Affektiivisuuden laskennallinen määritteleminen

One of the most tangled fields of research is the field of defining and modeling affective concepts, i. e. concepts regarding emotions and feelings. The subject can be approached from many disciplines. The main problem is lack of generally approved definitions. However, e.g. linguists have recently started to check the consistency of their theories with the help of computer simulations. Definitions of affective concepts are needed for performing similar simulations in behavioral sciences. In this thesis, preliminary computational definitions of affects for a simple utility-maximizing agent are given. The definitions have been produced by synthetizing ideas from theories from several fields of research. The class of affects is defined as a superclass of emotions and feelings. Affect is defined as a process, in which a change in an agent's expected utility causes a bodily change. If the process is currently under the attention of the agent (i.e. the agent is conscious of it), the process is a feeling. If it is not, but can in principle be taken into attention (i.e. it is preconscious), the process is an emotion. Thus, affects do not presuppose consciousness, but emotions and affects do. Affects directed at unexpected materialized (i.e. past) events are delight and fright. Delight is the consequence of an unexpected positive event and fright is the consequence of an unexpected negative event. Affects directed at expected materialized (i.e. past) events are happiness (expected positive event materialized), disappointment (expected positive event did not materialize), sadness (expected negative event materialized) and relief (expected negative event did not materialize). Affects directed at expected unrealized (i.e. future) events are fear and hope. Some other affects can be defined as directed towards originators of the events. The affect classification has also been implemented as a computer program, the purpose of which is to ensure the coherence of the definitions and also to illustrate the capabilities of the model. The exact content of bodily changes associated with specific affects is not considered relevant from the point of view of the logical structure of affective phenomena. The utility function need also not be defined, since the target of examination is only its dynamics.

"Towards 'a real reunion'?" : Archbishop Aleksi Lehtonen's efforts for closer relations with the Church of England 1945–1951

Mika KT Pajusen väitös "Towards 'a real reunion'?" – Archbishop Aleksi Lehtonen's efforts for closer relations with the Church of England 1945–1951 on yleiseen kirkkohistoriaan lukeutuva tutkimus Englannin kirkon ja Suomen evankelis-luterilaisen kirkon välisistä suhteista Aleksi Lehtosen arkkipiispakaudella 1945–1951. Suhteita on tutkittu kolmesta näkökulmasta: ekumeenisesta, poliittisesta ja kirkkopoliittisesta. Tutkimuskausi alkaa pastori H.M. Waddamsin joulukuussa 1944 Suomeen tekemän vierailun jälkimainingeista ja päättyy arkkipiispa Lehtosen kuolemaan pääsiäisenä 1951. Kirkollisten suhteiden kehitystä rytmittivät lukuisat vierailut, jotka osoittivat Englannin kirkon asenteen muuttumisen sodan aikaisesta neuvostomyönteisyydestä kylmän sodan aikaiseen täysin vastakkaiseen kantaan. Englantilaiset vieraat kohtasivat Suomessa sekä kirkon että yhteiskunnan ylimmän johdon. Molemmat maat olivat valmiita tukemaan hyviä kirkollisia suhteita tilanteen niin salliessa, joskaan eivät kovin suunnitelmallisesti. Suomen evankelis-luterilainen kirkko käytti hyviä suhteita Englannin kirkkoon saadakseen tukea ja ymmärrystä omalle kirkolleen ja yhteiskunnalleen kokemaansa Neuvostoliiton uhkaa vastaan erityisesti vaaran vuosina 1944–1948. Englannin kirkko halusi tukea suomalaista sisarkirkkoaan, mutta varoi, ettei tuottaisi tuellaan enemmän haittaa kuin hyötyä suhteessa Neuvostoliittoon. Sodan jälkeinen ekumeeninen jälleenrakentaminen lähensi kirkkoja toisiinsa. Lehtonen pyrki jatkamaan 1930-luvun kirkkojen välisiä, ehtoollisvieraanvaraisuuden saavuttaneita neuvotteluita kohti täyttä kirkollista yhteyttä. Häntä motivoi sekä evankelis-katolinen teologia että pyrkimys tukea oman maan ja kirkon läntisiä yhteyksiä. Tämä haastoi Englannin kirkon ekumeenisen linjan, joka Suomen kirkon sijasta pyrki jatkamaan neuvotteluja Tanskan, Norjan ja Islannin luterilaisten kirkkojen kanssa, joilla ei vielä ollut virallista ekumeenista sopimusta Englannin kirkon kanssa. Lehtosen pyrkimyksistä huolimatta Englannin kirkko päätyi jättämään Suomen tilanteen hautumaan. Sillä se tarkoitti suhteiden koetinkivenä olleen historiallisen piispuuden leviämistä läpi Suomen kirkon ennen kuin katsoi olevansa valmis jatkamaan kohti täyttä kirkollista yhteyttä. Molemmissa kirkoissa vaikutti pieni, innokkaiden, lähempiä suhteita toivoneiden kirkollisten vaikuttajien ydinjoukko. Englantilaisia Suomen-ystäviä motivoi tarve auttaa Suomea hankalassa poliittisessa tilanteessa. Suomessa arkkipiispa Lehtonen tuki korkeakirkollista liturgista liikettä, jolla oli läheinen yhteys anglikaanisuuteen, mutta joka sai vastaansa vanhoilliset pietistit. Suomen kirkon yleinen mielipide asettui etupäässä pietistiselle kannalle, jolle anglikaanisuus näyttäytyi teologisesti sekä liian katolisena että liian reformoituna. Kirkolliset suhteet tasaantuivat vuoden 1948 Lambeth-konferenssin jälkeen, joka rohkaisi anglikaanisia kirkkoja hyväksymään 1930-luvun neuvottelujen lähempiin kirkollisiin suhteisiin tähtäävät suositukset. Lehtonen näytti tyytyvän tähän. Samaan aikaan lähempää kirkollista kanssakäymistä tukenut ekumeeninen jälleenrakennus tuli tiensä päähän. Lehtonen jatkoi läheisempien suhteiden edistämistä, mutta hänen intonsa hiipui yhdessä heikkenevän terveydentilan kanssa. Osoituksena Lehtosen linjan kapeudesta Suomen evankelis-luterilaisen kirkon piispoista ei löytynyt hänen kuoltuaan ketään, joka olisi jatkanut hänen aktiivista anglikaanimyönteistä linjaansa.

