Methicillin Resistance In Staphylococci : Horizontal Transfer of Mobile Genetic Element (SCCmec) Between Staphylococcal Species

Regardless of the existence of antibiotics, infectious diseases are the leading causes of death in the world. Staphylococci cause many infections of varying severity, although they can also exist peacefully in many parts of the human body. Most often Staphylococcus aureus colonises the nose, and that colonisation is considered to be a risk factor for spread of this bacterium. S. aureus is considered to be the most important Staphylococcus species. It poses a challenge to the field of medicine, and one of the most problematic aspects is the drastic increase of the methicillin-resistant S. aureus (MRSA) strains in hospitals and community world-wide, including Finland. In addition, most of the clinical coagulase-negative staphylococcus (CNS) isolates express resistance to methicillin. Methicillin-resistance in S. aureus is caused by the mecA gene that encodes an extra penicillin-binding protein (PBP) 2a. The mecA gene is found in a mobile genomic island called staphylococcal chromosome cassette mec (SCCmec). The SCCmec consists of the mec gene and cassette chromosome recombinase (ccr)gene complexes. The areas of the SCCmec element outside the ccr and mec complex are known as the junkyard J regions. So far, eight types of SCCmec(SCCmec I- SCCmec VIII) and a number of variants have been described. The SCCmec island is an acquired element in S. aureus. Lately, it appears that CNS might be the storage place of the SCCmec that aid the S. aureus by providing it with the resistant elements. The SCCmec is known to exist only in the staphylococci. The aim of the present study was to investigate the horizontal transfer of SCCmec between the S. aureus and CNS. One specific aim was to study whether or not some methicillin-sensitive S. aureus (MSSA) strains are more inclined to receive the SCCmec than others. This was done by comparing the genetic background of clinical MSSA isolates in the health care facilities of the Helsinki and Uusimaa Hospital District in 2001 to the representatives of the epidemic MRSA (EMRSA) genotypes, which have been encountered in Finland during 1992-2004. Majority of the clinical MSSA strains were related to the EMRSA strains. This finding suggests that horizontal transfer of SCCmec from unknown donor(s) to several MSSA background genotypes has occurred in Finland. The molecular characteristics of representative clinical methicillin-resistant S. epidermidis (MRSE) isolates recovered in Finnish hospitals between 1990 and 1998 were also studied, examining their genetic relation to each other and to the internationally recognised MRSE clones as well, so as to ascertain the common traits between the SCCmec elements in MRSE and MRSA. The clinical MRSE strains were genetically related to each other; eleven PFGE types were associated with sequence type ST2 that has been identified world-wide. A single MRSE strain may possess two SCCmec types III and IV, which were recognised among the MRSA strains. Moreover, six months after the onset of an outbreak of MRSA possessing a SCCmec type V in a long-term care facility in Northern Finland (LTCF) in 2003, the SCCmec element of nasally carried methicillin-resistant staphylococci was studied. Among the residents of a LTCF, nasal carriage of MR-CNS was common with extreme diversity of SCCmec types. MRSE was the most prevalent CNS species. Horizontal transfer of SCCmec elements is speculated to be based on the sharing of SCCmec type V between MRSA and MRSE in the same person. Additionally, the SCCmec element of the clinical human S. sciuri isolates was studied. Some of the SCCmec regions were present in S. sciuri and the pls gene was common in it. This finding supports the hypothesis of genetic exchange happening between staphylococcal species. Evaluation of the epidemiology of methicillin-resistant staphylococcal colonisation is necessary in order to understand the apparent emergence of these strains and to develop appropriate control strategies. SCCmec typing is essential for understanding the emergence of MRSA strains from CNS, considering that the MR-CNS may represent the gene pool for the continuous creation of new SCCmec types from which MRSA might originate.

