Oppilaan oma uskonto perusopetuksen uskonnon opetussuunnitelman perusteissa

Vuonna 2003 voimaan tulleen uuden uskonnonvapauslain muutosprosessin yhteydessä tehtiin muutoksia myös uskonnonopetusta määrittäviin lakeihin. Aikaisemmin uskonnonopetusta kuvannut termi ” oppilaan tunnustuksen mukainen” korvattiin käsitteellä ”oppilaan oman uskonnon opetus” (Lukiolaki 455/2003, Perusopetuslaki 454/2003). Tässä tutkimuksessa tarkasteltiin oppilaan omaa uskontoa perusopetuksen uskonnon opetussuunnitelmissa. Analyysi kohdistui kaikille uskontokuntasidonnaisille ryhmille yhteiseen opetussuunnitelman osaan sekä evanekelisluterilaisen, ortodoksisen, islamin ja Myöhempien Aikojen Jeesuksen Kristuksen kirkon uskonnon opetussuunnitelman perusteisiin. Tutkimus toteutettiin sisällön analyysin keinoin. -- Ensimmäisessä tutkimuskysymyksessä keskityttiin siihen, miten oma uskonto näyttäytyy opetussuunnitelman kaikille uskontosidonnaisille ryhmille yhteisessä osassa. Oppilaan oma uskonto ei siinä ole opetuksen päämäärää tai lähtökohtia määrittävässä asemassa. Omaan uskontoon tutustuminen määritetään yhdeksi opetuksen tavoitteista, jota ei kuitenkaan korosteta muihin tavoitteisiin nähden. Muita tavoitteita on esimerkiksi tutustuminen muihin uskontoihin ja uskonnon ymmärtäminen kulttuurisen ilmiön tasolla. Toisessa tutkimuskysymyksessä tarkasteltiin oppilaan oman uskonnon ilmenemistä uskontokohtaisissa opetussuunnitelmissa. Opetussuunnitelmia vertailtaessa huomattiin merkittäviä eroja siinä, millaisessa asemassa oma uskonto on opetussuunnitelman kokonaisuudessa. Vähemmistöuskontojen opetussuunnitelmissa oppilaan omaan uskontoon tutustuminen ilmenee opetuksen päämääränä. Evankelisluterilaisen uskonnon opetussuunnitelmassa sillä on välineellinen arvo pyrittäessä ymmärtämään uskontoa kulttuurisena ilmiönä. Oman uskonnon ilmenemistä opetussuunnitelmissa voi tutkimuksen perusteella hahmottaa juridisesta, kulttuurisesta ja identiteetin näkökulmasta. Merkittävimmät erot löytyivät identiteetin näkökulmasta hahmotetusta oppilaan omasta uskonnosta. Toisissa opetussuunnitelmissa oppilaan uskonnollinen identiteetti määritetään oppilaan juridisen oman uskonnon pohjalta. Toisissa suunnitelmissa oppilaan uskonnollista identiteettiä lähestytään individualisoituneen uskonnollisuuden näkökulmasta ja oppilaan henkilökohtainen uskonnollinen identiteetti jätetään oppilaan oman määrittämisen varaan. Kolmannessa tutkimuskysymyksessä keskityttiin siihen, minkä eri ulottuvuuksien kautta omaa uskontoa opetuksessa käsitellään. Aineistosta hahmotettiin viisi ulottuvuutta: instituution, rituaalinen, opillinen, narratiivinen ja eettinen ulottuvuus. Ulottuvuuksien painotuksiin eri uskontojen opetussuunnitelmissa voi nähdä vaikuttavan esimerkiksi uskonnon asema Suomessa ja uskonnon itseymmärrys uskonnon keskeisistä piirteistä sekä se, kuinka tiiviisti koulun uskonnonopetus on yhteydessä uskonnolliseen yhdyskuntaan. -- Tutkimus lisää tietoa uskonnon opetussuunnitelmien perusteista ja eri uskontojen suunnitelmien välisistä eroista. Tutkimuksen perusteella voidaan huomata, että eri opetussuunnitelmissa omalla uskonnolla on hyvin erilainen asema opetuksen lähtökohtia ja sisältöä määritettäessä. Toisissa opetussuunnitelmissa omaan uskontoon tutustuminen näyttäytyy opetuksen päämääränä ja lähtökohtana, mutta toisissa opetussuunnitelmissa opetuksen lähtökohdat kumpuavat uskontotieteellisestä ja individualisoituneen uskonnollisuuden viitekehyksestä. Näin ollen kouluissa annettavaa uskonnonopetusta ei voi pitää luonteeltaan yhdenmukaisena. Myös opetuksen kuvaaminen luonteeltaa ”oppilaan oman uskonnon opetuksena” näyttäytyy ristiriitaisena eri uskontojen opetussuunnitelmien valossa.