Pitäjän hyödyllisin mies? : Pehr Kalmin käsitys papiston yhteiskunnallisesta tehtävästä

Turun akatemian ensimmäisen talousopin professorin Pehr Kalmin johdolla tarkastettiin vuonna 1757 väitöskirja aiheesta Mitä pappi voi tehdä talouden parantamiseksi? Muutamaa vuotta myöhemmin ilmestyi väitöskirja papiston mahdollisuuksista pastoraalilääketieteen alalla. Molemmat julkaisut käsittelivät papiston yhteiskunnallista tehtävää. Pehr Kalmin mukaan papisto saattoi toiminnallaan näyttää hyvää esimerkkiä seurakuntalaisilleen. Tarkoituksena oli, että säätyläiset, joihin papistokin kuului, olisivat itse kokeilleet tiloillaan uudenlaisia viljelymenetelmiä. Nähdessään pappiensa yritysten onnistuvan, Kalm uskoi talonpoikien seuraavan näiden esimerkkiä. Kalm toivoi, että talonpojat olisivat siten luopuneet valtakunnan taloutta uhkaavista vääränlaisista viljelymenetelmistä. Erityisesti kaskeamisen uskottiin uhkaavan valtakunnan suurinta resurssia, metsiä. Kalmin ajatukset papiston yhteiskunnallisesta tehtävästä perustuivat Johannes Browalliuksen ja Carl Linnæuksen aikaisempiin kirjoituksiin. Ruotsin valtakunta oli menettänyt suurvalta-asemansa Suuressa Pohjan sodassa. Vapaudenajalla poliittisen suurvalta-aseman sijaan ryhdyttiin tavoittelemaan taloudellista valtaa. Aseet taottiin englantilaisen fysiokratismin hengessä auroiksi. Taloudellisen nousun edellytyksenä oli valtakunnan omavaraisuus. Ruotsin uskottiin olevan luonnonvarojensa puolesta poikkeuksellisen rikas maa. Näiden luonnonvarojen selvittäminen edellytti luonnontieteellistä tutkimista. Tämä johti tieteelliseen murrokseen, jonka tuloksena valtakunnantaloudellista hyötyä edistävät luonnontieteet nousivat Carl Linnæuksen ja Kuninkaallisen Tiedeakatemian johdolla kukoistukseen. Luonnontieteisiin kuului myös "jumalainen talousoppi". Taloudelliset uudistukset olivat ennen kaikkea uuden valtiopäiväpuolueen, hattujen ideologian mukaisia. Uutta aatevirtausta voidaan nimittää hyötypatriotismiksi. Pehr Kalm kuului hyötypatrioottien joukkoon. Hänen merkittävin tieteellinen saavutuksensa oli Tiedeakatemian tuella tehty tutkimusmatka Pohjois-Amerikkaan. Matkan tavoitteena oli silkinviljelyn aloittaminen Ruotsissa. Matkan jälkeen Kalm toimi Turun akatemian talousopinprofessorina. Vuonna 1757 hänet vihittiin papiksi. Kalm valitsi papin toimen nähtävästi taloudellisten syiden vuoksi. Kalm sai palkkapitäjästä tarvitsemansa lisätulot. Kalm oli myös Turun tuomiokapitulin jäsen. Kalmilla oli papiston yhteiskunnallisesta tehtävästä selkeä käsitys, joka näkyi paitsi hänen opetuksessaan myös hänen omassa työssään kirkkoherrana. Papin tehtävänä oli valtakunnan taloudellisen hyödyn edistäminen. Tässä mielessä papiston yhteiskunnallinen tehtävä ei lainkaan muuttunut suurvaltakaudelta vapaudenajalle siirryttäessä. Molempina aikoina keskusvalta määräsi tahdit, joiden mukaan papiston oli marssittava. Avainsanat: oppihistoria : Ruotsi : 1700-luku - valistus : papit - hyödyn aikakausi : papit - Pehr Kalm

