Individuals on the Move : Body Condition Dependent Dispersal and Quasi-Local Competition in Metapopulations

Individual movement is very versatile and inevitable in ecology. In this thesis, I investigate two kinds of movement body condition dependent dispersal and small-range foraging movements resulting in quasi-local competition and their causes and consequences on the individual, population and metapopulation level. Body condition dependent dispersal is a widely evident but barely understood phenomenon. In nature, diverse relationships between body condition and dispersal are observed. I develop the first models that study the evolution of dispersal strategies that depend on individual body condition. In a patchy environment where patches differ in environmental conditions, individuals born in rich (e.g. nutritious) patches are on average stronger than their conspecifics that are born in poorer patches. Body condition (strength) determines competitive ability such that stronger individuals win competition with higher probability than weak individuals. Individuals compete for patches such that kin competition selects for dispersal. I determine the evolutionarily stable strategy (ESS) for different ecological scenarios. My models offer explanations for both dispersal of strong individuals and dispersal of weak individuals. Moreover, I find that within-family dispersal behaviour is not always reflected on the population level. This supports the fact that no consistent pattern is detected in data on body condition dependent dispersal. It also encourages the refining of empirical investigations. Quasi-local competition defines interactions between adjacent populations where one population negatively affects the growth of the other population. I model a metapopulation in a homogeneous environment where adults of different subpopulations compete for resources by spending part of their foraging time in the neighbouring patches, while their juveniles only feed on the resource in their natal patch. I show that spatial patterns (different population densities in the patches) are stable only if one age class depletes the resource very much but mainly the other age group depends on it.

Construction of a global map of human gene expression : the process, tools and analysis

This thesis studies human gene expression space using high throughput gene expression data from DNA microarrays. In molecular biology, high throughput techniques allow numerical measurements of expression of tens of thousands of genes simultaneously. In a single study, this data is traditionally obtained from a limited number of sample types with a small number of replicates. For organism-wide analysis, this data has been largely unavailable and the global structure of human transcriptome has remained unknown. This thesis introduces a human transcriptome map of different biological entities and analysis of its general structure. The map is constructed from gene expression data from the two largest public microarray data repositories, GEO and ArrayExpress. The creation of this map contributed to the development of ArrayExpress by identifying and retrofitting the previously unusable and missing data and by improving the access to its data. It also contributed to creation of several new tools for microarray data manipulation and establishment of data exchange between GEO and ArrayExpress. The data integration for the global map required creation of a new large ontology of human cell types, disease states, organism parts and cell lines. The ontology was used in a new text mining and decision tree based method for automatic conversion of human readable free text microarray data annotations into categorised format. The data comparability and minimisation of the systematic measurement errors that are characteristic to each lab- oratory in this large cross-laboratories integrated dataset, was ensured by computation of a range of microarray data quality metrics and exclusion of incomparable data. The structure of a global map of human gene expression was then explored by principal component analysis and hierarchical clustering using heuristics and help from another purpose built sample ontology. A preface and motivation to the construction and analysis of a global map of human gene expression is given by analysis of two microarray datasets of human malignant melanoma. The analysis of these sets incorporate indirect comparison of statistical methods for finding differentially expressed genes and point to the need to study gene expression on a global level.

Empirical Analysis of Competition in Procurement Auctions

A vast amount of public services and goods are contracted through procurement auctions. Therefore it is very important to design these auctions in an optimal way. Typically, we are interested in two different objectives. The first objective is efficiency. Efficiency means that the contract is awarded to the bidder that values it the most, which in the procurement setting means the bidder that has the lowest cost of providing a service with a given quality. The second objective is to maximize public revenue. Maximizing public revenue means minimizing the costs of procurement. Both of these goals are important from the welfare point of view. In this thesis, I analyze field data from procurement auctions and show how empirical analysis can be used to help design the auctions to maximize public revenue. In particular, I concentrate on how competition, which means the number of bidders, should be taken into account in the design of auctions. In the first chapter, the main policy question is whether the auctioneer should spend resources to induce more competition. The information paradigm is essential in analyzing the effects of competition. We talk of a private values information paradigm when the bidders know their valuations exactly. In a common value information paradigm, the information about the value of the object is dispersed among the bidders. With private values more competition always increases the public revenue but with common values the effect of competition is uncertain. I study the effects of competition in the City of Helsinki bus transit market by conducting tests for common values. I also extend an existing test by allowing bidder asymmetry. The information paradigm seems to be that of common values. The bus companies that have garages close to the contracted routes are influenced more by the common value elements than those whose garages are further away. Therefore, attracting more bidders does not necessarily lower procurement costs, and thus the City should not implement costly policies to induce more competition. In the second chapter, I ask how the auctioneer can increase its revenue by changing contract characteristics like contract sizes and durations. I find that the City of Helsinki should shorten the contract duration in the bus transit auctions because that would decrease the importance of common value components and cheaply increase entry which now would have a more beneficial impact on the public revenue. Typically, cartels decrease the public revenue in a significant way. In the third chapter, I propose a new statistical method for detecting collusion and compare it with an existing test. I argue that my test is robust to unobserved heterogeneity unlike the existing test. I apply both methods to procurement auctions that contract snow removal in schools of Helsinki. According to these tests, the bidding behavior of two of the bidders seems consistent with a contract allocation scheme.