Genetic Basis of Pituitary Adenoma Predisposition

Much of the global cancer research is focused on the most prevalent tumors; yet, less common tumor types warrant investigation, since A rare disorder is not necessarily an unimportant one . The present work discusses a rare tumor type, the benign adenomas of the pituitary gland, and presents the advances which, during the course of this thesis work, contributed to the elucidation of a fraction of their genetic background. Pituitary adenomas are benign neoplasms of the anterior pituitary lobe, accounting for approximately 15% of all intracranial tumors. Pituitary adenoma cells hypersecrete the hormones normally produced by the anterior pituitary tissue, such as growth hormone (GH) and prolactin (PRL). Despite their non-metastasizing nature, these adenomas can cause significant morbidity and have to be adequately treated; otherwise, they can compromise the patient s quality of life, due to conditions provoked by hormonal hypersecretion, such as acromegaly in the case of GH-secreting adenomas, or due to compressive effects to surrounding tissues. The vast majority of pituitary adenomas arise sporadically, whereas a small subset occur as component of familial endocrine-related tumor syndromes, such as Multiple Endocrine Neoplasia type 1 (MEN1) and Carney complex (CNC). MEN1 is caused by germline mutations in the MEN1 tumor suppressor gene (11q13), whereas the majority of CNC cases carry germline mutations in the PRKAR1A gene (17q24). Pituitary adenomas are also encountered in familial settings outside the context of MEN1 and CNC, but unlike in the latter syndromes, their genetic background until recently remained elusive. Evidence in previous literature supported the notion that a tumor suppressor gene on 11q13, residing very close to but still distinct from MEN1, causes genetic susceptibility to pituitary tumors. The aim of the study was to identify the genetic cause of a low penetrance form of Pituitary Adenoma Predisposition (PAP) in families from Northern Finland. The present work describes the methodological approach that led to the identification of aryl hydrocarbon receptor interacting protein (AIP) as the gene causing PAP. Combining chip-based technologies (SNP and gene expression arrays) with traditional gene mapping methods and genealogy data, we showed that germline AIP mutations cause PAP in familial and sporadic settings. PAP patients were diagnosed with mostly adenomas of the GH/PRL-secreting cell lineage. In Finland, two AIP mutations accounted for 16% of all patients diagnosed with GH-secreting adenomas, and for 40% of patients being younger than 35 years of age at diagnosis. AIP is suggested to act as a tumor suppressor gene, a notion supported by the nature of the identified mutations (most are truncating) and the biallelic inactivation of AIP in the tumors studied. AIP has been best characterized as a cytoplasmic interaction partner of aryl hydrocarbon receptor (AHR), also known as dioxin receptor, but it has other partners as well. The mechanisms that underlie AIP-mediated pituitary tumorigenesis are to date largely unknown and warrant further investigation. Because AIP was identified in the genetically homogeneous Finnish population, it was relevant to examine its contribution to PAP in other, more heterogeneous, populations. Analysis of pituitary adenoma patient series of various ethnic origins and differing clinical settings revealed germline AIP mutations in all cohorts studied, albeit with low frequencies (range 0.8-7.4%). Overall, PAP patients were typically diagnosed at a young age (range 8-41 years), mainly with GH-secreting adenomas, without strong family history of endocrine disease. Because many PAP patients did not display family history of pituitary adenomas, detection of the condition appeared challenging. AIP immunohistochemistry was tested as a molecular pre-screening tool on mutation-positive versus mutation-negative tumors, and proved to be a potentially useful predictor of PAP. Mutation screening of a large cohort of colorectal, breast, and prostate tumors did not reveal somatic AIP mutations. These tumors, apart from being the most prevalent among men and women worldwide, have been associated with acromegaly, particularly colorectal neoplasia. In this material, AIP did not appear to contribute to the pathogenesis of these common tumor types and other genes seem likely to play a role in such tumorigenesis. Finally, the contribution of AIP in pediatric onset pituitary adenomas was examined in a unique population-based cohort of sporadic pituitary adenoma patients from Italy. Germline AIP mutations may account for a subset of pediatric onset GH-secreting adenomas (in this study one of seven GH-secreting adenoma cases or 14.3%), and appear to be enriched among young (≤25 years old) patients. In summary, this work reveals a novel tumor susceptibility gene, namely AIP, which causes genetic predisposition to pituitary adenomas, in particular GH-secreting adenomas. Moreover, it provides molecular tools for identification of individuals predisposed for PAP. Further elaborate studies addressing the functional role of AIP in normal and tumor cells will hopefully expand our knowledge on endocrine neoplasia and reveal novel cellular mechanisms of pituitary tumorigenesis, including potential drug targets.