Essays on Illegal Activities, Institutions and Economic Growth

In public economics, two extremist views on the functions of a government compete: one emphasizes government working for the public interest to provide value for the citizens, while another regards government mainly as a workhorse for private interests. Moreover, as the sole legitimate authority, the government has the right to define the rules and laws as well as to enforce them. With respect to regulation, two extremes arise: from too little regulation to too much of it. If the government does not function or ceases to exist, the state falls into anarchy or chaos (Somalia). If it regulates too much, it will completely suffocate private activities, which might be considered extralegal (the former Soviet Union). In this thesis I scrutinize the government s interventionist policies and evaluate the question of how to best promote economic well-being. The first two essays assume that the government s policies promote illegal activity. The first paper evaluates the interaction between the government and the mafia, and pays attention to the law enforcement of underground production. We show that the revenue-maximizing government will always monitor the shadow economy, as monitoring contributes to the government s revenue. In general, both legal and illegal firms are hurt by the entry of the mafia. It is, however, plausible that legal firms might benefit by the entry of the mafia if it competes with the government. The second paper tackles the issue of the measurement of the size of the shadow economy. To formulate policies it is essential to know what drives illegal economic activity; is it the tax burden, excess regulation, corruption or a weak legal environment? In this paper we propose an additional explanation for tax evasion and shadow production, namely cultural factors as manifested by religion as determinants of tax morality. According to our findings, Catholic and Protestant countries do not differ in their tax morale. The third paper contributes to the literature discussing the role of the government in promoting economic and productivity growth. Our main result is that, given the complex relationship between economic growth and economic freedom, marketization has not necessarily been beneficial in terms of growth. The last paper builds on traditional growth literature and revisits the debate on convergence clubs arising from demographic transition. We provide new evidence against the idea that countries within a club would converge over time. Instead, we propose that since the demographic transition is a dynamic process, one can expect countries to enter the last regime of stable, modern growth in stages.

Studies on the international effects of real shocks

This doctoral thesis addresses the macroeconomic effects of real shocks in open economies in flexible exchange rate regimes. The first study of this thesis analyses the welfare effects of fiscal policy in a small open economy, where private and government consumption are substitutes in terms of private utility. The main findings are as follows: fiscal policy raises output, bringing it closer to its efficient level, but is not welfare-improving even though government spending directly affects private utility. The main reason for this is that the introduction of useful government spending implies a larger crowding-out effect on private consumption, when compared with the `pure waste' case. Utility decreases since one unit of government consumption yields less utility than one unit of private consumption. The second study of this thesis analyses the question of how the macroeconomic effects of fiscal policy in a small open economy depend on optimal intertemporal behaviour. The key result is that the effects of fiscal policy depend on the size of the elasticity of substitution between traded and nontraded goods. In particular, the sign of the current account response to fiscal policy depends on the interplay between the intertemporal elasticity of aggregate consumption and the elasticity of substitution between traded and nontraded goods. The third study analyses the consequences of productive government spending on the international transmission of fiscal policy. A standard result in the New Open Economy Macroeconomics literature is that a fiscal shock depreciates the exchange rate. I demonstrate that the response of the exchange rate depends on the productivity of government spending. If productivity is sufficiently high, a fiscal shock appreciates the exchange rate. It is also shown that the introduction of productive government spending increases both domestic and foreign welfare, when compared with the case where government spending is wasted. The fourth study analyses the question of how the international transmission of technology shocks depends on the specification of nominal rigidities. A growing body of empirical evidence suggests that a positive technology shock leads to a temporary decline in employment. In this study, I demonstrate that the open economy dimension can enhance the ability of sticky price models to account for the evidence. The reasoning is as follows. An improvement in technology appreciates the nominal exchange rate. Under producer-currency pricing, the exchange rate appreciation shifts global demand toward foreign goods away from domestic goods. This causes a temporary decline in domestic employment.

