and international criminal law, have overtaken portions of the state responsibility domain. Functionality is presented as the most suitable appraisal framework for an analysis of how state responsibility rules respond to challenges such as globalization and its actors, the diversity of responsibility ideas, and the normative fragmentation of responsibility in international law. The thesis thus prioritizes the practical usefulness of a responsibility regime in international law rather than abstract system-building functions. The exploration into the functionality of rules for state responsibility is preceded by an excursion into the evolution of state responsibility. The historical outlook will reveal how the rules developed, what dilemmas have characterized the state responsibility project and how these tensions have affected the formation of the ILC state responsibility rules. The critical appraisal of the general rules of state responsibility addresses a range of problems that relate to legal pluralization, form, function and implementation. It is argued that the state responsibility rules are unable to respond to the multi-actor and multi-issue world, to norm differentiation, and to the diversity of function, all of which contributes to the marginalization of the law of state responsibility. The thesis contends that the decision to create one all-embracing responsibility system for all kinds of violations of international law in order to build a credible international legal order actually led to lessened functionality. Real-world problems of crucial importance to the international community cannot be effectively handled under the state responsibility regime. From the demerits of state responsibility, the thesis proceeds to explore particularized responsibility regimes that have developed and strengthened as alternatives to state responsibility. International liability and international criminal law rules are explored from specific points of functionality that state responsibility is ill-suited to handle, i.e. social control, collectivity and the signalling effect. It is submitted that their particularity is an asset that allows regimes to deal with real problems in flexible and creative ways. They have thus effectively contributed to the side-tracking of state responsibility.
  • Kangas, Kaisa (Helsingin yliopisto, 2015)
    The starting point for this dissertation is whether the concept of Zariski geometry, introduced by Hrushovski and Zilber, could be generalized to the context of non-elementary classes. This leads to the axiomatization of Zariski-like structures. As our main result, we prove that if the canonical pregeometry of a Zariski-like structure is non locally modular, then the structure interprets either an algebraically closed field or a non-classical group. This is a counterpart to the result by Hrushovski and Zilber which states that an algebraically closed field can be found in a non locally modular Zariski geometry. It demonstrates that the concept of a Zariski-like structure captures one of the most essential features of a Zariski geometry. Finally, we give a non-trivial example by showing that the cover of the multiplicative group of an algebraically closed field of characteristic zero is Zariski-like. We define a Zariski-like structure as a quasiminimal pregeometry structure that has certain properties. Instead of assuming underlying topologies, we formulate the axioms for a countable collection C of Galois definable sets that have some of the properties of irreducible closed sets from the Zariski geometry context. Quasiminimal classes are abstract elementary classes (AECs) that arise from a quasiminimal pregeometry structure. They are uncountably categorical, and have both the amalgamation property (AP) and the joint embedding property (JEP), and thus also a model homogeneous universal monster model, which we denote by M. To adapt Hrushovski's and Zilber's proof to our setting, we first generalize Hrushovski's Group Configuration Theorem to the context of quasiminimal classes. For this, we develop an independence calculus that has all the usual properties of non-forking and works in our context. We then prove the group configuration theorem and apply it to find a 1-dimensional group, assuming that the canonical pregeometry obtained from the bounded closure operator is non-trivial. A field can be found under the further assumptions that M does not interpret a non-classical group and the canonical pregeometry is non locally modular. Finally, we show that the cover of the multiplicative group of an algebraically closed field, studied by e.g. Boris Zilber and Lucy Burton, provides a non-trivial example of a Zariski-like structure. Burton obtained a topology on the cover by taking sets definable by positive, quantifier-free first order formulae as the basic closed sets. This is called the PQF-topology, and the sets that are closed with respect to it are called the PQF-closed sets. We show that the cover becomes Zariski-like after adding names for a countable number of elements to the language. The axioms for a Zariski-like structure are then satisfied if the collection C is taken to consist of the PQF-closed sets that are definable over the empty set.