The DISC1 Pathway in the Genetic Etiology of Schizophrenia

Schizophrenia is a severe psychotic disorder affecting 0.5-1 % of the population. The disorder is characterized by hallucinations; delusions; disorganized behavior and speech; avolition; anhedonia; flattened affect and cognitive deficits. The etiology of the disorder is complex with evidence for multiple genes contributing to the onset of the disorder along with environmental factors. DISC1 is one of the most promising candidate genes for schizophrenia. It codes for a protein which takes part in numerous molecular interactions along several pathways. This network, termed as the DISC1 pathway, is evidently important for the development and maturation of the central nervous system from the embryo until young adulthood. Disruption at these pathways is thought to predispose schizophrenia. In the present study, we have studied the DISC1 pathway in the etiology of schizophrenia in the Finnish population. We have utilized large Finnish samples; the schizophrenia family sample where DISC1 was originally shown to associate with schizophrenia and the Northern Finland birth cohort 1966 (NFBC66). Several DISC1 binding partners displayed evidence for association in the family sample along with DISC1. Through a genome-wide linkage study, we found a significant linkage signal to a locus where a DISC1 binding partner NDE1 is located at the carriers of a certain DISC1 risk variant. In a follow-up study, genetic markers in NDE1 displayed significant evidence for association with schizophrenia. Further exploration of association between 11 genes of the DISC1 pathway and schizophrenia led to recognition of novel variants in NDEL1, PDE4B and PDE4D that significantly either increased or decreased the risk for schizophrenia. Further, we found evidence that DISC1 itself has a significant role in the human mental functioning even in the healthy population. Variants in DISC1 had a significant effect on anhedonia which is a trait present at everybody but is in its severe form one of the main symptoms of schizophrenia and correlates with the risk of developing the disorder. Further, utilizing genome-wide marker data, we recognized three genes; MIR620; CCDC141 and LCT; that are closely related to the DISC1 pathway but which effects on anhedonia were observable only at the individuals who carried these specific DISC1 variants. Our findings significantly add up to the previous evidence for the involvement of DISC1 and the DISC1 pathway in the etiology of schizophrenia and psychosis. Our results support the concept of a number of DISC1 pathway related genes contributing in the etiology of schizophrenia along with DISC1 and provide new candidates for the studies of schizophrenia. Our findings also significantly increase the importance of DISC1 itself as having a role in psychological functioning in the general population.