Uskon lyhyt oppimäärä : Suomen keskiaikaisten kivikirkkojen kalkkimaalausten välittämä raamattukuva

Tämä tutkielma on kokoava tutkimus Raamatusta nousevista kuva-aiheista, joita esiintyy 1400-luvun lopun ja 1510-luvun henkilömaalauksissa Suomen keskiaikaisissa kivikirkoissa. Aikaisempi Suomen keskiaikaista kirkkotaidetta koskeva tutkimus on ollut pääosin taidehistoriallista. Tutkielma pyrkii vastaamaan kysymykseen, millaisena Raamattu näyttäytyy suomalaisissa keskiaikaisissa kirkkomaalauksissa. Tutkimuskohteena ovat 1470–1490 –luvuilla Suomessa työskennelleen Taivassalon maalariryhmän maalaamat kirkot sekä 1500-luvun alussa maalatut kuvasarjat Länsi-Uudellamaalla, Hattulassa, Raumalla ja Pyhtäällä. Vanhasta testamentista peräisin olevia kuva-aiheita on vain 1510-luvulla tehdyissä maalaussarjoissa, joissa ne toimivat Kristukseen liittyvänä vertauskuvina ja hänen toimintansa selittäjinä. Vanhasta testamentista Suomen keskiaikaisten kirkkojen seinille on päässyt vanhan liiton keskeisiä henkilöitä, kuten Mooses, Daniel, Iisak ja Simson. Pelastushistoriaan kiinteästi liittyvät luominen ja syntiinlankeemus esiintyvät maalaussarjoissa kirkkosalin seinällä. Sen sijaan Kristuksen esikuvat löytyvät kirkkojen kattoholveista. Neitsyt Mariaan liittyvistä kuva-aiheista valtaosa on peräisin muualta kuin Raamatusta. Maalauksissa, jotka pohjaavat Raamattuun, Maria liittyy kiinteästi poikaansa Jeesukseen. Maria on yhdistetty Jeesuksen elämän loppuun ja alkuun; syntymäkertomuksiin ja kärsimykseen, kuolemaan ja ylösnousemukseen. Maria-aiheita esiintyy sekä Taivassalon ryhmän maalauksissa että 1500-luvun maalaussarjoissa. Jeesuksen elämä on havainnollistettu kalkkimaalauksissa yksityiskohtaisesti, etenkin Jeesuksen kärsimyshistorian osalta. Vain harvassa kirkossa Jeesus ei ole päässyt lainkaan osaksi maalauskokonaisuutta. Maalauksista välittyy kuva Jeesuksesta, joka etsii aikuisen ihmisen uskoa. Niin ikään maalausten Jeesus on pikemminkin ihmiskuntaa pelastamaan tullut taivaallinen Kristus kuin mies Jeesus Nasaretilainen. Viimeistä tuomiota esittävät maalauskokonaisuudet sekä evankelistasymbolit poikkeavat hyvin vähän eri kirkoissa. Molempien aiheiden kuvaamistavat olivat siinä määrin vakiintuneet, että erot kirkkojen välillä selittyvät käytetyillä mallikirjoilla ja maalareiden taidoilla. Viimeinen tuomio ja evankelista-symbolit kuuluvat suosituimpien kuva-aiheiden joukkoon. Jeesuksen opetuslapset on useassa kirkossa esitetty niin sanottuna credo-sarjana sekä osana kirkon pyhimysjoukkoa. Käsitellyn aineiston pohjalta nousee kuva Raamatusta, joka on lyhennetty ja tiivistetty. Keskiaikaisiin kuvaraamattuihin ja hartauskirjoihin verrattuna suomalaisissa kalkkimaalauksissa pitäydytään pitkälle olennaisessa kertomuksessa, eikä Jeesuksen elämää turhan tähden lähdetä selostamaan vaikeaselkoisin typologioin. Kuvakertomukset ovat ajan hengen mukaisesti vahvasti Kristus-keskeisiä. Kuitenkin kokonaisuutena Vanha testamentti pääsee maalauksissa paremmin esille kuin Uusi testamentti, josta huomio kiinnittyy lähes yksinomaan evankeliumeihin.