  • Leino, Aleksi (Helsingin yliopisto, 2015)
    Ions in the keV energy range are regularly used in the semiconductor industry for device fabrication. Irradiation with ions of higher energies can also induce favorable structural changes in the irradiated samples. Among these, irradiation effects of the so-called swift heavy ions (SHIs, heavy ions with specific energies in the 1 MeV / amu range) in electrically insulating materials are particularly interesting. Despite the wide range of existing applications (filters, printed circuit boards and geological dating) and application potential (fuel cells, cell mimicking membranes) of SHI irradiation, the mechanisms by which SHIs interact with insulators are still under debate. Modelling of SHIs is a very challenging task as, contrary to ions with lower energies, they mostly interact with electrons, inducing lots of electronic excitations. Incorporating the latter with atomistic dynamics is especially difficult in insulators, and the methods have not yet been fully established. SHIs can induce a cylindrical region of structural transformation known as an ion track. In crystalline silicon dioxide, a track consists of an amorphized region that is typically several microns long and has a radius of less than ten nanometers. Furthermore, it was recently found out that SHI irradiation can be used to induce a shape transformation in metal nanoclusters (NCs) that are embedded in amorphous silicon dioxide. Spherical NCs (radius 1-50 nm) elongate along the ion beam direction and are shaped into nanorods or prolate spheroids. The phenomenon can be exploited to produce large arrays of equally aligned nanoclusters within a solid substrate, which is difficult to achieve otherwise. In this thesis, ion track formation and the elongation of gold nanoclusters in silicon dioxide are studied using so called two-temperature molecular dynamics simulations. The structure of the tracks is studied and a mechanism is proposed for the nanoparticle elongation effect. The work presented here is a step towards the understanding of SHI related effects in a broader range of insulating materials for the SHI based applications.
  • Ilina, Polina (Helsingin yliopisto, 2015)
    Gene therapy offers promise for the treatment of both inherited and acquired diseases through the introduction of genetic material into target cells. The primary challenge for gene therapy is to develop a safe and efficient method for the delivery of therapeutic genetic material to the specific intracellular target. Non-viral carriers have received significant attention because of their potential to overcome the limitations of viral-based systems. However, their relatively low efficacy is a major obstacle to their clinical application. A thorough understanding of the key factors affecting the gene delivery process will provide clues on how to develop more effective carriers. This thesis focuses on the role of the carrier and the endocytic pathway in non-viral gene delivery, and also suggests improvements in the experimental methodology that would make it possible to obtain more reliable results in nanoparticle uptake studies. More effective carriers for gene delivery are very much needed. We tested the novel pentaspheric lysine-based dendrimer and its analog, modified with fatty acid residues, for their gene delivery capacity. We demonstrated that despite their relatively low in vitro transfection efficacy, lysine dendrimers have good plasmid DNA (pDNA) binding and protective properties, and can therefore be used as the basis for the development of more effective carriers. A detailed understanding of the cellular kinetics of gene delivery systems is critical to the further development of more effective carriers. We studied the impact of the carrier and of the endocytic pathway on cell uptake and the intracellular processing of genetic material (pDNA). The highly sensitive method of quantitative real-time PCR was applied to the study of the intracellular kinetics of pDNA introduced by the cationic polymer PEI, cationic lipid DOTAP, and CaP precipitates at multiple time points after transfection. The results obtained indicate that the carrier affects the cell uptake and the intracellular kinetics, and therefore predetermines the main transfection-limiting step. Furthermore, we demonstrated the important role of the post-nuclear processes in efficient non-viral gene delivery. The impact of a specific endocytic pathway was studied by the inhibition of either the clathrin- or dynamin-mediated endocytosis. Selective blockage of endocytosis was achieved by two approaches, namely the genetically manipulated cell lines and the chemical inhibitors of endocytosis. Analysis of the intracellular kinetics of pDNA in the genetically blocked cells revealed that neither the amount of pDNA taken up by the cell, nor the intracellular pDNA elimination, but the amount of pDNA delivered to the nucleus was indicative of the significance of the particular pathway in the resulting efficacy of the carrier. A comparison of chemical and genetic means for blocking endocytosis revealed the limitations