The role of AIP and CDKN1B/p27Kip1 in endocrine neoplasia

Identification of genes predisposing to tumor syndromes has raised general awareness of tumorigenesis. Genetic testing of tumor susceptibility genes aids the recognition of individuals at increased risk of tumors. Identification of novel predisposing genes enables further studies concerning the classification of potential associated tumors and the definition of target patient group. Pituitary adenomas are common, benign neoplasms accounting for approximately 15% of all intracranial tumors. Accurate incidence estimation is challenging since a great portion of these adenomas are small and asymptomatic. Clinically relevant adenomas, that cause symptoms due to the expansion of the cell mass or the over-secretion of normally produced hormones, occur in approximately one of 1 000 individuals. Although the majority of pituitary adenomas are sporadic, a minority occur as components of familial syndromes, such as Multiple Endocrine Neoplasia type 1 (MEN1) and Carney complex (CNC). MEN1 syndrome is caused by germ-line mutations in the MEN1 gene, whereas most of the CNC patients carry the mutated protein kinase A (PKA) regulatory subunit-1-α (PRKAR1A) gene. Recently, other conditions predisposing to endocrine tumors have been identified: Pituitary Adenoma Predisposition (PAP) and MEN type 4 (MEN4). PAP was originally identified in a genetically homogeneous Finnish population. In a population based cohort from Northern Finland, aryl hydrocarbon receptor-interacting protein (AIP) gene mutations were found in 16% of all patients diagnosed with growth hormone (GH) producing pituitary adenoma, and in 40% of the subset of patients who were diagnosed under the age of 35 years. Since AIP mutations were originally described in a defined, homogeneous population from Northern Finland, it was relevant to study whether mutations also occur in more heterogeneous populations. In patient cohorts with different ethnic origins and variable clinical phenotypes, germ-line AIP mutations were detectable at low frequencies (range 0.8-7.4%). AIP mutation-positive patients were often diagnosed with a GH-producing adenoma at a young age, and usually had no family history of endocrine tumors. The low frequency of AIP mutations in randomly selected patients, and the lack of any family history of pituitary adenomas create a challenge for the identification of PAP patients. Our preliminary study suggests that AIP immunohistochemistry may serve as a pre-screening tool to distinguish between the AIP mutation-negative and the mutation-positive tumors. Tumors of various endocrine glands are components of MEN1 and CNC syndromes. Somatic MEN1 and PRKAR1A mutations in sporadic pituitary adenomas are rare, but occur in some of the other tumors related to these syndromes. The role of AIP mutations in endocrine neoplasia was studied and our results indicated that somatic AIP mutations are rare or non-existent in sporadic tumors of endocrine glands (0 of 111). Furthermore, germ-line AIP mutations in prolactin producing adenomas (2 of 9) confirmed the role of this pituitary tumor type in the PAP phenotype. Thyroid disorders are common in the general population, and the majority of them are sporadic. Interestingly, it has been suggested that thyroid disorders might be more common in PAP families. For this reason we studied germ-line AIP mutations in 93 index cases from familial non-medullary thyroid cancer (NMTC) families. The underlying gene or genes for familial NMTC have not been identified yet. None of the patients had any potentially pathogenic AIP mutation. This suggests that AIP is unlikely to play a role in familial NMTCs. A novel multiple endocrine syndrome was originally described in rats with phenotypic features of human MEN type 1 and 2. Germ-line mutations of cyclin-dependent kinase inhibitor 1B (CDKN1B also known as p27Kip1) gene were reported later in these rats and a germ-line mutation was also identified in one human family with MEN1-like phenotype (later named MEN4). To confirm the importance of this gene’s mutations in humans, we performed a mutation screening in MEN-like patients and in patients with pituitary adenoma. Our results indicate that CDKN1B/p27Kip1 mutations appear in a small portion of MEN1-like patients (one of 36), and that such mutations are rare or non-existent in both familial (0 of 19) and sporadic pituitary adenoma patients (0 of 50). In conclusion, this work strengthens the tumor susceptibility role of AIP and CDKN1B/p27Kip1 in endocrine neoplasia. Clarifying the PAP phenotype facilitates the identification of potential AIP mutation carriers. Genetic counseling can be offered to the relatives and follow-up of the mutation carriers can be organized, hence an earlier diagnosis is feasible.

Nuorten männiköiden kehitys - taimikonhoito ja ensiharvennus

The aim of this study was to investigate the effects of location, site type, regeneration method and precommercial thinning on the characteristics and development of young, even-aged, pure Scots pine stands. In addition, the effects of timing and intensity of first commercial thinning on the yield and profitability during the rotation period were also studied. The stand characteristics and external quality of young Scots pine stands and stand-level growth models were based on extensive inventory data of the Finnish Forest Research Institute for young Scots pine stands (3 measurement times, 192 stands). The effect of precommercial thinning on stand development was examined on the basis of long-term experiments (13 stands, 169 plots). The effect of timing and intensity of the first commercial thinning on yield and profitability were based on measurements made in first commercial thinnings (27 stands of Metsähallitus), and the further stand development was modeled using the MOTTI simulator. The thesis was based on four articles and a summary. Stand level growth models were developed for young, even-aged Scots pine stands. The models reliably predicted the development up until the first commercial thinning stage. The stand density of young Scots pine stands in Finland was moderately low compared to the target values. In addition, the external quality of pines was low on average. The low stand density and poor external quality will result in the need for quality tree selection in thinnings, if high quality sawn timber is required. In Northern Finland, only 20% of the dominant trees were classified as normal. This will lead to the situation where external quality will remain relatively poor up until the end of rotation. Early and light precommercial thinning (Hdom 3 m, to a density of 3000 trees per hectare) increased the thinning removal by 40% compared to late and more intensive precommercial thinning (at 7 meters to a density of 2000 trees per hectare). A model for the effect of precommercial thinning on merchantable thinning removal at the first commercial thinning was developed for forest management planning purposes. When the recommended time of first commercial thinning was delayed from a dominant height of 12 m to 16 m, or by ten years, the yield of merchantable wood was doubled. Simultaneously, the current value of the stumpage revenues (with 4% interest rate) was increased on the average by 65% (330 € per hectare). Variation in stumpage prices or interest rates did not have any effect on the final results. Without exception, delaying the first commercial thinning by ten years seemed to be the most profitable method. This presupposes that precommercial thinning has been carried out at the right time and that tree quality aspects do not be specially considered. Furthermore, the wood yield and economic outcome from the entire rotation were similar regardless of whether the first thinning was performed at the time currently recommended or ten years later.