Luonnollisen lain sisältö Herbert Olssonin Luther-tulkinnassa

Tutkielmassa selvitetään Herbert Olssonin käsitystä luonnollisen lain sisällöstä, sen suhteesta rakkauden vaatimukseen ja kultaiseen sääntöön. Lähteinä ovat Olssonin kaikki Lutheria käsittelevät tekstit, keskeisimpinä hänen väitöskirjansa Grundproblemet i Luthers socialetik sekä pääteoksensa Schöpfung, Vernunft und Gesetz in Luthers Theologie . Tutkielma jakaantuu johdannon jälkeen taustalukuun, kolmeen analyysilukuun ja loppukatsaukseen, jossa esitellään tutkimuksen tulokset. Taustaluvussa tehdään katsaus luonnollisen lain tulkintahistoriaan aikaisemmassa Luther-tutkimuksessa. Luvussa tulevat ilmi erot pohjoismaisen ja mannereurooppalaisen Luther-tutkimuksen välillä, edellisen painottaessa vahvasti luonnollista lakia, sen identtisyyttä ilmoitetun lain ja rakkauden lain kanssa sekä sen universaalisuutta kaiken moraalisen toiminnan ohjaajana. Tämän jälkeen, luvussa kolme tutkitaan systemaattisen analyysin avulla Olssonin tulkintaa Lutherin luonnonoikeuskäsityksistä sekä luonnollisen käsitteestä ja luodaan pohjaa seuraaville tutkimusluvuille. Luvuissa neljä ja viisi käsitellään luonnollista lakia sinällään, ensin Olssonin väitöskirjasta ja sen jälkeen hänen myöhemmästä tuotannostaan välittyvän kuvan kautta. Jo alusta asti Olssonin tulkintaa leimaa käsitys siitä, että luonnollinen laki kuuluu kaikille ihmisille, se on sisällöltään identtinen dekalogin ja muun ilmoitetun lain kanssa, ja että sen vaatimus voidaan tiivistää rakkauden vaatimukseksi, ts. kultaiseksi säännöksi. Kultainen sääntö koskee Olssonilla sekä suhdetta lähimmäiseen että Jumalaan, kun sen aiemmin ajateltiin liittyvän vain lähimmäissuhteeseen. Olsson ei kuitenkaan vedä kovin monia johtopäätöksiä siitä, kuinka kultainen sääntö suhteessa Jumalaan pitäisi ymmärtää. Ensisijaisesti luonnollisen lain mukaan eläminen vaikuttaa olevan Olssonin tulkinnassa samaa kuin Jumalan tahdon mukaan eläminen. Tästä seuraa kysymys, millainen on Jumalan tahto ja sitä kautta Jumalan luonto, johon luonnollisen lain oletetaan viittaavan. Tutkimuksessa tullaan siihen tulokseen, että Olsson ei oleta Jumalan luonnon olevan lahjoittava rakkaus. Myöskään luonnollinen laki ei vaadi ihmistä luopumaan hyvän tavoittelusta, vaan se juuri vaatii suuntautumista kohti hyvää. Näin ollen Olssonin tulkinnan taustalla ei ole lahjoittavan rakkauden malli luomakunnan järjestyksenä, vaan Augustinukselta ja skolastikoilta tuttu ordo caritatis -malli. Ihmisessä reaalisesti läsnä oleva, pyyteettömästi palveleva Kristus ei tästä syystä sovi Olssonin tulkintaan lainkaan, ja näin ollen hänen käsityksensä vanhurskaudesta on oltava lähinnä forenssinen, ei efektiivinen. Ylipäätään Jumalan ja ihmisen suhteessa on hänellä kyse tahdon suhteesta, ei ontologisesta Kristuksen ja ihmisen yhdistymisestä. Lahjoittavan rakkauden ja Jumala-suhteen ontologisen ulottuvuuden puuttumisen takia siis Olssonin Luther-kuva muodostuu voluntaristiseksi, Jumalan absoluuttista tahtoa korostavaksi. Toisaalta hänen tulkintansa saa myös skolastisia painotuksia, johtuen juuri hyvään suuntautuvan rakkauden korostamisesta. Kultainen sääntö vaikuttaa olevan enemmän vastavuoroisuuden sääntö kuin pyyteettömän rakkauden sääntö. Summa summarum: mainituista syistä johtuen luonnollisen lain, rakkauden vaatimuksen ja kultaisen säännön sisällöllinen ykseys jää Olssonilla hänen tavoitteistaan huolimatta saavuttamatta.

Evidence of Partisan Emphasis on EMU during 1994–1999 : Comparing Finnish Parties

This study explores the EMU stand taken by the major Finnish political parties from 1994 to 1999. The starting point is the empirical evidence showing that party responses to European integration are shaped by a mix of national and cross-national factors, with national factors having more explanatory value. The study is the first to produce evidence that classified party documents such as protocols, manifestos and authoritative policy summaries may describe the EMU policy emphasis. In fact, as the literature review demonstrates, it has been unclear so far what kind of stand the three major Finnish political parties took during 1994–1999. Consequently, this study makes a substantive contribution to understanding the factors that shaped EMU party policies, and eventually, the national EMU policy during the 1990s. The research questions addressed are the following: What are the main factors that shaped partisan standpoints on EMU during 1994–1999? To what extent did the policy debate and themes change in the political parties? How far were the policies of the Social Democratic Party, the Centre Party and the National Coalition Party shaped by factors unique to their own national contexts? Furthermore, to what extent were they determined by cross-national influences from abroad, and especially from countries with which Finland has a special relationship, such as Sweden? The theoretical background of the study is in the area of party politics and approaches to EU policies, and party change, developed mainly by Kevin Featherstone, Peter Mair and Richard Katz. At the same time, it puts forward generic hypotheses that help to explain party standpoints on EMU. It incorporates a large quantity of classified new material based on primary research through content analysis and interviews. Quantitative and qualitative methods are used sequentially in order to overcome possible limitations. Established content-analysis techniques improve the reliability of the data. The coding frame is based on the salience theory of party competition. Interviews with eight party leaders and one independent expert civil servant provided additional insights and improve the validity of the data. Public-opinion surveys and media coverage are also used to complete the research path. Four major conclusions are drawn from the research findings. First, the quantitative and the interview data reveal the importance of the internal influences within the parties that most noticeably shaped their EMU policies during the 1990s. In contrast, international events play a minor role. The most striking feature turned out to be the strong emphasis by all of the parties on economic goals. However, it is important to note that the factors manifest differences between economic, democratic and international issues across the three major parties. Secondly, it seems that the parties have transformed into centralised and professional organisations in terms of their EMU policy-making. The weight and direction of party EMU strategy rests within the leadership and a few administrative elites. This could imply changes in their institutional environment. Eventually, parties may appear generally less differentiated and more standardised in their policy-making. Thirdly, the case of the Social Democratic Party shows that traditional organisational links continue to exist between the left and the trade unions in terms of their EMU policy-making. Hence, it could be that the parties have not yet moved beyond their conventional affiliate organisations. Fourthly, parties tend to neglect citizen opinion and demands with regard to EMU, which could imply conflict between the changes in their strategic environment. They seem to give more attention to the demands of political competition (party-party relationships) than to public attitudes (party-voter relationships), which would imply that they have had to learn to be more flexible and responsive. Finally, three suggestions for institutional reform are offered, which could contribute to the emergence of legitimised policy-making: measures to bring more party members and voter groups into the policy-making process; measures to adopt new technologies in order to open up the policy-formation process in the early phase; and measures to involve all interest groups in the policy-making process.