of both these methods. A careful optimization of the method and the use of several alternative approaches is recommended in order to obtain more reliable data. We suggest that the characterization of in vitro cell models for the expression and activity of specific endocytic pathways (endocytic profiling) would facilitate the interpretation of the data obtained in nanoparticle uptake studies. Endothelial and epithelial cells are widely utilized in such studies because they form substantial barriers en route from the administration site to the target tissue. We performed endocytic profiling of the epithelial CaCo-2 cells and the endothelial hCMEC/D3 cells at different stages of differentiation. The expression of genes involved in specific endocytic pathways was analysed at the mRNA level by quantitative real-time PCR, and at the protein level by Western blotting. The endocytic activities of the cells were analyzed by flow cytometry. We concluded that the mRNA expression of the endogenous proteins involved in particular endocytic pathways can be indicative of the expression and activity of these pathways. Furthermore, we showed that the differentiation status of the cells affects their endocytic activity, and must therefore be taken into account when designing nanoparticle uptake and transcellular permeability experiments. A comparison of the endocytic profiles of cell lines with primary cells revealed clear discrepancies, pointing to the importance of careful selection of in vitro models for endocytosis-related studies. Overall, our study has improved the mechanistic understanding of the non-viral gene delivery process. We have described how the carrier and endocytic pathway both affect intracellular kinetics and the efficacy of gene transfer. Furthermore, we have demonstrated the importance of method optimization and the endocytic profiling of in vitro cell models in improving the quality of the obtained data.
  • Liikanen, Elina (Helsingin yliopisto, 2015)
    The role of literature in the construction of cultural memory: three modes of representing the Spanish Civil War and the Franco dictatorship in recent Spanish novels This dissertation focuses on representations of the Spanish Civil War and the Franco dictatorship in Spanish novels written by third-generation authors (born between 1960 and 1975) and published between 1997 and 2008, during the so-called memory boom . I argue that these novels function as media of cultural memory in Spanish memory culture by producing, shaping and disseminating images, versions and interpretations of Spain s violent past. According to Astrid Erll, certain combinations of formal features constitute different modes of representation within the medium of literature, which in turn may elicit different modes of cultural remembering among readers. Based on the narratological analysis of ten novels, I propose in this study that there are three main modes of representing the past in third-generation Spanish novels on the Civil War and the Franco dictatorship. I call these modes the experiential mode, the reconstructive mode and the dissenting mode. The experiential mode represents the past as a lived-through experience. Rather than examining the process of remembering and narrating the past in the present, experiential novels create the illusion of direct access to the past and the protagonists minds so that the reader can share in the subjective experience of the narrated events. In this way, the novels turn the reader into a vicarious witness to the past. The main function of the experiential mode is to promote empathy and understanding towards certain individuals or groups in the past. The reconstructive mode focuses on the process of investigation and reconstruction of past events led by a narrator-character set in the present day. The novels examine the intergenerational transmission of memories and emphasise the meaning of the past as a source of personal and collective identity in the present. Even if the novels represent the past as a subjective reconstruction and often employ metafictional and auto-fictional techniques to explore the limits of art and reality, history and fiction, they still end up presenting one version of the past as the true one. By depicting the narrator-character s search for historical truth and personal and cultural self-understanding, the reconstructive mode tries to convince the reader of the importance of knowing the past in order to understand the present. The dissenting mode does not offer the reader a supposedly true story of the past, but examines instead the role of literature in the production of knowledge about the past. Dissenting novels are highly self-reflexive texts that reject verisimilitude and disrupt the illusion of reality in order to problematize both prevailing practices of representation and dominant interpretation of the past. The dissenting mode encourages readers to adopt a more critical stance towards the representations of the past that circulate in society by revealing how literature affects our understanding of the past through narrative forms and the stories it tells.