Bat habitat requirements : implications for land use planning

Knowledge of the habitat requirements of bat species is needed in decision making in land use planning. Bats' hibernation requirements were studied both in Estonia and in southern Finland. In both countries, the northern bat and the brown long-eared bat hibernated in colder and drier locations, whereas Daubenton's bat and Brandt's/whiskered bats hibernated in warmer and more humid locations. In Estonia, the pond bat hibernated in the warmest and most humid conditions, whereas Natterer's bat hibernated in the coldest and driest conditions. Hibernacula were at their coldest in mid-season and became warmer towards the end of the season. The results suggest that bats made an active choice of colder hibernation temperatures at the seasons end. They minimised the negative effects of hibernation early in the hibernation season by hibernating in warmer locations and energy expenditure late in the hibernation season by hibernating in colder locations. The use of foraging habitats was studied in northern and southern Finland. The northern bat used foraging sites opportunistically. Daubenton's bat foraged mainly in water habitats, whereas Brandt's/whiskered bats and the brown long-eared bat foraged mainly in forest habitats. In northern Finland, Daubenton's bats foraged almost exclusively on rivers and typically together with the northern bat. Daubenton's bats and Brandt's/whiskered bats were found only where there were lower ambient light levels. One of the most important things in the management of foraging areas for them is to keep them shady. Hibernacula in Finland typically housed few bats, suggesting that hibernation sites used by even a small number of bats are important. Bats typically used natural stone for hibernation suggesting that natural underground sites in rocks or cliffs or man-made underground sites built using natural stone are important for them. The results suggest that appropriate timing of surveys may vary according to the species and latitude.

Plant secondary compounds and soil microbial processes in carbon and nitrogen cycling in relation to tree species

The aim of this study was to explore soil microbial activities related to C and N cycling and the occurrence and concentrations of two important groups of plant secondary compounds, terpenes and phenolic compounds, under silver birch (Betula pendula Roth), Norway spruce (Picea abies (L.) Karst) and Scots pine (Pinus sylvestris L.) as well as to study the effects of volatile monoterpenes and tannins on soil microbial activities. The study site, located in Kivalo, northern Finland, included ca. 70-year-old adjacent stands dominated by silver birch, Norway spruce and Scots pine. Originally the soil was very probably similar in all three stands. All forest floor layers (litter (L), fermentation layer (F) and humified layer (H)) under birch and spruce showed higher rates of CO2 production, greater net mineralisation of nitrogen and higher amounts of carbon and nitrogen in microbial biomass than did the forest floor layers under pine. Concentrations of mono-, sesqui-, di- and triterpenes were higher under both conifers than under birch, while the concentration of total water-soluble phenolic compounds as well as the concentration of condensed tannins tended to be higher or at least as high under spruce as under birch or pine. In general, differences between tree species in soil microbial activities and in concentrations of secondary compounds were smaller in the H layer than in the upper layers. The rate of CO2 production and the amount of carbon in the microbial biomass correlated highly positively with the concentration of total water-soluble phenolic compounds and positively with the concentration of condensed tannins. Exposure of soil to volatile monoterpenes and tannins extracted and fractionated from spruce and pine needles affected carbon and nitrogen transformations in soil, but the effects were dependent on the compound and its molecular structure. Monoterpenes decreased net mineralisation of nitrogen and probably had a toxic effect on part of the microbial population in soil, while another part of the microbes seemed to be able to use monoterpenes as a carbon source. With tannins, low-molecular-weight compounds (also compounds other than tannins) increased soil CO2 production and nitrogen immobilisation by soil microbes while the higher-molecular-weight condensed tannins had inhibitory effects. In conclusion, plant secondary compounds may have a great potential in regulation of C and N transformations in forest soils, but the real magnitude of their significance in soil processes is impossible to estimate.