Generations and turnout : The generational effect in electoral participation in Finland

The relationship between age and turnout has been curve-linear as electoral participation first increases with age, remains relatively stable throughout middle-age and then gradually declines as certain physical infirmities set in (see e.g. Milbrath 1965). Alongside this life-cycle effect in voting, recent pooled cross-sectional analyses (see e.g. Blais et al. 2004; Lyons and Alexander 2000) have shown that there is also a generational effect, referring to lasting differences in turnout between various age groups. This study firstly examines the extent to which the generational effect applies in the Finnish context. Secondly, it investigates the factors accounting for that effect. The first article, based on individual-level register data from the parliamentary elections of 1999, shows that turnout differences between the different age groups would be even larger if there were no differences in social class and education. The second article examines simultaneously the effects of age, generation and period in the Finnish parliamentary elections of 1975-2003 based on pooled data from Finnish voter barometers (N = 8,634). The results show that there is a clear life cycle, generational and period effect. The third article examines the role of political socialisation in accounting for generational differences in electoral participation. Political socialisation is defined as the learning process in which an individual adopts various values, political attitudes, and patterns of actions from his or her environment. The multivariate analysis, based on the Finnish national election study 2003 (N=1,270), indicated that if there were no differences in socialisation between the youngest and the older generations, the difference in turnout would be much larger than if only sex and socioeconomic factors are controlled for. The fourth article examines other possible factors related to generational effect in voting. The results mainly apply to the Finnish parliamentary elections of 2003 in which we have data available. The results show that the sense of duty by far accounts for the generational effect in voting. Political interest, political knowledge and non-parliamentary participation also narrowed the differences in electoral participation between the youngest and the second youngest generations. The implication of the findings is that the lower turnout among the current youth is not a passing phenomenon that will diminish with age. Considering voting a civic duty and understanding the meaning of collective action are both associated with the process of political socialisation which therefore has an important role concerning the generational effect in turnout.

How terrorist campaigns end : The Rode Jeugd in the Netherlands and the Symbionese Liberation Army in the United States

This study explores the decline of terrorism by conducting source-based case studies on two left-wing terrorist campaigns in the 1970s, those of the Rode Jeugd in the Netherlands and the Symbionese Liberation Army in the United States. The purpose of the case studies is to bring more light into the interplay of different external and internal factors in the development of terrorist campaigns. This is done by presenting the history of the two chosen campaigns as narratives from the participants’ points of view, based on interviews with participants and extensive archival material. Organizational resources and dynamics clearly influenced the course of the two campaigns, but in different ways. This divergence derives at least partly from dissimilarities in organizational design and the incentive structure. Comparison of even these two cases shows that organizations using terrorism as a strategy can differ significantly, even when they share ideological orientation, are of the same size and operate in the same time period. Theories on the dynamics of terrorist campaigns would benefit from being more sensitive to this. The study also highlights that the demise of a terrorist organization does not necessarily lead to the decline of the terrorist campaign. Therefore, research should look at the development of terrorist activity beyond the lifespan of a single organization. The collective ideological beliefs and goals functioned primarily as a sustaining force, a lens through which the participants interpreted all developments. On the other hand, it appears that the role of ideology should not be overstated. Namely, not all participants in the campaigns under study fully internalized the radical ideology. Rather, their participation was mainly based on their friendship with other participants. Instead of ideology per se, it is more instructive to look at how those involved described their organization, themselves and their role in the revolutionary struggle. In both cases under study, the choice of the terrorist strategy was not merely a result of a cost-benefit calculation, but an important part of the participants’ self-image. Indeed, the way the groups portrayed themselves corresponded closely with the forms of action that they got involved in. Countermeasures and the lack of support were major reasons for the decline of the campaigns. However, what is noteworthy is that the countermeasures would not have had the same kind of impact had it not been for certain weaknesses of the groups themselves. Moreover, besides the direct impact the countermeasures had on the campaign, equally important was how they affected the attitudes of the larger left-wing community and the public in general. In this context, both the attitudes towards the terrorist campaign and the authorities were relevant to the outcome of the campaigns.

Tiedon jakaminen tiimityössä

Knowledge-sharing in a teamwork The study examines the link between knowledge-sharing that takes place in a team and the dimensions and objectives of the team s activities. The question the study poses is: How does knowledge-sharing in a team relate to the team s activities? The exchange of knowledge is examined using knowledge-sharing networks and the conversion model, which describes the process of knowledge formation. The answer to the question is sought through four empirical articles describing the activities of a team from the viewpoint of quality, fairness, power related to knowledge management, and performance. One of the articles used in the study describes the role of networks in work life more generally. It attempts to shed light on the manner in which team-related networks operate as part of a more extensive structure of organizational networks. Finland is one of the most eager users of teamwork, if numbers are used as a yardstick. About half of all Finnish wage earners worked in teams in 2009, and comparisons show that the use of teams in Finland is above the EU average. This study focuses on so-called semi-autonomous teams, which carry out permanent work tasks. In such teams, tasks are interdependent, and teams are jointly responsible for ensuring that the work is done. Team members may also, at least to some extent, agree between themselves on how the tasks are carried out and are able to take part in the decision-making process. Such teamwork makes knowledge-sharing an important element for the team s activities. Knowledge and knowledge-sharing have become a major resource, allowing organizations to operate and even compete in today s increasingly competitive markets. A single team or a single organization cannot, however, possess all the knowledge required for carrying out the tasks assigned to it. Although it is difficult to copy the knowledge generated in an organization, it is important to share the knowledge within and between organizations. External links supply teams and organizations with important knowledge that allows them to keep their operations up-to-date and their structures well-functioning. In fact, knowledge provides teams and organizations with an intangible resource that improves their capacity to interact with their environment and to adjust to it. For this reason, it is important to examine both the internal and external knowledge-sharing taking place in a team. The findings of the study show that in terms of quality, fairness, performance and the knowledge management issues concerning a team, its social network structure is both internally and externally connected with its activities. A team structure that is internally coherent and at the same time open to external contacts, is, with certain restrictions, connected with the quality, fairness, and performance of the team. The restrictions concern differences between procedural and interactional justice, public and private sectors, and the team leaders and ordinary team members. The role of the team leader is closely connected with the management of networks that are considered valuable. The results of the study indicate that teamwork is supervisor-dominated. Thus, teamwork does not substantially strengthen the influence of individual employees as players in knowledge-transfer networks. However, ordinary team members possess important peer contacts inside the organization. Teamwork clearly allows employees to interact in a democratic manner, and here the transfer of tacit knowledge plays an important role. Keywords: teamwork, knowledge-sharing, social networks, organization