  • Hannula, Miika (Helsingin yliopisto, 2015)
    Dependence logic is a novel logical formalism that has connections to database theory, statistics, linguistics, social choice theory, and physics. Its aim is to provide a systematic and mathematically rigorous tool for studying notions of dependence and independence in different areas. Recently many variants of dependence logic have been studied in the contexts of first-order, modal, and propositional logic. In this thesis we examine independence and inclusion logic that are variants of dependence logic extending first-order logic with so-called independence or inclusion atoms, respectively. The work consists of two parts in which we study either axiomatizability or expressivity hierarchies regarding these logics. In the first part we examine whether there exist some natural parameters of independence and inclusion logic that give rise to infinite expressivity or complexity hierarchies. Two main parameters are considered. These are arity of a dependency atom and number of universal quantifiers. We show that for both logics, the notion of arity gives rise to strict expressivity hierarchies. With respect to number of universal quantifiers however, strictness or collapse of the corresponding hierarchies turns out to be relative to the choice of semantics. In the second part we turn attention to axiomatizations. Due to their complexity, dependence and independence logic cannot have a complete recursively enumerable axiomatization. Hence, restricting attention to partial solutions, we first axiomatize all first-order consequences of independence logic sentences, thus extending an analogous result for dependence logic. We also consider the class of independence and inclusion atoms, and show that it can be axiomatized using implicit existential quantification. For relational databases this implies a sound and complete axiomatization of embedded multivalued and inclusion dependencies taken together. Lastly, we consider keys together with so-called pure independence atoms and prove both positive and negative results regarding their finite axiomatizability.
  • Österman, Janina (Helsingin yliopisto, 2015)
    Nitrogen is an indispensable element for plants and animals to be able to synthesise essential biological compounds such as amino acids and nucleotides. Although there is plenty of nitrogen in the form of nitrogen gas (N2) in the Earth s atmosphere, it is not readily available to plants but needs to be converted (fixed) into ammonia before it can be utilised. Nitrogen-fixing bacteria living freely in the soil or in symbiotic association with legume plants, fix N2 into ammonia used by the plants. This is known as biological nitrogen fixation (BNF). In contrast to industrial nitrogen fixation, an energy-demanding process using high temperature and pressure to produce chemical fertilizers, BNF makes use of solar energy alone to complete the same reaction. However, the requirements on compatibility of plants and nitrogen-fixing micro-organism, the rate of conversion and the ability of the micro-organisms to survive in stressful environments are limiting factors of this system. The current demand for more sustainable food production makes BNF an attractive alternative. However, optimization of existing BNF systems as well as development of new highly productive ones is necessary, to be able to replace the use of chemical fertilisers. In order to develop new alternatives, we need to gain more knowledge on the requirements set by both plants and micro-organisms for successful and efficient nitrogen fixation to occur. In this thesis, the nitrogen-fixing legume host Galega (goat s rue) and its symbiotic microbial partner Neorhizobium galegae were used as a model system to investigate the features defining good symbiotic nitrogen fixation. Studies of genetic diversity within the host plant showed that there are genetic traits making a distinction between the two species G. orientalis and G. officinalis, both at a whole-genome level and at the level of specific symbiosis-related genes. Genome sequencing of ten strains of N. galegae provided a useful dataset for studying i) the genomic features separating N. galegae from related nitrogen-fixing bacteria (rhizobia) and ii) the genetically encoded characteristics that divide strains of N. galegae into two separate symbiovars (symbiotic variants that show different phenotypes on the two different Galega host plant species). These studies provided new information on genes possibly involved in determining host specificity and efficiency of nitrogen fixation. In addition, previously unrecognised genetic contents provided insight into the ecology of N. galegae. Most importantly, genome sequencing enabled identification of the noeT gene, responsible for acetylation of the N. galegae Nod factor (signal molecule required for symbiosis). Although the noeT gene did not turn out to be the crucial determinant enabling nodulation of Galega spp. as previously anticipated, these results are important for future studies on mechanisms behind the selectiveness (host specificity) observed in nitogen-fixing symbioses between Galega and N. galegae.
