Effect of type of forage offered and breed on performance of crossbred suckler heifers and their calves

Selostus: Karkearehuruokinnan vaikutus risteytyshiehojen ja niiden vasikoiden kehitykseen sisäruokinta- ja laidunkaudella

Conservation Genetics of Exploited Finnish Salmonid Fishes

Genetic diversity is one of the levels of biodiversity that the World Conservation Union (IUCN) has recognized as being important to preserve. This is because genetic diversity is fundamental to the future evolution and to the adaptive flexibility of a species to respond to the inherently dynamic nature of the natural world. Therefore, the key to maintaining biodiversity and healthy ecosystems is to identify, monitor and maintain locally-adapted populations, along with their unique gene pools, upon which future adaptation depends. Thus, conservation genetics deals with the genetic factors that affect extinction risk and the genetic management regimes required to minimize the risk. The conservation of exploited species, such as salmonid fishes, is particularly challenging due to the conflicts between different interest groups. In this thesis, I conduct a series of conservation genetic studies on primarily Finnish populations of two salmonid fish species (European grayling, Thymallus thymallus, and lake-run brown trout, Salmo trutta) which are popular recreational game fishes in Finland. The general aim of these studies was to apply and develop population genetic approaches to assist conservation and sustainable harvest of these populations. The approaches applied included: i) the characterization of population genetic structure at national and local scales; ii) the identification of management units and the prioritization of populations for conservation based on evolutionary forces shaping indigenous gene pools; iii) the detection of population declines and the testing of the assumptions underlying these tests; and iv) the evaluation of the contribution of natural populations to a mixed stock fishery. Based on microsatellite analyses, clear genetic structuring of exploited Finnish grayling and brown trout populations was detected at both national and local scales. Finnish grayling were clustered into three genetically distinct groups, corresponding to northern, Baltic and south-eastern geographic areas of Finland. The genetic differentiation among and within population groups of grayling ranged from moderate to high levels. Such strong genetic structuring combined with low genetic diversity strongly indicates that genetic drift plays a major role in the evolution of grayling populations. Further analyses of European grayling covering the majority of the species’ distribution range indicated a strong global footprint of population decline. Using a coalescent approach the beginning of population reduction was dated back to 1 000-10 000 years ago (ca. 200-2 000 generations). Forward simulations demonstrated that the bottleneck footprints measured using the M ratio can persist within small populations much longer than previously anticipated in the face of low levels of gene flow. In contrast to the M ratio, two alternative methods for genetic bottleneck detection identified recent bottlenecks in six grayling populations that warrant future monitoring. Consistent with the predominant role of random genetic drift, the effective population size (N_e) estimates of all grayling populations were very low with the majority of N_e estimates below 50. Taken together, highly structured local populations, limited gene flow and the small N_e of grayling populations indicates that grayling populations are vulnerable to overexploitation and, hence, monitoring and careful management using the precautionary principles is required not only in Finland but throughout Europe. Population genetic analyses of lake-run brown trout populations in the Inari basin (northernmost Finland) revealed hierarchical population structure where individual populations were clustered into three population groups largely corresponding to different geographic regions of the basin. Similar to my earlier work with European grayling, the genetic differentiation among and within population groups of lake-run brown trout was relatively high. Such strong differentiation indicated that the power to determine the relative contribution of populations in mixed fisheries should be relatively high. Consistent with these expectations, high accuracy and precision in mixed stock analysis (MSA) simulations were observed. Application of MSA to indigenous fish caught in the Inari basin identified altogether twelve populations that contributed significantly to mixed stock fisheries with the Ivalojoki river system being the major contributor (70%) to the total catch. When the contribution of wild trout populations to the fisheries was evaluated regionally, geographically nearby populations were the main contributors to the local catches. MSA also revealed a clear separation between the lower and upper reaches of Ivalojoki river system – in contrast to lower reaches of the Ivalojoki river that contributed considerably to the catch, populations from the upper reaches of the Ivalojoki river system (>140 km from the river mouth) did not contribute significantly to the fishery. This could be related to the available habitat size but also associated with a resident type life history and increased cost of migration. The studies in my thesis highlight the importance of dense sampling and wide population coverage at the scale being studied and also demonstrate the importance of critical evaluation of the underlying assumptions of the population genetic models and methods used. These results have important implications for conservation and sustainable fisheries management of Finnish populations of European grayling and brown trout in the Inari basin.