Suomen maankuoren kivilajikoostumuksen tulkinta seismisistä aineistoista

Aim of this study is to investigate composition of the crust in Finland using seismic wide-angle velocity models and laboratory measurements on P- and S-wave velocities of different rock types. The velocities adopted from wide-angle velocity models were compared with laboratory velocities of different rock types corrected for the crustal PT conditions in the study area. The wide-angle velocity models indicate that the P-wave velocity does not only increase step-wise at boundaries of major crustal layers, but there is also gradual increase of velocity within the layers. On the other hand, the laboratory measurements of velocities indicate that no single rock type is able to provide the gradual downward increasing trends. Thus, there must be gradual vertical changes in rock composition. The downward increase of velocities indicates that the composition of the crust becomes gradually more mafic with increasing depth. Even though single rock types cannot simulate the wide-angle model velocities, it can be done with a mixture of rock types. There are a large number of rock type mixtures giving the correct P-wave velocities. Therefore, the inverse solution of rock types and their proportions from velocities is a non-unique problem if only P-wave velocities is available. Amount of the possible rock type mixtures can be limitted using S-wave velocities, reflection seismic results and other geological and geophysical results of the study area. Crustal model FINMIX-2 is presented in this study and it suggest that the crustal velocity profiles can be simulated with rock type mixtures, where the upper crust consists of felsic gneisses and granitic-granodioritic rocks with a minor contribution of quartzite, amphibolite and diabase. In the middle crust the amphibolite proportion increases. The lower crust consists of tonalitic gneiss, mafic garnet granulite, hornblendite, pyroxenite and minor mafic eclogite. This composition model is in agreement with deep crustal kimberlite-hosted xenolith data in eastern Finland and reflectivity of the FIRE (Finnish Reflection Experiment). According to FINMIX-2 model the Moho is deeper and the crustal composition is a more mafic than an average global continental model would suggest. Composition models of southern Finland are quite similar than FINMIX-2 model. However, there are minor differencies between the models, which indicates areal differences of composition. Models of northern Finland shows that the crustal thickness is smaller than southern Finland and composition of the upper crust is different. Density profiles calculated from the lithological models suggest that there is practically no density contrast at Moho in areas of the high-velocity lower crust. This implies that crustal thickness in the central Fennoscandian Shield may have been controlled by the densities of the lower crustal and upper mantle rocks.

Aika ennen aluepolitiikkaa. Pohjoisen kehityshankkeet ja alueiden kehittämisen idea 1950-luvun Suomessa

The dissertation focuses on the recognition of the problems of uneven regional development in Finland in the 1950s, and the way the idea of controlling this development was introduced to Finnish politics. Since it is often stated that Finnish regional policy only began in the mid-1960s, the period at hand is considered to fall in the time before regional policy. However, various ideas, plans and projects of regional development as well as different aims of development were brought forward and discussed already in the 1950s. These give an interesting perspective to the ideas of later regional development. In the 1950s, many Finnish politicians became more conscious of the unavoidable societal change. The need for overall modernisation of the society made it reasonable to expect a growing level of unemployment and eagerness to migration. The uneven distribution of well-being was also feared to cause discontent and political changes. International experience proved interfering in the regional development possible when using the argument of public interest ; the measures taken increased the level of well-being, helped sustain societal balance, and supported national economy. Many of the development projects of the 1950s focused on Northern Finland, the natural resources of which were considered an important reserve and the political climate of which was regarded unstable. After the late 1940s, regional development was discussed frequently both on the national and the regional level. Direct and indirect support was given to less developed areas and the government outlined thorough investigations in order to relieve the regional problem. Towards the end of the decade, the measures taken were already often connected to the idea of equality. In the 1950s the conflicts within and between the largest Finnish political parties significantly affected the decisions of regional development. There are three case studies in this qualitative research based on the narrative method. The case studies clarify the characteristics of the 1950s regional development. In the first one, the representatives of the northern region and the state first discuss the location of a state-run nitrogen fertilizer factory and later the location of a new university. In the second, the aims and perspectives of private entrepreneurs and the state collide due to ideas of statist industrialisation projects and later due to an idea of a tax relief targeting northern industry. In the third case, the main role is given to the changing rural areas, in relation to which societal development and urbanisation were often measured. The regional development of the 1950s laid groundwork for the new, more established regional policy. The early problem solving actions were aimed both at the prevailing situation and the future and thus showed the way for the upcoming actions. Regional development policy existed already before regional policy.