Context-dependent mate choice in the sand goby, Pomatoschistus minutus

The results presented in this thesis show that all females of a given population do not necessarily choose similar mating partners. Specifically, partner preferences of a fish, the sand goby (Pomatoschistus minutus), varied among individual females and depended on the social context at the time of choice. I also show that females assess multiple mate choice cues simultaneously; partner preferences were based more strongly on an interaction effect between different choice cues than on any individual cue. Furthermore, I found that preferred matings involved fitness benefits in the form of increased offspring success, but these benefits were not significantly affected by mate compatibility. Hence, mate choice for partner compatibility does not appear to be an important determinant of the observed variation in female mate preferences in this species. The context-dependency of female mating preferences revealed is relevant to how genetic variation in sexually selected traits might be maintained: as the mating success of a certain male type varies according to the choice context, directional sexual selection on male traits is shown to be less intense than generally thought making for a slower loss of genetic variation in these traits. Mating preferences of sand gobies were assessed by giving females a binary choice between males that differed in body size and/or other focus traits. These association preferences were found to be sexually motivated, repeatable and to correspond to actual mating decisions.

Phylogeography and Population Genetics of the Moor frog (Rana arvalis Nilsson) in Northern Europe

The ongoing climate change along with increasing levels of pollutants, diseases, habitat loss and fragmentation constitute global threats to the persistence of many populations, species and ecosystems. However, for the long-term persistence of local populations, one of the biggest threats is the intrinsic loss of genetic variation. In order to adapt to changes in the environment, organisms must have a sufficient supply of heritable variation in traits important for their fitness. With a loss of genetic variation, the risk of extinction will increase. For conservational practices, one should therefore understand the processes that shape the genetic population structure and also the broader (historical) phylogenetic patterning of the species in focus. In this thesis, microsatellite markers were applied to study genetic diversity and population differentiation of the protected moor frog (Rana arvalis) in Fennoscandia from both historical (evolutionary) and applied (conservation) perspectives. The results demonstrate that R. arvalis populations are highly structured over rather short geographic distances. Moreover, the results suggest that R. arvalis recolonized Fennoscandia from two directions after the last ice age. This has had implications for the genetic structuring and population differentiation, especially in the northernmost parts where the two lineages have met. Compared to more southern populations, the genetic variation decreases and the interpopulation differentiation increases dramatically towards north. This could be an outcome of serial population bottlenecking along the recolonization route. Also, current isolation and small population sizes increase the effect of drift, thus reinforcing the observed pattern. The same pattern can also be seen in island populations. However, though R. arvalis on the island of Gotland has lost most of its neutral genetic variability, our results indicate that the levels of additive genetic variation have remained high. This conforms to the conjecture that though neutral markers are widely used in conservation purposes, they may be quite uninformative about the levels of genetic variation in ecologically important traits. Finally, the evolutionary impact of the typical amphibian mating behaviour on genetic diversity was investigated. Given the short time available for larval development, it is important that mating takes place as early as possible. The genetic data and earlier capture-recapture data suggest that R. arvalis gather at mating grounds they are familiar with. However, by forming leks in random to relatedness, and having multiple paternities in single clutches, the risk of inbreeding may be minimized in this otherwise highly philopatric species.