Local and Regional Scale Determinants of Biodiversity Patterns in Boreal Agricultural Landscapes

This thesis examines the local and regional scale determinants of biodiversity patterns using existing species and environmental data. The research focuses on agricultural environments that have experienced rapid declines of biodiversity during past decades. Existing digital databases provide vast opportunities for habitat mapping, predictive mapping of species occurrences and richness and understanding the speciesenvironment relationships. The applicability of these databases depends on the required accuracy and quality of the data needed to answer the landscape ecological and biogeographical questions in hand. Patterns of biodiversity arise from confounded effects of different factors, such as climate, land cover and geographical location. Complementary statistical approaches that can show the relative effects of different factors are needed in biodiversity analyses in addition to classical multivariate models. Better understanding of the key factors underlying the variation in diversity requires the analyses of multiple taxonomic groups from different perspectives, such as richness, occurrence, threat status and population trends. The geographical coincidence of species richness of different taxonomic groups can be rather limited. This implies that multiple geographical regions should be taken into account in order to preserve various groups of species. Boreal agricultural biodiversity and in particular, distribution and richness of threatened species is strongly associated with various grasslands. Further, heterogeneous agricultural landscapes characterized by moderate field size, forest patches and non-crop agricultural habitats enhance the biodiversity of rural environments. From the landscape ecological perspective, the major threats to Finnish agricultural biodiversity are the decline of connected grassland habitat networks, and general homogenization of landscape structure resulting from both intensification and marginalization of agriculture. The maintenance of key habitats, such as meadows and pastures is an essential task in conservation of agricultural biodiversity. Furthermore, a larger landscape context should be incorporated in conservation planning and decision making processes in order to respond to the needs of different species and to maintain heterogeneous rural landscapes and viable agricultural diversity in the future.

Phylogeography and population genetics of north European Atlantic salmon (Salmo salar L.)

Although abundant in the number of individuals, the Atlantic salmon may be considered as a threatened species in many areas of its native distribution range. Human activities such as building of power plant dams, offshore overfishing, pollution, clearing of riverbeds for timber floating and badly designed stocking regimes have diminished the distribution of Atlantic salmon. As a result of this, many of the historical populations both in Europe and northern America have gone extinct or are severely depressed. In fact, only 1% of Atlantic salmon existing today are of natural origin, the rest being farmed salmon. All of this has lead to a vast amount of research and many restoration programmes aiming to bring Atlantic salmon back to rivers from where it has vanished. However, many of the restoration programmes conducted thus far have been unsuccessful due to inadequate scientific research or lack of its implementation, highlighting the fact that more research is needed to fully understand the biology of this complex species. The White and Barents Seas in northwest Russia are among the last regions in Europe where Atlantic salmon populations are still stable, thus forming an important source of biodiversity for the entire European region. Salmon stocks from this area are also of immense economic and social importance for the local people in the form of fishing tourism. The main aim of this thesis was to elucidate the post-glacial history and population genetic structure of north European and particularly northwest Russian Atlantic salmon, both of which are aspects of great importance for the management and conservation of the species. Throughout the whole thesis, these populations were studied by utilizing microsatellites as the main molecular tool. One of the most important discoveries of the thesis was the division of Atlantic salmon from the White and Barents Seas into four separate clusters, which has not been observed in previous studies employing nuclear markers although is supported by mtDNA studies. Populations from the western Barents Sea clustered together with the northeast Atlantic populations into a clearly distinguishable group while populations from the White Sea and eastern Barents Sea were separated into three additional groups. This has important conservation implications as this thesis clearly indicates that conservation of populations from all of the observed clusters is warranted in order to conserve as much of the genetic diversity as possible in this area. The thesis also demonstrates how differences in population life histories within a species, migratory behaviour in this case, and in their phylogeographic origin affect the genetic characteristics of populations, namely diversity and divergence levels. The anadromous populations from the Atlantic Ocean, White Sea and Barents Sea possessed higher levels of genetic diversity than the anadromous populations form the Baltic Sea basin. Among the non-anadromous populations the result was the opposite: the Baltic freshwater populations were more variable. This emphasises the importance of taking the life history of a population into consideration when developing conservation strategies: due to the limited possibilities for new genetic diversity to be generated via gene flow, it is expected that freshwater Atlantic salmon populations would be more vulnerable to extinction following a population crash and thus deserve a high conservation status. In the last chapter of this thesis immune relevant marker loci were developed and screened for signatures of natural selection along with loci linked to genes with other functions or no function at all. Also, a novel landscape genomics method, which combines environmental information with molecular data, was employed to investigate whether immune relevant markers displayed significant correlations to various environmental variables more frequently than other loci. Indications of stronger selection pressure among immune-relevant loci compared to non-immune relevant EST-linked loci was found but further studies are needed to evaluate whether it is a common phenomenon in Atlantic salmon.