  • Vihervuori , Liisa (Helsingin yliopisto, 2015)
    Many risks and environmental concerns have been linked with the cultivation of genetically modified (GM) trees. Among the most frequently mentioned risks are the unintentional/pleiotropic effects of transgenes on organisms or plant properties that are not the targets of genetic modification. Risks in forest ecosystems are difficult to predict, due to the long life cycles of trees and their complex ecological interactions. This thesis is focused on the interactions between insect and mammal herbivores and transgenic trees. The studied silver birch (Betula pendula Roth) carried a sugar beet chitinase IV gene and the aspen and hybrid aspen (Populus sp.) carried a pine pinosylvin synthase gene, both aiming to improve the trees resistance to fungal diseases, but also potentially affecting insect and mammal herbivores. Tree growth and quality, and insect density and composition were studied during the field trial. The palatability of transgenic trees was studied in the feeding experiments using the lepidopteran herbivores rusty tussock moth and buff-tip (Orgyia antiqua L., Phalera bucephala L.) and the mammalian herbivores mountain hare and roe deer (Lepus timidus L., Capreolus capreolus L.). The studies revealed that sugar beet chitinase IV caused unintentional effects on silver birch: the transgenic trees were smaller and their red leaf colour indicated stress. There were differences in herbivore pressure between transgenic and wild-type birches: GM trees had a higher aphid density but a lower diversity of insect species and visible leaf damages. The relative growth rate of O. antiqua on transgenic birches was lower in comparison to wild-type birches. No difference in the palatability to mammals was found between transgenic and wild-type trees. The results suggest that the impact on herbivores is species- dependent. The pleiotropic effects of the transgenes affecting plant-herbivore interaction can explain these impacts. These results should be taken into account when considering the biosafety of GM trees.
  • Patokoski, Johanna (Helsingin yliopisto, 2015)
    Volatile organic compounds (VOCs) are known to be key players in the atmospheric processes. They are emitted both from natural and non-biogenic sources. Although the biogenic sources are generally dominant compared with the anthropogenic sources in some circumstances the anthropogenic emissions can dominate e.g. in densely populated areas or during pollution plumes. VOCs are ambient trace gases including a vast group of compounds. Some of the VOCs are very reactive, participating in atmospheric transformation processes e.g. secondary organic aerosol formation and growth. Some VOCs are also known to be harmful air pollutants for humans. Thus VOCs can have direct effects on air quality and secondary effects on climate. In this thesis, the aim was to define sources of VOCs, determine influence of transport and transformation of trace gases and aerosols. VOCs were measured mainly by a real time proton transfer reaction mass spectrometer. In addition, gas and liquid chromatography were used. The measurements were conducted at the rural SMEAR II and the urban background SMEAR III sites. Concentrations of VOCs were observed to have inter-annual, seasonal and diurnal variations due to meteorological factors, photochemistry and different sources. The anthropogenic influence was dominant in winter at both sites, while the biogenic influence with enhanced photochemical reactions increased in spring. The mixing and dilution effect also increased in spring and it led to decreased levels of volume mixing ratios (VMRs) in daytime especially in short-lived compounds. The VMRs of aromatic compounds were lower in spring than in winter due to lesser sources and enhanced photochemistry at both sites. At the urban background site there were more local sources than rural site and thus more variation of the diurnal patterns; higher VMRs of VOCs were observed. At both sites long-range transport was found to be an important source for long-lived VOCs. The source areas of long-lived VOCs were studied with trajectory analysis. Eastern Europe including West Russia was found to be major source area for all studied VOCs. In addition, some of them had specific source areas of their own related e.g. to their use in the solvent industry. During measurements there were two different forest fire episodes in Eastern Europe and a one-day prescribed biomass burning experiment. VMRs of VOCs related to burning were observed to increase during these pollution plumes. Atmospheric oxidation of BVOCs (biogenic VOCs) revealed that monoterpenes and their oxidation products by ozone were limiting factors for nucleation particle growth. Study of total OH reactivity showed that there are a number of unmeasured biogenic compounds which may have effects on the local atmospheric chemical processes. Keywords: variations of VOC concentrations, source analysis, long-range transport, forest fires, atmospheric chemistry
  • Tuulos, Antti (Helsingin yliopisto, 2015)