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.

Essays on Misplanning Wealth and Health: A Behavioural Approach

The dissertation consists of three essays on misplanning wealth and health accumulation. The conventional economics assumes that individual's intertemporal preferences are exponential (exponential preferences, EP). Recent findings in behavioural economics have shown that, actually, people do discount near future relatively heavier than distant future. This implies hyperbolic intertemporal preferences (HP). Essays I and II concentrate especially on the effects of a delayed completion of tasks, a feature of behaviour that HP enables. Essay III uses current Finnish data to analyse the evolvement of the quality adjusted life years (QALYs) and inconsistencies in measuring that. Essay I studies the existence effects of a lucrative retirement savings program (SP) on the retirement savings of different individual types having HP. If the individual does not know that he will have HP also in the future, i.e. he is the naïve, for certain conditions, he delays the enrolment on SP until he abandons it. Very interesting finding is that the naïve retires then poorer in the presence than in the absence of SP. For the same conditions, the individual who knows that he will have HP also in the future, i.e. he is the sophisticated, gains from the existence of SP, and retires with greater retirement savings in the presence than in the absence of SP. Finally, capabilities to learn from past behaviour and about intertemporal preferences improve possibilities to gain from the existence but an adequate time to learn must be then guaranteed. Essay II studies delayed doctor's visits, theirs effects on the costs of a public health care system and government's attempts to control patient behaviour and fund the system. The controlling devices are a consultation fee and a deductible for that. The deductible is effective only for a patient whose diagnosis reveals a disease that would not get cured without the doctor's visit. The naives delay their visits the longest while EP-patients are the quickest visitors. To control the naives, the government should implement a low fee and a high deductible, while for the sophisticates the opposite is true. Finally, if all the types exist in an economy then using an incorrect conventional assumption that all individuals have EP leads to worse situation and requires higher tax rates than assuming incorrectly but unconventionally that only the naives exists. Essay III studies the development of QALYs in Finland 1995/96-2004. The essay concentrates on developing a consistent measure, i.e. independent of discounting, for measuring the age and gender specific QALY-changes and their incidences. For the given time interval, use of a relative change out of an attainable change seems to be almost intact to discounting and reveals that the greatest gains are for older age groups.

Participation or further exclusion? : Contestations over forest conservation and control in the East Usambara Mountains, Tanzania