Spatiotemporal Aspects of Environmental Monitoring in the Complex Coastal Region of Southwest Finland

The management and conservation of coastal waters in the Baltic is challenged by a number of complex environmental problems, including eutrophication and habitat degradation. Demands for a more holistic, integrated and adaptive framework of ecosystem-based management emphasize the importance of appropriate information on the status and changes of the aquatic ecosystems. The thesis focuses on the spatiotemporal aspects of environmental monitoring in the extensive and geomorphologically complex coastal region of SW Finland, where the acquisition of spatially and temporally representative monitoring data is inherently challenging. Furthermore, the region is subject to multiple human interests and uses. A holistic geographical approach is emphasized, as it is ultimately the physical conditions that set the frame for any human activity. Characteristics of the coastal environment were examined using water quality data from the database of the Finnish environmental administration and Landsat TM/ETM+ images. A basic feature of the complex aquatic environment in the Archipelago Sea is its high spatial and temporal variability; this foregrounds the importance of geographical information as a basis of environmental assessments. While evidence of a consistent water turbidity pattern was observed, the coastal hydrodynamic realm is also characterized by high spatial and temporal variability. It is therefore also crucial to consider the spatial and temporal representativeness of field monitoring data. Remote sensing may facilitate evaluation of hydrodynamic conditions in the coastal region and the spatial extrapolation of in situ data despite their restrictions. Additionally, remotely sensed images can be used in the mapping of many of those coastal habitats that need to be considered in environmental management. With regard to surface water monitoring, only a small fraction of the currently available data stored in the Hertta-PIVET register can be used effectively in scientific studies and environmental assessments. Long-term consistent data collection from established sampling stations should be emphasized but research-type seasonal assessments producing abundant data should also be encouraged. Thus a more comprehensive coordination of field work efforts is called for. The integration of remote sensing and various field measurement techniques would be especially useful in the complex coastal waters. The integration and development of monitoring system in Finnish coastal areas also requires further scientific assesement of monitoring practices. A holistic approach to the gathering and management of environmental monitoring data could be a cost-effective way of serving a multitude of information needs, and would fit the holistic, ecosystem-based management regimes that are currently being strongly promoted in Europe.