Dental Service Utilization, Dental Health Production and Equity in Dental Care: the Finnish Experience

This thesis is grounded on four articles. Article I generally examines the factors affecting dental service utilization. Article II studies the factors associated with sector-specific utilization among young adults entitled to age-based subsidized dental care. Article III explores the determinants of dental ill-health as measured by the occurrence of caries and the relationship between dental ill-health and dental care use. Article IV measures and explains income-related inequality in utilization. Data employed were from the 1996 Finnish Health Care Survey (I, II, IV) and the 1997 follow-up study included in the longitudinal study of the Northern Finland 1966 Birth Cohort (III). Utilization is considered as a multi-stage decision-making process and measured as the number of visits to the dentist. Modified count data models and concentration and horizontal equity indices were applied. Dentist s recall appeared very efficient at stimulating individuals to seek care. Dental pain, recall, and the low number of missing teeth positively affected utilization. Public subvention for dental care did not seem to statistically increase utilization. Among young adults, a perception of insufficient public service availability and recall were positively associated with the choice of a private dentist, whereas income and dentist density were positively associated with the number of visits to private dentists. Among cohort females, factors increasing caries were body mass index and intake of alcohol, sugar, and soft drinks and those reducing caries were birth weight and adolescent school achievement. Among cohort males, caries was positively related to the metropolitan residence and negatively related to healthy diet and education. Smoking increased caries, whereas regular teeth brushing, regular dental attendance and dental care use decreased caries. We found equity in young adults utilization but pro-rich inequity in the total number of visits to all dentists and in the probability of visiting a dentist for the whole sample. We observed inequity in the total number of visits to the dentist and in the probability of visiting a dentist, being pro-poor for public care but pro-rich for private care. The findings suggest that to enhance equal access to and use of dental care across population and income groups, attention should focus on supply factors and incentives to encourage people to contact dentists more often. Lowering co-payments and service fees and improving public availability would likely increase service use in both sectors. To attain favorable oral health, appropriate policies aimed at improving dental health education and reducing the detrimental effects of common risk factors on dental health should be strengthened. Providing equal access with respect to need for all people ought to take account of the segmentation of the service system, with its two parallel delivery systems and different supplier incentives to patients and dentists.

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.

The role of lakes in carbon cycling in boreal catchments

Lakes are an important component of ecosystem carbon cycle through both organic carbon sequestration and carbon dioxide and methane emissions, although they cover only a small fraction of the Earth's surface area. Lake sediments are considered to be one of rather perma-nent sinks of carbon in boreal regions and furthermore, freshwater ecosystems process large amounts of carbon originating from terrestrial sources. These carbon fluxes are highly uncer-tain especially in the changing climate. -- The present study provides a large-scale view on carbon sources and fluxes in boreal lakes situated in different landscapes. We present carbon concentrations in water, pools in lake se-diments, and carbon gas (CO2 and CH4) fluxes from lakes. The study is based on spatially extensive and randomly selected Nordic Lake Survey (NLS) database with 874 lakes. The large database allows the identification of the various factors (lake size, climate, and catchment land use) determining lake water carbon concentrations, pools and gas fluxes in different types of lakes along a latitudinal gradient from 60oN to 69oN. Lakes in different landscapes vary in their carbon quantity and quality. Carbon (C) content (total organic and inorganic carbon) in lakes is highest in agriculture and peatland dominated areas. In peatland rich areas organic carbon dominated in lakes but in agricultural areas both organic and inorganic C concentrations were high. Total inorganic carbon in the lake water was strongly dependent on the bedrock and soil quality in the catchment, especially in areas where human influence in the catchment is low. In inhabited areas both agriculture and habitation in the catchment increase lake TIC concentrations, since in the disturbed soils both weathering and leaching are presumably more efficient than in pristine areas. TOC concentrations in lakes were related to either catchment sources, mainly peatlands, or to retention in the upper watercourses. Retention as a regulator of the TOC concentrations dominated in southern Finland, whereas the peatland sources were important in northern Finland. The homogeneous land use in the north and the restricted catchment sources of TOC contribute to the close relationship between peatlands and the TOC concentrations in the northern lakes. In southern Finland the more favorable climate for degradation and the multiple sources of TOC in the mixed land use highlight the importance of retention. Carbon processing was intensive in the small lakes. Both CO2 emission and the Holocene C pool in sediments per square meter of the lake area were highest in the smallest lakes. How-ever, because the total area of the small lakes on the areal level is limited, the large lakes are important units in C processing in the landscape. Both CO2 and CH4 concentrations and emissions were high in eutrophic lakes. High availability of nutrients and the fresh organic matter enhance degradation in these lakes. Eutrophic lakes are often small and shallow, enabling high contact between the water column and the sediment. At the landscape level, the lakes in agricultural areas are often eutrophic due to fertile soils and fertilization of the catchments, and therefore they also showed the highest CO2 and CH4 concentrations. Export from the catchments and in-lake degradation were suggested to be equally important sources of CO2 and CH4 in fall when the lake water column was intensively mixed and the transport of sub-stances from the catchment was high due to the rainy season. In the stagnant periods, especially in the winter, in-lake degradation as a gas source was highlighted due to minimal mixing and limited transport of C from the catchment. The strong relationship between the annual CO2 level of lakes and the annual precipitation suggests that climate change can have a major impact on C cycling in the catchments. Increase in precipitation enhances DOC export from the catchments and leads to increasing greenhouse gas emissions from lakes. The total annual CO2 emission from Finnish lakes was estimated to be 1400 Gg C a-1. The total lake sediment C pool in Finland was estimated to be 0.62 Pg, giving an annual sink in Finnish lakes of 65 Gg C a-1.