    Winter turnip rape is a biennial crucifer oilseed. A problem associated with winter turnip rape is early sowing time in July, when farms are short of available land. The aim of this study was to determine if winter turnip rape can be established by undersowing with spring cereals without decreasing cereal and winter turnip rape yields. It was also studied, if cutting the vegetative leaves of winter turnip rape in autumn could be performed without affecting the yield and whether the composition of leaf material was nutritionally acceptable to be used as forage. The ability of winter turnip rape to act as a mineral nitrogen scavenging catch crop was studied as well. Field experiments were conducted at University of Helsinki experimental farm in Finland during 2009-2011. Winter turnip rape was either undersown with spring cereal or as pure stands. Cereals were six-row barley, two-row barley, oat and wheat. One third of the winter turnip rape plots were cut after cereal harvest to simulate forage harvest. Winter turnip rape yield and its quality was not compromised due to undersowing with a cereal, when the overwintering conditions following cereal harvest were favorable. Cutting the winter turnip rape stands in autumn decreased seed yield in the following year. Winter turnip rape leaf forage has very high crude protein content and low crude fibre content. The glucosinolate content of winter turnip rape leaf forage is comparable to other forage crucifers. Cereal yield was not decreased by the undersown winter turnip rape and quality of wheat was only slightly affected. Seed yield of six-row barley and oat was increased by the undersown winter turnip rape, indicating a facilitative interaction between the species. Undersown winter turnip rape decreased subsoil nitrate content effectively in late autumn under moist conditions that favored mineralization. Winter turnip rape can be established by undersowing to a cereal without decreasing the yields of both crops. Some cultivars of barley and oat may even benefit from the undersown winter turnip rape possibly due to root interactions. Even though winter turnip rape is nutritionally suitable as forage, autumn forage cuts should be avoided. Winter turnip rape seems to be an effective catch crop, which may decrease the leaching of soil nitrate.
  • Helle, Inari (Helsingin yliopisto, 2015)
    Coastal and marine ecosystems across the globe are heavily impacted by various anthropogenic stressors, which has led to a significant loss of biodiversity and ecosystem services in recent decades. In order to find means to counteract this trend, there is a need to develop methods for assessing the environmental impacts of human activities and the effectiveness of management practices to mitigate the harmful effects. However, this is a challenging task due to the complex interactions within and between the ecosystems and human components, and various uncertainties related to them. Bayesian networks (BNs) are graphical models for reasoning under uncertainty. A BN consists of a set of probabilistic variables connected with links describing causalities within the system. As the states of the variables are described with probability distributions, uncertainty can be described in an explicit manner. BNs also enable integration of qualitative and quantitative knowledge from various sources such as observational data sets, models and expert knowledge. In this thesis I have developed BN models to study environmental risks related to anthropogenic stressors in the Gulf of Finland and the Finnish Archipelago Sea. The main aim is to quantify human impacts on the environment, and to assess the ability of different management measures to lessen these impacts. I focus especially on oil spills resulting from potential tanker accidents and I set out to fill various information gaps related to this recently emerged threat. The thesis includes five papers. In paper I, the main aim is to assess the spatial risk posed by oil spills in the Gulf of Finland and the Finnish Archipelago Sea, and identify species and habitat types with the highest risk. In paper II, I focus on the effectiveness of different oil combating methods to mitigate the negative impacts of oil spills on the ecosystem, and paper III widens the approach to a probabilistic cost-benefit analysis of preventive and post-spill measures. Paper IV deals with multiple risks as, in addition to oil spills, eutrophication and harvesting of species are studied. Paper V reviews and discusses various methods that can be applied to evaluate the uncertainty related to deterministic models, which could increase their usefulness in decision-making. The results suggest that risks related to tanker accidents are distributed unevenly between areas, habitats and species. Furthermore, the results support the current Finnish strategy to base oil combating primarily on offshore recovery vessels instead of chemical dispersants. However, as the efficiency of mechanical recovery is dependent on several factors, there is also a need to develop preventive measures. Although major oil accidents are estimated to be rare events, the costs can be very high, if a spill occurs. The work offers new insights to the oil spill risks in the study area and provides examples how Bayesian networks can be applied in environmental risk assessment. The thesis is a part of the work needed in order to develop comprehensive decision support tools related to environmental risk management in the northern Baltic Sea.