Statistical Methods for Conservation and Alignment Quality in Proteins

Construction of multiple sequence alignments is a fundamental task in Bioinformatics. Multiple sequence alignments are used as a prerequisite in many Bioinformatics methods, and subsequently the quality of such methods can be critically dependent on the quality of the alignment. However, automatic construction of a multiple sequence alignment for a set of remotely related sequences does not always provide biologically relevant alignments.Therefore, there is a need for an objective approach for evaluating the quality of automatically aligned sequences. The profile hidden Markov model is a powerful approach in comparative genomics. In the profile hidden Markov model, the symbol probabilities are estimated at each conserved alignment position. This can increase the dimension of parameter space and cause an overfitting problem. These two research problems are both related to conservation. We have developed statistical measures for quantifying the conservation of multiple sequence alignments. Two types of methods are considered, those identifying conserved residues in an alignment position, and those calculating positional conservation scores. The positional conservation score was exploited in a statistical prediction model for assessing the quality of multiple sequence alignments. The residue conservation score was used as part of the emission probability estimation method proposed for profile hidden Markov models. The results of the predicted alignment quality score highly correlated with the correct alignment quality scores, indicating that our method is reliable for assessing the quality of any multiple sequence alignment. The comparison of the emission probability estimation method with the maximum likelihood method showed that the number of estimated parameters in the model was dramatically decreased, while the same level of accuracy was maintained. To conclude, we have shown that conservation can be successfully used in the statistical model for alignment quality assessment and in the estimation of emission probabilities in the profile hidden Markov models.

Assessment of Coastal Bird Populations and Habitats on the Finnish Coast of the Baltic Sea: Implications for Monitoring and Management

Coastal birds are an integral part of coastal ecosystems, which nowadays are subject to severe environmental pressures. Effective measures for the management and conservation of seabirds and their habitats call for insight into their population processes and the factors affecting their distribution and abundance. Central to national and international management and conservation measures is the availability of accurate data and information on bird populations, as well as on environmental trends and on measures taken to solve environmental problems. In this thesis I address different aspects of the occurrence, abundance, population trends and breeding success of waterbirds breeding on the Finnish coast of the Baltic Sea, and discuss the implications of the results for seabird monitoring, management and conservation. In addition, I assess the position and prospects of coastal bird monitoring data, in the processing and dissemination of biodiversity data and information in accordance with the Convention on Biological Diversity (CBD) and other national and international commitments. I show that important factors for seabird habitat selection are island area and elevation, water depth, shore openness, and the composition of island cover habitats. Habitat preferences are species-specific, with certain similarities within species groups. The occurrence of the colonial Arctic Tern (Sterna paradisaea) is partly affected by different habitat characteristics than its abundance. Using long-term bird monitoring data, I show that eutrophication and winter severity have reduced the populations of several Finnish seabird species. A major demographic factor through which environmental changes influence bird populations is breeding success. Breeding success can function as a more rapid indicator of sublethal environmental impacts than population trends, particularly for long-lived and slowbreeding species, and should therefore be included in coastal bird monitoring schemes. Among my target species, local breeding success can be shown to affect the populations of the Mallard (Anas platyrhynchos), the Eider (Somateria mollissima) and the Goosander (Mergus merganser) after a time lag corresponding to their species-specific recruitment age. For some of the target species, the number of individuals in late summer can be used as an easier and more cost-effective indicator of breeding success than brood counts. My results highlight that the interpretation and application of habitat and population studies require solid background knowledge of the ecology of the target species. In addition, the special characteristics of coastal birds, their habitats, and coastal bird monitoring data have to be considered in the assessment of their distribution and population trends. According to the results, the relationships between the occurrence, abundance and population trends of coastal birds and environmental factors can be quantitatively assessed using multivariate modelling and model selection. Spatial data sets widely available in Finland can be utilised in the calculation of several variables that are relevant to the habitat selection of Finnish coastal species. Concerning some habitat characteristics field work is still required, due to a lack of remotely sensed data or the low resolution of readily available data in relation to the fine scale of the habitat patches in the archipelago. While long-term data sets exist for water quality and weather, the lack of data concerning for instance the food resources of birds hampers more detailed studies of environmental effects on bird populations. Intensive studies of coastal bird species in different archipelago areas should be encouraged. The provision and free delivery of high-quality coastal data concerning bird populations and their habitats would greatly increase the capability of ecological modelling, as well as the management and conservation of coastal environments and communities. International initiatives that promote open spatial data infrastructures and sharing are therefore highly regarded. To function effectively, international information networks, such as the biodiversity Clearing House Mechanism (CHM) under the CBD, need to be rooted at regional and local levels. Attention should also be paid to the processing of data for higher levels of the information hierarchy, so that data are synthesized and developed into high-quality knowledge applicable to management and conservation.