Conservation of Atlantic salmon by supplementary stocking of juvenile fish

Wild salmon stocks in the northern Baltic rivers became endangered in the second half of the 20th century, mainly due to recruitment overfishing. As a result, supplementary stocking was widely practised, and supplementation of the Tornionjoki salmon stock took place over a 25 year period until 2002. The stock has been closely monitored by electrofishing, smolt trapping, mark-recapture studies, catch samples and catch surveys. Background information on hatchery-reared stocked juveniles was also collected for this study. Bayesian statistics was applied to the data as this method offers the possibility of bringing prior information into the analysis and an advanced ability for incorporating uncertainty, and also provides probabilities for a multitude of hypotheses. Substantial divergences between reared and wild Tornionjoki salmon were identified in both demographic and phenological characteristics. The divergences tended to be larger the longer the duration spent in hatchery and the more favourable the hatchery conditions were for fast growth. Differences in environment likely induced most of the divergences, but selection of brood fish might have resulted in genotypic divergence in maturation age of reared salmon. Survival of stocked 1-year old juveniles to smolt varied from about 10% to about 25%. Stocking on the lower reach of the river seemed to decrease survival, and the negative effect of stocking volume on survival raises the concern of possible similar effects on the extant wild population. Post-smolt survival of wild Tornionjoki smolts was on average two times higher than that of smolts stocked as parr and 2.5 times higher than that of stocked smolts. Smolts of different groups showed synchronous variation and similar long-term survival trends. Both groups of reared salmon were more vulnerable to offshore driftnet and coastal trapnet fishing than wild salmon. Average survival from smolt to spawners of wild salmon was 2.8 times higher than that of salmon stocked as parr and 3.3 times higher than that of salmon stocked as smolts. Wild salmon and salmon stocked as parr were found to have similar lifetime survival rates, while stocked smolts have a lifetime survival rate over 4 times higher than the two other groups. If eggs are collected from the wild brood fish, stocking parr would therefore not be a sensible option. Stocking smolts instead would create a net benefit in terms of the number of spawners, but this strategy has serious drawbacks and risks associated with the larger phenotypic and demographic divergences from wild salmon. Supplementation was shown not to be the key factor behind the recovery of the Tornionjoki and other northern Baltic salmon stocks. Instead, a combination of restrictions in the sea fishery and simultaneous occurrence of favourable natural conditions for survival were the main reasons for the revival in the 1990 s. This study questions the effectiveness of supplementation as a conservation management tool. The benefits of supplementation seem at best limited. Relatively high occurrences of reared fish in catches may generate false optimism concerning the effects of supplementation. Supplementation may lead to genetic risks due to problems in brood fish collection and artificial rearing with relaxed natural selection and domestication. Appropriate management of fisheries is the main alternative to supplementation, without which all other efforts for long-term maintenance of a healthy fish resource fail.

CADASIL: molecular studies on the most common hereditary vascular dementing disorder

Cerebral Autosomal Dominant Arteriopathy with Subcortical Infarcts and Leukoencephalopathy (CADASIL) is the most common hereditary vascular dementia. CADASIL is a systemic disease of small and medium-sized arteries although the symptoms are almost exclusively neurological, including migraineous headache, recurrent ischemic episodes, cognitive impairment and, finally, subcortical dementia. CADASIL is caused by over 170 different mutations in the NOTCH3 gene, which encodes a receptor expressed in adults predominantly in the vascular smooth muscle cells. The function of NOTCH3 is not crucial for embryonic development but is needed after birth. NOTCH3 directs postnatal arterial maturation and helps to maintain arterial integrity. It is involved in regulation of vascular tone and in the wound healing of a vascular injury. In addition, NOTCH3 promotes cell survival by inducing expression of anti-apoptotic proteins. NOTCH3 is a membrane-spanning protein with a large extracellular domain (N3ECD) containing 34 epidermal growth factor-like (EGF) repeats and a smaller intracellular domain with six ankyrin repeats. All CADASIL mutations are located in the EGF repeats and the majority of the mutations cause gain or loss of one cysteine residue in one of these repeats leading to an odd number of cysteine residues, which in turn leads to misfolding of N3ECD. This misfolding most likely alters the maturation, targetting, degradation and/or function of the NOTCH3 receptor. CADASIL mutations do not seem to affect the canonical NOTCH3 signalling pathway. The main pathological findings are the accumulation of the NOTCH3 extracellular domain on degenerating vascular smooth muscle cells (VSMCs), accumulation of granular osmiophilic material (GOM) in the close vicinity of VSMCs as well as fibrosis and thickening of arterial walls. Narrowing of the arterial lumen and local thrombosis cause insufficient blood flow, mainly in small arteries of the cerebral white matter, resulting in tissue damage and lacunar infarcts. CADASIL is suspected in patients with a suggestive family history and clinical picture as well as characteristic white matter alterations in magnetic resonance imaging. A definitive verification of the diagnosis can be achieved by identifying a pathogenic mutation in the NOTCH3 gene or through the detection of GOM by electron microscopy. To understand the pathology underlying CADASIL, we have generated a unique set of cultured vascular smooth muscle cell (VSMC) lines from umbilical cord, placental, systemic and cerebral arteries of CADASIL patients and controls. Analyses of these VSMCs suggest that mutated NOTCH3 is misfolded, thus causing endoplasmic reticulum stress, activation of the unfolded protein response and increased production of reactive oxygen species. In addition, mutation in NOTCH3 causes alterations in actin cytoskeletal structures and protein expression, increased branching and abnormal node formation. These changes correlate with NOTCH3 expression levels within different VSMCs lines, suggesting that the phenotypic differences of SMCs may affect the vulnerability of the VSMCs and, therefore, the pathogenic impact of mutated NOTCH3 appears to vary in the arteries of different locations. Furthermore, we identified PDGFR- as an immediate downstream target gene of NOTCH3 signalling. Activation of NOTCH induces up-regulation of the PDGFR- expression in control VSMCs, whereas this up-regulation is impaired in CADASIL VSMCs and might thus serve as an alternative molecular mechanism that contributes to CADASIL pathology. In addition, we have established the congruence between NOTCH3 mutations and electron microscopic detection of GOM with a view to constructing a strategy for CADASIL diagnostics. In cases where the genetic analysis is not available or the mutation is difficult to identify, a skin biopsy is an easy-to-perform and highly reliable diagnostic method. Importantly, it is invaluable in setting guidelines concerning how far one should proceed with the genetic analyses.