This Ph.D. thesis Participation or Further Exclusion? Contestations over Forest Conservation and Control in the East Usambara Mountains, Tanzania describes and analyses the shift in the prevailing discourse of forest and biodiversity conservation policies and strategies towards more participatory approaches in Tanzania, and the changes in the practises of resource control. I explore the scope for and limits to the different actors and groups who are considered to form the community, to participate in resource control, in a specific historical and socio-economic context. I analyse whether, how and to which extent the targets of such participatory conservation interventions have been able to affect the formal rules and practices of resource control, and explore their different responses and discursive and other strategies in relation to conservation efforts. I approach the problematic through exploring certain participatory conservation interventions and related negotiations between the local farmers, government officials and the external actors in the case of two protected forest reserves in the southern part of the East Usambaras, Tanzania. The study area belongs to the Eastern Arc Mountains that are valued globally and nationally for their high level of biodiversity and number of endemic and near endemic species. The theoretical approach draws from theorising on power, participation and conservation in anthropology of development and post-structuralist political ecology. The material was collected in three stages between 2003 and 2008 by using an ethnographic approach. I interviewed and observed the actors and their resource use and control practices at the local level, including the representatives of the villagers living close to the protected forests and the conservation agency, but also followed the selected processes and engaged with the non-local agencies involved in the conservation efforts in the East Usambaras. In addition, the more recent processes of change and the actors strategies in resource control were contextualised against the social and environmental history of the study area and the evolvement of institutions of natural resource control. My findings indicate that the discourse of participation that has emerged in global conservation policy debate within the past three decades, and is being institutionalised in the national policies in many countries, including Tanzania, has shaped the practices of forest conservation in the East Usambaras, although in a fragmented and uneven way. Instrumental interpretation of participation, in which it is to serve the goals of improving the control of the forest and making it more acceptable and efficient, has prevailed among the governmental actors and conservation organisations. Yet, there is variation between the different projects and actors promoting participatory conservation regarding the goals and means of participation, e.g. to which extent the local people are to be involved in decision-making. The actors representing communities also have their diverse agendas, understandings and experiences regarding the rationality, outcomes and benefits of being involved in forest control, making the practices of control fluid. The elements of the exclusive conservation thinking and practices co-exist with the more recent participatory processes, and continue to shape the understandings and strategies of the actors involved in resource control. The ideas and narratives of the different discourses are reproduced and selectively used by the parties involved. The idea of forest conservation is not resisted as such by most of the actors at local level, quite the opposite. However, the strict regulations and rules governing access to resources, such as valuable timber species, continue to be disputed by many. Furthermore, the history of control, such as past injustices related to conservation and unfulfilled promises, undermines the participation of certain social groups in resource control and benefit sharing. This also creates controversies in the practices of conservation, and fuels conflicts regarding the establishment of new protected areas. In spite of this, the fact that the representatives of the communities have been invited to the arenas where information is shared, and principles and conditions of forest control and benefit sharing are discussed and partly decided upon, has created expectations among the participants, and opened up opportunities for some of the local actors to enhance their own, and sometimes wider interests in relation to resource control and the related benefits. The local actors experiences of the previous government and other interventions strongly affect how they position themselves in relation to conservation interventions, and their responses and strategies. However, my findings also suggest, in a similar way to research conducted in some other protected areas, that the benefits of participation in conservation and resource control tend to accrue unevenly between different groups of local people, e.g. due to unequal access to information and differences in their initial resources and social position.

Climate forcing on avian life history

In the 21st century, human-induced global climate change has been highlighted as one of the most serious threats to ecosystems worldwide. According to global climate scenarios, the mean temperature in Finland is expected to increase by 1.8 4.0°C by the end of the century. The regional and seasonal change in temperature has predicted to be spatially and temporally asymmetric, where the High-Arctic and Antarctic areas and winter and spring seasons have been projected to face the highest temperature increase. To understand how species respond to the ongoing climate change, we need to study how climate affects species in different phases of their life cycle. The impact of climate on breeding and migration of eight large-sized bird species was studied in this thesis, taking food availability into account. The findings show that climatic variables have considerable impact on the life-history traits of large-sized birds in northern Europe. The magnitude of climatic effects on migration and breeding was comparable with that of food supply, conventionally regarded as the main factor affecting these life-history traits. Based on the results of this thesis and the current climate scenarios, the following not mutually exclusive responses are possible in the near future. Firstly, asymmetric climate change may result in a mistiming of breeding because mild winters and early spring may lead to earlier breeding, whereas offspring are hatching into colder conditions which elevate mortality. Secondly, climate induced responses can differ between species with different breeding tactics (income vs. capital breeding), so that especially capital breeders can gain advantage on global warming as they can sustain higher energy resources. Thirdly, increasing precipitation has the potential to reduce the breeding success of many species by exposing nestlings to more severe post-hatching conditions and hampering the hunting conditions of parents. Fourthly, decreasing ice cover and earlier ice-break in the Baltic Sea will allow earlier spring migration in waterfowl. In eiders, this can potentially lead to more productive breeding. Fifthly, warming temperatures can favour parents preparing for breeding and increase nestling survival. Lastly, the climate-induced phenological changes in life history events will likely continue. Furthermore, interactions between climate and food resources can be complex and interact with each other. Eiders provide an illustrative example of this complexity, being caught in the crossfire between more benign ice conditions and lower salinity negatively affecting their prime food resource. The general conclusion is that climate is controlling not only the phenology of the species but also their reproductive output, thus affecting the entire population dynamics.