An experimental and theoretical study of multiphase flow in a circulating fluidized bed

A systematic averaging procedure has been derived in order to obtain an integral form of conservation equations for dispersed multiphase flow, especially applicable to fluidized beds. A similar averaging method is applied further to formulate macroscopic integral equations, which can be used in one-dimensional and macroscopic multi dimensional models. Circulating fluid bed hydrodynamics has been studied experimentally and both macroscopic and microscopic flow profiles have been measured in a cold model. As an application of the theory, the one dimensional model has been used to study mass and momentum conservation of gas and solid in a circulating fluid bed. Axial solid mixing has also been modelled by the one dimensional model and mixing parameters have been evaluated.

Analysis and modelling of chemical looping combustion process with and without oxygen uncoupling

Effective control and limiting of carbon dioxide (CO₂) emissions in energy production are major challenges of science today. Current research activities include the development of new low-cost carbon capture technologies, and among the proposed concepts, chemical combustion (CLC) and chemical looping with oxygen uncoupling (CLOU) have attracted significant attention allowing intrinsic separation of pure CO₂ from a hydrocarbon fuel combustion process with a comparatively small energy penalty. Both CLC and CLOU utilize the well-established fluidized bed technology, but several technical challenges need to be overcome in order to commercialize the processes. Therefore, development of proper modelling and simulation tools is essential for the design, optimization, and scale-up of chemical looping-based combustion systems. The main objective of this work was to analyze the technological feasibility of CLC and CLOU processes at different scales using a computational modelling approach. A onedimensional fluidized bed model frame was constructed and applied for simulations of CLC and CLOU systems consisting of interconnected fluidized bed reactors. The model is based on the conservation of mass and energy, and semi-empirical correlations are used to describe the hydrodynamics, chemical reactions, and transfer of heat in the reactors. Another objective was to evaluate the viability of chemical looping-based energy production, and a flow sheet model representing a CLC-integrated steam power plant was developed. The 1D model frame was succesfully validated based on the operation of a 150 kWth laboratory-sized CLC unit fed by methane. By following certain scale-up criteria, a conceptual design for a CLC reactor system at a pre-commercial scale of 100 MWth was created, after which the validated model was used to predict the performance of the system. As a result, further understanding of the parameters affecting the operation of a large-scale CLC process was acquired, which will be useful for the practical design work in the future. The integration of the reactor system and steam turbine cycle for power production was studied resulting in a suggested plant layout including a CLC boiler system, a simple heat recovery setup, and an integrated steam cycle with a three pressure level steam turbine. Possible operational regions of a CLOU reactor system fed by bituminous coal were determined via mass, energy, and exergy balance analysis. Finally, the 1D fluidized bed model was modified suitable for CLOU, and the performance of a hypothetical 500 MWth CLOU fuel reactor was evaluated by extensive case simulations.

Taxonomy and Species Richness of Amazonian Pimplinae and Rhyssinae (hymenoptera: ichneumonidae)