Human genetic variation in the Baltic Sea region: Features of population history and natural selection

In this thesis, the genetic variation of human populations from the Baltic Sea region was studied in order to elucidate population history as well as evolutionary adaptation in this region. The study provided novel understanding of how the complex population level processes of migration, genetic drift, and natural selection have shaped genetic variation in North European populations. Results from genome-wide, mitochondrial DNA and Y-chromosomal analyses suggested that the genetic background of the populations of the Baltic Sea region lies predominantly in Continental Europe, which is consistent with earlier studies and archaeological evidence. The late settlement of Fennoscandia after the Ice Age and the subsequent small population size have led to pronounced genetic drift, especially in Finland and Karelia but also in Sweden, evident especially in genome-wide and Y-chromosomal analyses. Consequently, these populations show striking genetic differentiation, as opposed to much more homogeneous pattern of variation in Central European populations. Additionally, the eastern side of the Baltic Sea was observed to have experienced eastern influence in the genome-wide data as well as in mitochondrial DNA and Y-chromosomal variation – consistent with linguistic connections. However, Slavic influence in the Baltic Sea populations appears minor on genetic level. While the genetic diversity of the Finnish population overall was low, genome-wide and Y-chromosomal results showed pronounced regional differences. The genetic distance between Western and Eastern Finland was larger than for many geographically distant population pairs, and provinces also showed genetic differences. This is probably mainly due to the late settlement of Eastern Finland and local isolation, although differences in ancestral migration waves may contribute to this, too. In contrast, mitochondrial DNA and Y-chromosomal analyses of the contemporary Swedish population revealed a much less pronounced population structure and a fusion of the traces of ancient admixture, genetic drift, and recent immigration. Genome-wide datasets also provide a resource for studying the adaptive evolution of human populations. This study revealed tens of loci with strong signs of recent positive selection in Northern Europe. These results provide interesting targets for future research on evolutionary adaptation, and may be important for understanding the background of disease-causing variants in human populations.

Lignin biosynthesis in Norway spruce: from a model system to the tree

Lignin is a hydrophobic polymer that is synthesised in the secondary cell walls of all vascular plants. It enables water conduction through the stem, supports the upright growth habit and protects against invading pathogens. In addition, lignin hinders the utilisation of the cellulosic cell walls of plants in pulp and paper industry and as forage. Lignin precursors are synthesised in the cytoplasm through the phenylpropanoid pathway, transported into the cell wall and oxidised by peroxidases or laccases to phenoxy radicals that couple to form the lignin polymer. This study was conducted to characterise the lignin biosynthetic pathway in Norway spruce (Picea abies (L.) Karst.). We focused on the less well-known polymerisation stage, to identify the enzymes and the regulatory mechanisms that are involved. Available data for lignin biosynthesis in gymnosperms is scarce and, for example, the latest improvements in precursor biosynthesis have only been verified in herbaceous plants. Therefore, we also wanted to study in detail the roles of individual gene family members during developmental and stress-induced lignification, using EST sequencing and real-time RT-PCR. We used, as a model, a Norway spruce tissue culture line that produces extracellular lignin into the culture medium, and showed that lignin polymerisation in the tissue culture depends on peroxidase activity. We identified in the culture medium a significant NADH oxidase activity that could generate H2O2 for peroxidases. Two basic culture medium peroxidases were shown to have high affinity to coniferyl alcohol. Conservation of the putative substrate-binding amino acids was observed when the spruce peroxidase sequences were compared with other peroxidases with high affinity to coniferyl alcohol. We also used different peroxidase fractions to produce synthetic in vitro lignins from coniferyl alcohol; however, the linkage pattern of the suspension culture lignin could not be reproduced in vitro with the purified peroxidases, nor with the full complement of culture medium proteins. This emphasised the importance of the precursor radical concentration in the reaction zone, which is controlled by the cells through the secretion of both the lignin precursors and the oxidative enzymes to the apoplast. In addition, we identified basic peroxidases that were reversibly bound to the lignin precipitate. They could be involved, for example, in the oxidation of polymeric lignin, which is required for polymer growth. The dibenzodioxocin substructure was used as a marker for polymer oxidation in the in vitro polymerisation studies, as it is a typical substructure in wood lignin and in the suspension culture lignin. Using immunolocalisation, we found the structure mainly in the S2+S3 layers of the secondary cell walls of Norway spruce tracheids. The structure was primarily formed during the late phases of lignification. Contrary to the earlier assumptions, it appears to be a terminal structure in the lignin macromolecule. Most lignin biosynthetic enzymes are encoded for by several genes, all of which may not participate in lignin biosynthesis. In order to identify the gene family members that are responsible for developmental lignification, ESTs were sequenced from the lignin-forming tissue culture and developing xylem of spruce. Expression of the identified lignin biosynthetic genes was studied using real-time RT-PCR. Candidate genes for developmental lignification were identified by a coordinated, high expression of certain genes within the gene families in all lignin-forming tissues. However, such coordinated expression was not found for peroxidase genes. We also studied stress-induced lignification either during compression wood formation by bending the stems or after Heterobasidion annosum infection. Based on gene expression profiles, stress-induced monolignol biosynthesis appeared similar to the developmental process, and only single PAL and C3H genes were specifically up-regulated by stress. On the contrary, the up-regulated peroxidase genes differed between developmental and stress-induced lignification, indicating specific responses.