  • Österman, Janina (Helsingin yliopisto, 2015)
    Nitrogen is an indispensable element for plants and animals to be able to synthesise essential biological compounds such as amino acids and nucleotides. Although there is plenty of nitrogen in the form of nitrogen gas (N2) in the Earth s atmosphere, it is not readily available to plants but needs to be converted (fixed) into ammonia before it can be utilised. Nitrogen-fixing bacteria living freely in the soil or in symbiotic association with legume plants, fix N2 into ammonia used by the plants. This is known as biological nitrogen fixation (BNF). In contrast to industrial nitrogen fixation, an energy-demanding process using high temperature and pressure to produce chemical fertilizers, BNF makes use of solar energy alone to complete the same reaction. However, the requirements on compatibility of plants and nitrogen-fixing micro-organism, the rate of conversion and the ability of the micro-organisms to survive in stressful environments are limiting factors of this system. The current demand for more sustainable food production makes BNF an attractive alternative. However, optimization of existing BNF systems as well as development of new highly productive ones is necessary, to be able to replace the use of chemical fertilisers. In order to develop new alternatives, we need to gain more knowledge on the requirements set by both plants and micro-organisms for successful and efficient nitrogen fixation to occur. In this thesis, the nitrogen-fixing legume host Galega (goat s rue) and its symbiotic microbial partner Neorhizobium galegae were used as a model system to investigate the features defining good symbiotic nitrogen fixation. Studies of genetic diversity within the host plant showed that there are genetic traits making a distinction between the two species G. orientalis and G. officinalis, both at a whole-genome level and at the level of specific symbiosis-related genes. Genome sequencing of ten strains of N. galegae provided a useful dataset for studying i) the genomic features separating N. galegae from related nitrogen-fixing bacteria (rhizobia) and ii) the genetically encoded characteristics that divide strains of N. galegae into two separate symbiovars (symbiotic variants that show different phenotypes on the two different Galega host plant species). These studies provided new information on genes possibly involved in determining host specificity and efficiency of nitrogen fixation. In addition, previously unrecognised genetic contents provided insight into the ecology of N. galegae. Most importantly, genome sequencing enabled identification of the noeT gene, responsible for acetylation of the N. galegae Nod factor (signal molecule required for symbiosis). Although the noeT gene did not turn out to be the crucial determinant enabling nodulation of Galega spp. as previously anticipated, these results are important for future studies on mechanisms behind the selectiveness (host specificity) observed in nitogen-fixing symbioses between Galega and N. galegae.
  • Björklund, Heidi (Helsingin yliopisto, 2015)
    Human-caused habitat changes have led to declines of many species by impairing their occurrence, reproduction and interactions with other species. In Northern Europe, intensive forestry has transformed boreal forests and worsened conditions of species dependent on old forest, but the mechanisms by which habitat changes affect species are often unknown. In my dissertation, I examined habitat changes, their effects and habitat-associated breeding performances of three declining forest-dwelling hawks, the northern goshawk (Accipiter gentilis), the common buzzard (Buteo buteo) and the honey buzzard (Pernis apivorus). These species can compete for nest sites and goshawks can also predate the buzzards. Raptors may lack nest sites in managed forests which is why their nesting has been supported by constructing artificial nests. I analysed the breeding success of the hawks in artificial nests since their benefits as a conservation measure were not previously analysed. Finally, I studied the conditions when dominant species can exclude subordinate ones in competition for suitable nest sites. According to my results, old forests have decreased and young forests increased throughout southern Finland. These changes are adverse for the goshawk whose breeding success improves when old spruce forests are more abundant and young forests scarce around their nests. Breeding performances of the common buzzard and honey buzzard were not significantly associated with habitats. However, common buzzards were reluctant to re-occupy nests surrounded by a vastness of old forest. The breeding success of the goshawk and common buzzard was lower in artificial nests than in natural ones. Thus, this measure aimed at enabling hawk breeding in managed forests seems to conflict with its conservation objectives. I discuss the possible contributing factors and directions for further investigations. Interspecific interactions with other raptors proved to be important when a subordinate hawk decided on territory occupancy. Common buzzards prefer to occupy safe territories, where threats of goshawk predation and interference competition with other raptors are small. Thus, interspecific raptors that are predators or competitors may exclude subordinate raptors from their territories, and thereby contribute to their occurrence. Adverse interactions with other raptors may even impede a subordinate raptor from fully exploiting the periodic food peaks. My dissertation shows that for each species, it is crucial to identify the most relevant spatiotemporal scales in order to identify, how external disturbances affect them most. Long-term data is of paramount importance to detect these scales.