The Amazonian region, the biggest rain forest of our planet, is known for its extraordinary biodiversity. Most of this diversity is still unexplored and new species of different taxa are regularly found there. In this region, as in most areas of the world, insects are some of the most abundant organisms. Therefore, studying this group is important to promote the conservation of these highly biodiverse ecosystems of the planet. Among insects, parasitoid wasps are especially interesting because they have potential for use as biodiversity indicators and biological control agents in agriculture and forestry. The parasitoid wasp family Ichneumonidae is one of the most species rich groups among the kingdom Animalia. This group is still poorly known in many areas of the world; the Amazonian region is a clear example of this situation. Ichneumonids have been thought to be species poor in Amazonia and other tropical areas. However, recent studies are suggesting that parasitoid wasps may be quite abundant in Amazonia and possibly in most tropical areas of the world. The aim of my doctoral thesis is to study the species richness and taxonomy of two of the best known ichneumonid subfamilies in the Neotropical region, Pimplinae and Rhyssinae. To do this I conducted two extensive sampling programs in the Peruvian Amazonia. I examined also a large number of Neotropical ichneumonids deposited to different natural history museums. According to the results of my thesis, the species richness of these parasitoids in the Amazonian region is considerably higher than previously reported. In my research, I firstly further develop the taxonomy of these parasitoids by describing many new species and reporting several new faunistic records (I, II, III). In this first part I focus on two genera (Xanthopimpla and Epirhyssa) which were thought to be rather species poor. My thesis demonstrates that these groups are actually rather species rich in the Amazonian region. Secondly, I concentrate on the species richness of these parasitoids in a global comparison showing that the Neotropical region and especially the Peruvian Amazonia is one of the most species rich areas of Pimpliformes ichneumonids (V). Furthermore, I demonstrate that with the data available to date no clear latitudinal gradient in species richness is visible. Thirdly, increasing the macroecological knowledge of these parasitoids I show that some previously unreported ichneumonid subfamilies are present in the Amazonian region (IV). These new insights and the results of the global comparison of ichneumonid inventories suggest that the previous belief of low diversity in the tropics is most likely related to a lack of sampling effort in the region. Overall, my research increases the knowledge of Neotropical ichneumonids highlighting the importance of Peruvian Amazonia as one of the diversity hotspots of parasitoid wasps.

Bumblebees in a Changing Climate: Evaluating the Effects of Temperature on Queen Performance

Bumblebees are a very essential group of pollinating insects, but their populations have declined drastically during the past decades. We need to understand why their numbers are decreasing and what can be done to reverse this trend. Climate change-related phenomena, such as changes in the overwintering temperatures and spring conditions, are among the most prominent threats to bumblebees. Queens have a special role in the lifecycle of bumblebees because they overwinter and start new colonies the next year. Their successful performance: survival, overwintering ability, longevity, immune competence, and nest establishing capability in spring, is highly important for bumblebee populations. However, the effects of climate change on bumblebee queen performance remain unknown. The main objective of this thesis was to assess how temperature affects the performance of bumblebee queens during and after overwintering. The effects of warm temperature predicted by climate change scenarios on queen survival and stress-tolerance were studied by a four-month artificial diapause of bumblebee queens at two temperatures (9°C and 1.8°C). Bumblebee colonies were also reared in a laboratory and factors affecting colony characteristics were examined. In addition, queen performance during spring was studied in a starvation experiment using two temperatures (15°C as normal; 24°C as warmer than average) and queens collected from nature right after their emergence. My research revealed how temperature affects queen performance, and queen size was found to be an important factor determining the direction of some of these effects. We found a 0.4g weight threshold for bumblebee queens to be able to survive overwintering. In addition, during mild winters, larger queens have a higher chance than smaller ones to survive through winter and also to cope with immunological stresses after overwintering. During cold conditions, which are normal in the current climatic situation, this advantage disappears. In the spring starvation experiment, the starved queens survived approximately eight days longer in 15°C than in 24°C, which means that starvation risk rises significantly with increasing spring temperature, in a situation where food is scarce due to for example frost damage or asynchrony between bumblebees and their important food plants. These results could mean that in the future climate, larger queens are better able to survive the winter, initiate their nests and start rearing their offspring. This may be problematic, because I also detected two alternative strategies of colony development that differ between large and small queens; larger queens start to lay eggs earlier at nest initiation, their colonies mature later, they produce more workers, and they have a more strongly male biased sex allocation compared with smaller queens. If larger queens have a greater change of producing offspring after a mild winter, this could lead to a significant decline in the total production of new queens at a population level. Thus, it seems that queen size could act as one mechanism regulating the population level outcomes in different temperatures. The new information presented in my thesis reinforces that basic research, monitoring, and local species conservation of bumblebees both in Finland and globally must be increased to ensure that this highly important pollinator group survives in the face of climate change.