Harmful algae in the planktonic food web of the Baltic Sea

This study deals with algal species occurring commonly in the Baltic Sea: haptophyte Prymnesium parvum, dinoflagellates Dinophysis acuminata, D. norvegica and D. rotundata, and cyanobacterium Nodularia spumigena. The hypotheses are connected to the toxicity of the species, to the factors determining toxicity, to the consequences of toxicity and to the transfer of toxins in the aquatic food web. Since the Baltic Sea is severely eutrophicated, the fast-growing haptophytes have potential in causing toxic blooms. In our studies, the toxicity (as haemolytic activity) of the haptophyte P. parvum was highest under phosphorus-limited conditions, but the cells were toxic also under nitrogen limitation and under nutrient-balanced growth conditions. The cellular nutrient ratios were tightly related to the toxicity. The stoichiometric flexibility for cellular phosphorus quota was higher than for nitrogen, and nitrogen limitation led to decreased biomass. Negative allelopathic effects on another algae (Rhodomonas salina) could be observed already at low P. parvum cell densities, whereas immediate lysis of R. salina cells occurred at P. parvum cell densities corresponding to natural blooms. Release of dissolved organic carbon from the R. salina cells was measured within 30 minutes, and an increase in bacterial number and biomass was measured within 23 h. Because of the allelopathic effect, formation of a P. parvum bloom may accelerate after a critical cell density is reached and the competing species are eliminated. A P. parvum bloom indirectly stimulates bacterial growth, and alters the functioning of the planktonic food web by increasing the carbon transfer through the microbial loop. Our results were the first reports on DSP toxins in Dinophysis cells in the Gulf of Finland and on PTX-2 in the Baltic Sea. Cellular toxin contents in Dinophysis spp. ranged from 0.2 to 149 pg DTX-1 cell-1 and from 1.6 to 19.9 pg PTX-2 cell-1 in the Gulf of Finland. D. norvegica was found mainly around the thermocline (max. 200 cells L-1), whereas D. acuminata was found in the whole mixed layer (max. 7 280 cells L-1). Toxins in the sediment trap corresponded to 1 % of DTX-1 and 0.01 % PTX-2 of the DSP pool in the suspended matter. This indicates that the majority of the DSP toxins does not enter the benthic community, but is either decomposed in the water column, or transferred to higher trophic levels in the planktonic food chain. We found that nodularin, produced by Nodularia spumigena, was transferred to the copepod Eurytemora affinis through three pathways: by grazing on filaments of small Nodularia, directly from the dissolved pool, and through the microbial food web by copepods grazing on ciliates, dinoflagellates and heterotrophic nanoflagellates. The estimated proportion of the microbial food web in nodularin transfer was 22-45 % and 71-76 % in our two experiments, respectively. This highlights the potential role of the microbial food web in the transfer of toxins in the planktonic food web.

The Human UDP-Glucuronosyltransferases: Studies on Substrate Binding and Catalytic Mechanism

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Novel CDNF/MANF protein family : Molecular structure, expression and neurotrophic activity

Neurotrophic factors (NTFs) are secreted proteins which promote the survival of neurons, formation and maintenance of neuronal contacts and regulate synaptic plasticity. NTFs are also potential drug candidates for the treatment of neurodegenerative diseases. Parkinson’s disease (PD) is mainly caused by the degeneration of midbrain dopaminergic neurons. Current therapies for PD do not stop the neurodegeneration or repair the affected neurons. Thus, search of novel neurotrophic factors for midbrain dopaminergic neurons, which could also be used as therapeutic proteins, is highly warranted. In the present study, we identified and characterized a novel protein named conserved dopamine neurotrophic factor (CDNF), a homologous protein to mesencephalic astrocyte-derived neurotrophic factor (MANF). Others have shown that MANF supports the survival of embryonic midbrain dopaminergic neurons in vitro, and protects cultured cells against endoplasmic reticulum (ER) stress. CDNF and MANF form a novel evolutionary conserved protein family with characteristic eight conserved cysteine residues in their primary structure. The vertebrates have CDNF and MANF encoding genes, whereas the invertebrates, including Drosophila and Caenorhabditis have a single homologous CDNF/MANF gene. In this study we show that CDNF and MANF are secreted proteins. They are widely expressed in the mammalian brain, including the midbrain and striatum, and in several non-neuronal tissues. We expressed and purified recombinant human CDNF and MANF proteins, and tested the neurotrophic activity of CDNF on midbrain dopaminergic neurons using a 6-hydroxydopamine (6-OHDA) rat model of PD. In this model, a single intrastriatal injection of CDNF protected midbrain dopaminergic neurons and striatal dopaminergic fibers from the 6-OHDA toxicity. Importantly, an intrastriatal injection of CDNF also restored the functional activity of the nigrostriatal dopaminergic system when given after the striatal 6-OHDA lesion. Thus, our study shows that CDNF is a potential novel therapeutic protein for the treatment of PD. In order to elucidate the molecular mechanisms of CDNF and MANF activity, we resolved their crystal structure. CDNF and MANF proteins have two domains; an amino (N)-terminal saposin-like domain and a presumably unfolded carboxy (C)-terminal domain. The saposin-like domain, which is formed by five α-helices and stabilized by three intradomain disulphide bridges, may bind to lipids or membranes. The C-terminal domain contains an internal cysteine bridge in a CXXC motif similar to that of thiol/disulphide oxidoreductases and isomerases, and may thus facilitate protein folding in the ER. Our studies suggest that CDNF and MANF are novel potential therapeutic proteins for the treatment of neurodegenerative diseases. Future studies will reveal the neurotrophic and cytoprotective mechanisms of CDNF and MANF in more detail.