The range expansion of the European map butterfly in Finland

Climate change will influence the living conditions of all life on Earth. For some species the change in the environmental conditions that has occurred so far has already increased the risk of extinction, and the extinction risk is predicted to increase for large numbers of species in the future. Some species may have time to adapt to the changing environmental conditions, but the rate and magnitude of the change are too great to allow many species to survive via evolutionary changes. Species responses to climate change have been documented for some decades. Some groups of species, like many insects, respond readily to changes in temperature conditions and have shifted their distributions northwards to new climatically suitable regions. Such range shifts have been well documented especially in temperate zones. In this context, butterflies have been studied more than any other group of species, partly for the reason that their past geographical ranges are well documented, which facilitates species-climate modelling and other analyses. The aim of the modelling studies is to examine to what extent shifts in species distributions can be explained by climatic and other factors. Models can also be used to predict the future distributions of species. In this thesis, I have studied the response to climate change of one species of butterfly within one geographically restricted area. The study species, the European map butterfly (Araschnia levana), has expanded rapidly northwards in Finland during the last two decades. I used statistical and dynamic modelling approaches in combination with field studies to analyse the effects of climate warming and landscape structure on the expansion. I studied possible role of molecular variation in phosphoglucose isomerase (PGI), a glycolytic enzyme affecting flight metabolism and thereby flight performance, in the observed expansion of the map butterfly at two separate expansion fronts in Finland. The expansion rate of the map butterfly was shown to be correlated with the frequency of warmer than average summers during the study period. The result is in line with the greater probability of occurrence of the second generation during warm summers and previous results on this species showing greater mobility of the second than first generation individuals. The results of a field study in this thesis indicated low mobility of the first generation butterflies. Climatic variables alone were not sufficient to explain the observed expansion in Finland. There are also problems in transferring the climate model to new regions from the ones from which data were available to construct the model. The climate model predicted a wider distribution in the south-western part of Finland than what has been observed. Dynamic modelling of the expansion in response to landscape structure suggested that habitat and landscape structure influence the rate of expansion. In southern Finland the landscape structure may have slowed down the expansion rate. The results on PGI suggested that allelic variation in this enzyme may influence flight performance and thereby the rate of expansion. Genetic differences of the populations at the two expansion fronts may explain at least partly the observed differences in the rate of expansion. Individuals with the genotype associated with high flight metabolic rate were most frequent in eastern Finland, where the rate of range expansion has been highest.

Meadow plant growth and competition under elevated ozone and carbon dioxide

The main aim of my thesis project was to assess the impact of elevated ozone (O3) and carbon dioxide (CO2) on the growth, competition and community of meadow plants in northern Europe. The thesis project consisted of three separate O3 and CO2 exposure experiments that were conducted as open-top-chamber (OTC) studies at Jokioinen, SW Finland, and a smaller-scale experiment with different availabilities of resources in greenhouses in Helsinki. The OTC experiments included a competition experiment with two- and three-wise interactions, a mesocosm-scale meadow community with a large number of species, and a pot experiment that assessed intraspecific differences of Centaurea jacea ecotypes. The studied lowland hay meadow proved to be an O3-sensitive biotope, as the O3 concentrations used (40-50 ppb) were moderate, and yet, six out of nine species (Campanula rotundifolia, Centaurea jacea, Fragaria vesca, Ranunculus acris, Trifolium medium, Vicia cracca) showed either significant reductions in biomass or reproductive development, visible O3 injury or any two as a response to elevated O3. The plant species and ecotypes exhibited large intra- and interspecific variation in their response to O3, but O3 and CO2 concentrations did not cause changes in their interspecific competition or in community composition. However, the largest O3-induced growth reductions were seen in the least abundant species (C. rotundifolia and F. vesca), which may indicate O3-induced suppression of weak competitors. The overall effects of CO2 were relatively small and mainly restricted to individual species and several measured variables. Based on the present studies, most of the deleterious effects of tropospheric O3 are not diminished by a moderate increase in CO2 under low N availability, and variation exists between different species and variables. The present study indicates that the growth of several herb species decreases with increasing atmospheric O3 concentrations, and that these changes may pose a threat to the biodiversity of meadows. Ozone-induced reductions in the total community biomass production and N pool are likely to have important consequences for the nutrient cycling of the ecosystem.

