Genetic diversity of Atlantic salmon (Salmo salar) and European whitefish (Coregonus lavaretus) in the Baltic Sea

We described the patterns and extent of microsatellite DNA variation in historical and present-day Atlantic salmon (Salmo salar L.) stocks in the Baltic Sea and neighbouring areas, and in European whitefish (Coregonus lavaretus) ecotypes, populations and run-timing types in Finland. Moreover, the amount and pattern of genetic diversity in historical salmon populations before human impact were described, and the proportion of diversity maintained in the present hatchery stocks evaluated. Salmon populations in the Baltic Sea were, on average, significantly less variable than eastern Atlantic populations, and the diversity of landlocked populations (Lakes Vänern, Saimaa, Onega and Ladoga) was in turn significantly lower than that of anadromous salmon populations in the Baltic Sea populations. Within the Baltic Sea, the anadromous populations of Atlantic salmon formed three clear groups, corresponding to the northern (Gulf of Bothnia), eastern (Gulf of Finland and eastern Baltic Main Basin) and southern (western Baltic Main Basin) regions. Based on microsatellite data, three salmon population groups in the Baltic Sea were considered potentially different colonization lineages. In short- and long-term breeding programmes of Atlantic salmon, the average observed rate of loss of alleles was 4.9% and 2.0% per generation and the average rate of loss of heterozygosity was 1.4% and 1% per generation, respectively. When comparing the genetic parameters of stocks before and after hatchery breeding of several successive generations (Rivers Iijoki and Oulujoki), statistically significant changes in allele frequencies were common, while large wild stock in the Teno River has remained temporally very stable over 56 years. Despite the observed losses of genetic diversity in broodstock breeding, a large proportion of the genetic resources of the extirpated stocks are still conserved in the broodstocks. Genetic differentiation among European whitefish ecotypes was generally low, thus giving support to the hypothesis of one native European whitefish species in Fennoscandia. Among the ecotypes, the northern, large sparsely rakered, bottom-dwelling whitefish was the most unique. The known genetic differences in quantitative traits have thus either developed independently of potential phylogenetic lineages, or the lineages have mixed and the quantitative traits of the ecotypes, like gill-raker number, have later changed according to environment and selection pressures. Overall, genetic distances between the anadromous whitefish populations along the Finnish coast, especially in the Bothnian Bay area, were small. Wild whitefish populations studied had slightly higher allelic diversity than hatchery-reared populations in corresponding rivers.

From the river to the open sea - a critical life phase of young Atlantic salmon migrating from the Simojoki river

Long-term monitoring data collected from wild smolts of Atlantic salmon (Salmo salar) in the Simojoki river, northern Finland, were used in studying the relationships between the smolt size and age, smolt and postsmolt migration, environmental conditions and postsmolt survival. The onset of the smolt run was significantly dependent on the rising water temperature and decreasing discharge of the river in the spring. The mean length of smolts migrating early in the season was commonly higher and the mean age always older than among smolts migrating later. Many of the smolts migrating early in the season and almost all smolts migrating later had started their new growth in spring in the river before their sea entry. Among postsmolts, the time required for emigration from the estuary was dependent on the sea surface temperature (SST) off the river, being significantly shorter in years with warm than cold sea temperatures. After leaving the estuary, the postsmolts migrated southwards along the eastern coast of the northern Gulf of Bothnia, the geographical distribution of the tag recoveries coinciding with the warm thermal zone in spring in the coastal area. After arriving in the southern Gulf of Bothnia in late summer the postsmolts mostly migrated near the western coast, reaching the Baltic Main Basin in late autumn. Until the early 1990s there was only a weak positive association between smolt length and postsmolt survival. However, following a subsequent decrease in the mean smolt size, a significant positive dependence was observed between smolt size and the reported recapture rate of tagged salmon. The differences in recapture rates between smolts tagged during the first and second half of the annual migration season were insignificant, indicating that the seasonal variation in smolt size and age seem to be too small to affect survival. Among the climatic factors examined, the summer SST in the Gulf of Bothnia was most clearly related to the survival of the wild postsmolts. Postsmolt survival appeared to be highest in years when the SST in June in the Bothnian Bay varied between 9 and 12 ºC. In addition, the survival of wild postsmolts showed a significant positive dependence on the SST in July in the Bothnian Sea, but not on the abundance of the prey fish (0+ herring, Clupea harengus and sprat, Sprattus sprattus) in the Bothnian Sea and in the Baltic Main Basin. The results suggest, that if the incidence of extreme weather conditions were to increase due to climatic changes, it would probably reduce the postsmolt survival of wild salmon populations. For improving the performance of hatchery-reared smolts, it could be useful to examine opportunities to produce smolts that are in their smolt traits and abilities more similar to the wild smolts described in this thesis.