Cytokinin signalling in the regulation of cambial development

Secondary growth of plants is of pivotal importance in terrestrial ecosystems, providing a significant carbon sink in the form of wood. As plant biomass accumulation results largely from the cambial growth, it is surprising that quite little is known about the hormonal or genetic control of this important process in any plant species. The central aim of my thesis studies was to explore the function of cytokinin in the regulation of cambial development. Since their discovery as regulators of plant cell divisions, cytokinins have been assumed to participate in the control of cambial development. Evidence for this action was deduced from hormone treatment experiments, where exogenously applied cytokinin was shown to enhance cambial cell divisions in diverse plant organs and species. In my thesis work, the conservation of cytokinin signalling and homeostasis genes between a herbaceous plant, Arabidopsis, and a hardwood tree species, Populus trichocarpa. Presumably reflecting the ancient origin of cytokinin signalling system, the Populus genome contains orthologs for all Arabidopsis cytokinin signalling and homeostasis genes. Thus, genes belonging to five main families of isopentenyl transferases (IPTs), cytokinin oxidases (CKXs), two-component receptors, histidine containing phosphotransmitters (HPts) and response regulators (RRs) were identified from the Populus genome. Three subfamilies associated with cytokinin signal transduction, the CKI1-like family of two-component receptors, the AHP4-like HPts, and the ARR22-like atypical RRs, were significantly larger in Populus genome than in Arabidopsis. Potential contribution to the extensive secondary development of Populus by the members of these considerably expanded gene families will be discussed. Representatives of all cytokinin signal transduction elements were expressed in the Populus cambial zone, and most of the expressed genes appeared to be slightly more abundant on the phloem side of the meristem. The abundance of cytokinin related genes in the cambium emphasizes the important role of this hormone in the regulation of the extensive secondary growth characteristic of tree species. The function of the pseudo HPts in primary vascular development was studied in Arabidopsis root vasculature. It was demonstrated that the pseudo HPt AHP6 has a role in locally inhibiting cytokinin signalling in the protoxylem position in the Arabidopsis root, thus enabling differentiation of the protoxylem cell file. The possible role of pseudo HPts in cambial development will be discussed. The expression peak of cytokinin signalling genes in the tree cambial zone strongly indicates that cytokinin has a role in the regulation of this meristem function. To address whether cytokinin signalling is required for cambial activity, transgenic Populus trees with modified cytokinin signalling were produced. These trees were expressing a cytokinin catabolic gene from Arabidopsis, CYTOKININ OXIDASE 2, (AtCKX2) under the promoter of a Betula CYTOKININ RECEPTOR 1 (BpCRE1). The pBpCRE1::CKX2 transgenic Populus trees showed a reduced concentration of a biologically active cytokinin, correlating with their impaired cytokinin response. Furthermore, the radial growth of these trees was compromised, as illustrated by a smaller stem diameter than in wild-type trees of the same height. Moreover, the level of cambial cytokinin signalling was down-regulated in these thin-stemmed trees. The reduced signalling correlated with a decreased number of meristematic cambial cells, implicating cytokinin activity as a direct regulator of cambial cell division activity. Together, the results of my study indicate that cytokinins are major hormonal regulators required for cambial development.