The Finnish Competition State and Entrepreneurial Policies in the Helsinki Region

The thesis examines the intensification and characteristics of a policy that emphasises economic competitiveness in Finland during the 1990s and early 2000s. This accentuation of economic objectives is studied at the level of national policy-making as well as at the regional level through the policies and strategies of cities and three universities in the Helsinki region. By combining the analysis of state policies, urban strategies and university activities, the study illustrates the pervasiveness of the objective of economic competitiveness and growth across these levels and sheds light on the features and contradictions of these policies on a broad scale. The thesis is composed of five research articles and a summary article. At the level of national policies, the central focus of the thesis is on the growing role of science and technology policy as a state means to promote structural economic change and its transformation towards a broader, yet ambivalent concept of innovation policy. This shift brings forward a tension between an increasing emphasis on economic aspects – innovations and competitiveness – as well as the expanding scope of issues across a wide range of policy sectors that are being subsumed under this market- and economy oriented framework. Related to science and technology policy, attention is paid to adjustments in university policy in which there has been increasing pressure for efficiency, rationalisation and commercialisation of academic activities. Furthermore, political efforts to build an information society through the application of information and communication technologies are analysed with particular attention to the balance between economic and social objectives. Finally, changes in state regional policy priorities and the tendency towards competitiveness are addressed. At the regional level, the focus of the thesis is on the policies of the cities in Finland’s capital region as well as strategies of three universities operating in the region, namely the University of Helsinki, Helsinki University of technology and Helsinki School of Economics. As regards the urban level, the main focus is on the changes and characteristics of the urban economic development policy of the City of Helsinki. With respect to the universities, the thesis examines their attempts to commercialise research and thus bring academic research closer to economic interests, and pays particular attention to the contradictions of commercialisation. Related to the universities, the activities of three intermediary organisations that the universities have established in order to increase cooperation with industry are analysed. These organisations are the Helsinki Science Park, Otaniemi International Innovation Centre and LTT Research Ltd. The summary article provides a synthesis of the material presented in the five original articles and relates the results of the articles to a broader discussion concerning the emergence of competition states and entrepreneurial cities and regions. The main points of reference are Bob Jessop’s and Neil Brenner’s theses on state and urban-regional restructuring. The empirical results and considerations from Finland and the Helsinki region are used to comment on, specify and criticise specific parts of the two theses.

Effect of GPS on understanding relationship between mobile map and environment : A field experiment

Objectives: GPS technology enables the visualisation of a map reader s location on a mobile map. Earlier research on the cognitive aspects of map reading identified that searching for map-environment points is an essential element for the process of determining one s location on a mobile map. Map-environment points refer to objects that are visualized on the map and are recognizable in the environment. However, because the GPS usually adds only one point to the map that has a relation to the environment, it does not provide a sufficient amount of information for self-location. The aim of the present thesis was to assess the effect of GPS on the cognitive processes involved in determining one s location on a map. Methods: The effect of GPS on self-location was studied in a field experiment. The subjects were shown a target on a mobile map, and they were asked to point in the direction of the target. In order for the map reader to be able to deduce the direction of the target, he/she has to locate himself/herself on the map. During the pointing tasks, the subjects were asked to think aloud. The data from the experiment were used to analyze the effect of the GPS on the time needed to perform the task. The subjects verbal data was used to assess the effect of the GPS on the number of landmark concepts mentioned during a task (landmark concepts are words referring to objects that can be recognized both on the map and in the environment). Results and conclusions: The results from the experiment indicate that the GPS reduces the time needed to locate oneself on a map. The analysis of the verbal data revealed that the GPS reduces the number of landmark concepts in the protocols. The findings suggest that the GPS guides the subject s search for the map-environment points and narrows the area on the map that must be searched for self-location.