Nile Basin Initiative diskursiivisena areenana

Pro gradu -tutkielma käsittelee Niilin alueen poliittista kehitystä käyttäen viitekehyksenä Barry Buzanin, Ole Wæverin sekä Jaap de Wilden muodostaman Kööpenhaminan koulukunnan teoriaa turvallistamisesta sekä turvallisuuskomplekseista. Niilin kymmenen valtiota kattavan jokialueen poliittista historiaa on vuosikymmenet hallinnut valtioiden välinen epäluottamus, ja erityisesti Egypti, joka on täysin riippuvainen Niilin vesivaroista, on julistanut Niilin vesivarojen olevan sille kansallinen turvallisuuskysymys, ja se on uhannut käyttää myös voimakeinoja puolustaakseen vedenkäyttöoikeuksiaan. Egyptiä on myös voitu Niilin alueen hegemonina, sillä vanhat, alun perin siirtomaa-ajalta peräisin olevat sopimukset ovat antaneet sille lähes yksinoikeuden Niilin vesivarojen käyttöön. Samanaikaisesti erityisesti Etiopia, jonka alueelta suurin osa joen vesivaroista on peräisin, on pitänyt vanhoja sopimuksia epäreiluina, ja vedonnut maassa kasvavaan nälänhätään ja ruokapulaan, mutta myös useat muut alueen valtiot ovat esittäneet tyytymättömyytensä vanhoihin sopimuksiin. Vuonna 1999 kehityksessä tapahtui kuitenkin käänne, sillä jokilaakson valtiot perustivat tuolloin Niilin jokialueen yhteistyöjärjestön Nile Basin Initiativen (NBI). Tutkielma keskittyy NBI:n perustamisen jälkeiseen ajanjaksoon. Keskeinen tutkimuskysymys on, millä tavalla Niilin alueen jäsenvaltiot pyrkivät 2000-luvun alkupuolella keskinäisellä toiminnallaan muuttamaan alueella vallitsevaa kilpailu- ja konfliktiasetelmaa rauhanomaiseen suuntaan. Aineistona käytetään NBI:n omia kausijulkaisuja vuosilta 2006–2010, ja tutkimuksen menetelmänä käytetään diskurssianalyysia. Tutkitun aineiston perusteella on esitettävissä, että Niilin alueen poliittisesta kehityksestä on selvästi erotettavissa diskursseja, jotka ovat NBI:n toiminnan taustalla. Nämä ovat luottamuksen rakentamisen, yhteisten haasteiden voittamisen, yhteistyön tärkeyden sekä saavutusten diskurssit. Aineiston perusteella on nähtävissä, että valtioiden välisen luottamuksen vähitellen lisääntyessä yhteistyötä aletaan yhä enemmän legitimoida aikaisemmin yhteistyöstä saavutetuilla hyödyillä. Toisaalta diskurssianalyysin perusteella on myös nähtävissä, että alueella on ratkaisematta yhä fundamentaalisia ongelmia; uutta vesisopimusta ei ole vielä vuoteen 2010 mennessä aikaansaatu ja NBI:n jäsenvaltioiden suhteissa on myös nähtävissä kiistaa periaatteista, joihin yhteistyön olisi pohjauduttava. Tutkielma pyrkii analyysin avulla kehittämään myös Kööpenhaminan koulukunnan teoriaa turvallistamisesta. Kööpenhaminan koulukunta jättää turvallistamisen purkamiskehityksen kirjoituksissaan taka-alalle keskustelun painottuessa siihen, kuinka jokin asia tuodaan esille julkiseen turvallisuuskeskusteluun. Tältä osin tutkimuksen tavoitteena on esittää mahdollinen kehityskulku turvallistamisen purkamisdiskurssista. Teoriaosuudessa käytetyt keskeiset lähteet: Allan 2001,2009; Buzan 1991; Buzan ja Wæver 2003; Buzan, Wæver ja de Wilde 1998; Jutila 2008; Schulz 1995; Stritzel 2007; Turton 2001; Wæver 1995; Wendt 1992, 1999.