Molecular Alterations in Asbestos-Related Lung Cancer

Background: Asbestos is a well known cancer-causing mineral fibre, which has a synergistic effect on lung cancer risk in combination with tobacco smoking. Several in vitro and in vivo experiments have demonstrated that asbestos can evoke chromosomal damage and cause alterations as well as gene expression changes. Lung tumours, in general, have very complex karyotypes with several recurrently gained and lost chromosomal regions and this has made it difficult to identify specific molecular changes related primarily to asbestos exposure. The main aim of these studies has been to characterize asbestos-related lung cancer at a molecular level. Methods: Samples from asbestos-exposed and non-exposed lung cancer patients were studied using array comparative genomic hybridization (aCGH) and fluorescent in situ hybridization (FISH) to detect copy number alterations (CNA) as well as microsatellite analysis to detect allelic imbalance (AI). In addition, asbestos-exposed cell lines were studied using gene expression microarrays. Results: Eighteen chromosomal regions showing differential copy number in the lung tumours of asbestos-exposed patients compared to those of non-exposed patients were identified. The most significant differences were detected at 2p21-p16.3, 5q35.3, 9q33.3-q34.11, 9q34.13-q34.3, 11p15.5, 14q11.2 and 19p13.1-p13.3 (p<0.005). The alterations at 2p and 9q were validated and characterized in detail using AI and FISH analysis in a larger study population. Furthermore, in vitro studies were performed to examine the early gene expression changes induced by asbestos in three different lung cell lines. The results revealed specific asbestos-associated gene expression profiles and biological processes as well as chromosomal regions enriched with genes believed to contribute to the common asbestos-related responses in the cell lines. Interestingly, the most significant region enriched with asbestos-response genes was identified at 2p22, close to the previously identified region showing asbestos-related CNA in lung tumours. Additionally, in this thesis, the dysregulated biological processes (Gene Ontology terms) detected in the cell line experiment were compared to dysregulated processes identified in patient samples in a later study (Ruosaari et al., 2008a). Commonly affected processes such as those related to protein ubiquitination, ion transport and surprisingly sensory perception of smell were identified. Conclusions: The identification of specific CNA and dysregulated biological processes shed some light on the underlying genes acting as mediators in asbestos-related lung carcinogenesis. It is postulated that the combination of several asbestos-specific molecular alterations could be used to develop a diagnostic method for the identification of asbestos-related lung cancer.

Benthic-pelagic coupling in the northern Baltic Sea: importance of bioturbation and benthic predation

Benthic-pelagic coupling describes processes that operate across and between the seafloor and open-water ecosystems. In soft-sediment communities, bioturbation by sediment-dwelling and epibenthic organisms may strongly shape habitat characteristics and influence processes, e.g. biogeochemical cycling, which supplies bioavailable nutrients to pelagic primary producers. In addition, benthic fauna may mediate benthic-pelagic coupling by affecting the survival and hatching of zooplankton dormant eggs in the sediment. In the shallow waters and seasonally fluctuating environment of the Baltic Sea, emergence from the seafloor essentially contributes to the dynamics of zooplankton pelagic populations. In this thesis, I examine how benthic organisms with different functional traits affect the link between the benthic and pelagic systems in the northern Baltic Sea. By means of experimental laboratory studies, the effects of sediment-dwelling (Monoporeia affinis, Macoma balthica and Marenzelleria spp.) and nectobenthic (Mysis spp.) taxa on the survival and hatching of zooplankton benthic eggs and on benthic nutrient fluxes and sediment structure were investigated. In the predation studies, the nectobenthic mysids Mysis spp. preyed upon benthic eggs of the cladoceran Bosmina longispina maritima (syn. B. coregoni maritima), both in pelagic and benthic environments. Of the sediment-dwelling species, the amphipod M. affinis and the bivalve M. balthica reduced the number of cladoceran eggs in the sediment, whereas the polychaetes Marenzelleria spp. had no effects on cladoceran eggs. Both M. balthica and M. affinis also increased the mortality rates of benthic eggs of copepods and rotifers. It was estimated that zooplankton eggs provide an additional carbon source for food-limited benthic communities. The results indicate that predation pressure on zooplankton benthic eggs may be strong, but varies widely depending on the season and the functional characteristics of the macrofauna. Macoma balthica buried cladoceran eggs and a fluorescent tracer from the sediment surface to a depth of 3 4 cm, indicating efficient sediment mixing. In contrast, the other taxa had fewer effects on particle distributions. In addition to organic matter mineralization, particle mixing is crucial to the success of benthic recruitment of zooplankton, since only eggs close to the sediment surface may hatch. Macoma balthica and M. affinis altered the patterns of zooplankton emergence from the sediment. In general, the highest emergence rates were observed in the absence of macroscopic fauna, and M. balthica exerted a stronger suppressive effect than M. affinis. Moreover, copepods were less severely affected than cladocerans, while only one species (Temora longicornis) clearly benefited from the presence of the macrofauna. These differences probably result from species-specific differences in the resistance of eggs to disturbances. The results show that benthic fauna may considerably alter the patterns of zooplankton emergence from the seafloor, thereby shaping zooplankton pelagic populations. The semi-motile M. balthica and Marenzelleria spp. increased the fluxes of phosphate and ammonium from the sediment to the water, whereas the motile M. affinis and Mysis mixta had a contrasting effect. In the eutrophied Baltic Sea, efficient internal cycling of bioavailable nutrients forms a strong feedback inhibiting the recovery of the ecosystem. Based on the results, a change in species dominance from the two motile taxa, susceptible to oxygen deficiency, to the more tolerant semi-motile taxa provides additional feedback, strengthening internal nutrient cycling and accelerating eutrophication, with deteriorating near-bottom oxygen conditions and changes in the benthic communities. In shallow-water ecosystems, benthic nutrient regeneration plays a key role in determining the overall productivity of the ecosystem. In addition, the results of this study show that the communities in the benthos may essentially contribute to the structure of those in the plankton.