Species-based and community-level approaches to conservation prioritization

One major reason for the global decline of biodiversity is habitat loss and fragmentation. Conservation areas can be designed to reduce biodiversity loss, but as resources are limited, conservation efforts need to be prioritized in order to achieve best possible outcomes. The field of systematic conservation planning developed as a response to opportunistic approaches to conservation that often resulted in biased representation of biological diversity. The last two decades have seen the development of increasingly sophisticated methods that account for information about biodiversity conservation goals (benefits), economical considerations (costs) and socio-political constraints. In this thesis I focus on two general topics related to systematic conservation planning. First, I address two aspects of the question about how biodiversity features should be valued. (i) I investigate the extremely important but often neglected issue of differential prioritization of species for conservation. Species prioritization can be based on various criteria, and is always goal-dependent, but can also be implemented in a scientifically more rigorous way than what is the usual practice. (ii) I introduce a novel framework for conservation prioritization, which is based on continuous benefit functions that convert increasing levels of biodiversity feature representation to increasing conservation value using the principle that more is better. Traditional target-based systematic conservation planning is a special case of this approach, in which a step function is used for the benefit function. We have further expanded the benefit function framework for area prioritization to address issues such as protected area size and habitat vulnerability. In the second part of the thesis I address the application of community level modelling strategies to conservation prioritization. One of the most serious issues in systematic conservation planning currently is not the deficiency of methodology for selection and design, but simply the lack of data. Community level modelling offers a surrogate strategy that makes conservation planning more feasible in data poor regions. We have reviewed the available community-level approaches to conservation planning. These range from simplistic classification techniques to sophisticated modelling and selection strategies. We have also developed a general and novel community level approach to conservation prioritization that significantly improves on methods that were available before. This thesis introduces further degrees of realism into conservation planning methodology. The benefit function -based conservation prioritization framework largely circumvents the problematic phase of target setting, and allowing for trade-offs between species representation provides a more flexible and hopefully more attractive approach to conservation practitioners. The community-level approach seems highly promising and should prove valuable for conservation planning especially in data poor regions. Future work should focus on integrating prioritization methods to deal with multiple aspects in combination influencing the prioritization process, and further testing and refining the community level strategies using real, large datasets.

Ecological consequences of genetic modifications - an invasion analysis approach

Biological invasions are considered as one of the greatest threats to biodiversity, as they may lead to disruption and homogenization of natural communities, and in the worst case, to native species extinctions. The introduction of gene modified organisms (GMOs) to agricultural, fisheries and forestry practices brings them into contact with natural populations. GMOs may appear as new invasive species if they are able to (1) invade into natural habitats or (2) hybridize with their wild relatives. The benefits of GMOs, such as increased yield or decreased use of insecticides or herbicides in cultivation, may thus be reduced due the potential risks they may cause. A careful ecological risk analysis therefore has to precede any responsible GMO introduction. In this thesis I study ecological invasion in relation to GMOs, and what kind of consequences invasion may have in natural populations. A set of theoretical models that combine life-history evolution, population dynamics, and population genetics were developed for the hazard identification part of ecological risks assessment of GMOs. In addition, the potential benefits of GMOs in management of an invasive pest were analyzed. In the first study I showed that a population that is fluctuating due to scramble-type density dependence (due to, e.g., nutrient competition in plants) may be invaded by a population that is relatively more limited by a resource (e.g., light in plants) that is a cause of contest-type density dependence. This result emphasises the higher risk of invasion in unstable environments. The next two studies focused on escape of a growth hormone (GH) transgenic fish into a natural population. The results showed that previous models may have given too pessimistic a view of the so called Trojan gene -effect, where the invading genotype is harmful for the population as a whole. The previously suggested population extinctions did not occur in my studies, since the changes in mating preferences caused by the GH-fish were be ameliorated by decreased level of competition. The GH-invaders may also have to exceed a threshold density before invasion can be successful. I also showed that the prevalence of mature parr (aka. sneaker) strategy among GH-fish may have clear effect on invasion outcome. The fourth study assessed the risks and developed methods against the invasion of the Colorado Potato Beetle (CPB, Leptinotarsa decemlineata). I showed that the eradication of CPB is most important for the prevention of their establishment, but the cultivation of transgenic Bt-potato could also be effective. In general, my results emphasise that invasion of transgenic species or genotypes to be possible under certain realistic conditions and resulting in competitive exclusion, population decline through outbreeding depression and genotypic displacement of native species. Ecological risk assessment should regard the decline and displacement of the wild genotype by an introduced one as a consequence that is as serious as the population extinction. It will also be crucial to take into account different kinds of behavioural differences among species when assessing the possible hazards that GMOs may cause if escaped. The benefits found of GMO crops effectiveness in pest management may also be too optimistic since CPB may evolve resistance to Bt-toxin. The models in this thesis could be further applied in case specific risk assessment of GMOs by supplementing them with detailed data of the species biology, the effect of the transgene introduced to the species, and also the characteristics of the populations or the environments in the risk of being invaded.