Fast- and drift-ice communities in the Bothnian Bay and the impact of UVA radiation on the Baltic Sea ice ecology

The aim of this thesis was to study ecology of Baltic Sea ice from two perspectives. In the first two studies, sea-ice ecology from riverine-influenced fast ice to drift ice in the Bothnian Bay was investigated, whereas the last two studies focus on the sensitivity of sea-ice bacteria and algae to UVA examined in situ. The seasonal sea ice cover is one of the main characteristics of the Baltic Sea, and despite the brackish parental water, the ice structure is similar to polar ice with saline brine inclusions, the sea ice habitat. The decreasing seawater salinity from the northern Baltic Sea to the Bothnian Bay translates to decreasing brine volumes along the gradient, governing the size and community structure of the food webs in ice. However, the drift and fast ice in the Bothnian Bay may differ greatly in this sense, as drift ice may have been formed at more southern locations. Rafting and the formation of snow ice are common processes in the ice field of the Bothnian Bay. As evidenced in this thesis, rafting altered the vertical distribution of organisms and snow-ice formation provided habitable space in the better-illuminated, nitrogen-rich surface layer. The divergence between fast and drift ice became apparent at the more advanced stages, and chlorophyte biomass decreased from fast to drift ice, while the opposite held true for protozoan and metazoan biomass. The brine volumes affected the communities somewhat, and a higher percentage of flagellate species was generally linked to lower brine volumes, whereas chain-forming diatoms were mostly concentrated in layers with larger brine volumes. These results add to knowledge of the ecological significance of the ice cover lasting up to 7 months per year in this area. Sea-ice food webs are generally light-limited, but while increasing light irradiances typically enhance the primary production and further, the secondary production in sea ice, any increase in solar radiation also includes an increase in harmful UVA radiation. The Baltic Sea ice microbial communities were clearly sensitive to UVA and the responses were strongly linked to the earlier light history, as well as to the solar irradiances they were exposed to. The increased biomass of chlorophytes and pennate diatoms, when UVA was excluded, indicates that their normally minor contribution to the biomass in the upper layers of sea ice might be partly dictated by UVA. The effects of UVA on bacterial production in Baltic Sea ice mostly followed the responses in algal growth, but occasionally the exposure to UVA even enhanced the bacterial production. The dominant bacterial class, Flavobacteria, seemed to be UVA-tolerant, whereas all the Alpha-, Beta- and Gammaproteobacteria present in the surface layer showed UVA sensitivity. These results indicate that changes in the light field of ice may alter the community structure and affect the functioning of ice food webs, and are of importance when the effects of thinning of the ice cover are assessed.

Effect of drainage basin characteristics on the diffuse load of phosphorus and nitrogen.

Tiivistelmä: Valuma-alueen vaikutus fosforin ja typen hajakuormitukseen.

Spread of the waters from the River Siikajoki in the Bothnian Bay.

Tiivistelmä: Siikajoen vesien leviäminen.