Export of organic matter, sulphate and base cations from boreal headwater catchments downstream to the coast: impacts of land use and climate

The terrestrial export of dissolved organic matter (DOM) is associated with climate, vegetation and land use, and thus is under the influence of climatic variability and human interference with terrestrial ecosystems, their soils and hydrological cycles. The present study provides an assessment of spatial variation of DOM concentrations and export, and interactions between DOM, catchment characteristics, land use and climatic factors in boreal catchments. The influence of catchment characteristics, land use and climatic drivers on the concentrations and export of total organic carbon (TOC), total organic nitrogen (TON) and dissolved organic phosphorus (DOP) was estimated using stream water quality, forest inventory and climatic data from 42 Finnish pristine forested headwater catchments, and water quality monitoring, GIS land use, forest inventory and climatic data from the 36 main Finnish rivers (and their sub-catchments) flowing to the Baltic Sea. Moreover, the export of DOM in relation to land use along a European climatic gradient was studied using river water quality and land use data from four European areas. Additionally, the role of organic and minerogenic acidity in controlling pH levels in Finnish rivers and pristine streams was studied by measuring organic anion, sulphate (SO4) and base cation (Ca, Mg, K and Na) concentrations. In all study catchments, TOC was a major fraction of DOM, with much lower proportions of TON and DOP. Moreover, most of TOC and TON was in a dissolved form. The correlation between TOC and TON concentrations was strong and TOC concentrations explained 78% of the variation in TON concentrations in pristine headwater streams. In a subgroup of 20 headwater catchments with similar climatic conditions and low N deposition in eastern Finland, the proportion of peatlands in the catchment and the proportion of Norway spruce (Picea abies Karsten) of the tree stand had the strongest correlation with the TOC and TON concentrations and export. In Finnish river basins, TOC export increased with the increasing proportion of peatland in the catchment, whereas TON export increased with increasing extent of agricultural land. The highest DOP concentrations and export were recorded in river basins with a high extent of agricultural land and urban areas, reflecting the influence of human impact on DOP loads. However, the most important predictor for TOC, TON and DOP export in Finnish rivers was the proportion of upstream lakes in the catchment. The higher the upstream lake percentage, the lower the export indicating organic matter retention in lakes. Molar TOC:TON ratio decreased from headwater catchments covered by forests and peatlands to the large river basins with mixed land use, emphasising the effect of the land use gradient on the stoichiometry of rivers. This study also demonstrated that the land use of the catchments is related to both organic and minerogenic acidity in rivers and pristine headwater streams. Organic anion dominated in rivers and streams situated in northern Finland, reflecting the higher extent of peatlands in these areas, whereas SO4 dominated in southern Finland and on western coastal areas, where the extent of fertile areas, agricultural land, urban areas, acid sulphate soils, and sulphate deposition is highest. High TOC concentrations decreased pH values in the stream and river water, whereas no correlation between SO4 concentrations and pH was observed. This underlines the importance of organic acids in controlling pH levels in Finnish pristine headwater streams and main rivers. High SO4 concentrations were associated with high base cation concentrations and fertile areas, which buffered the effects of SO4 on pH.

Decomposition of Scots pine fine woody debris in boreal conditions

"Litter quality and environmental effects on Scots pine (Pinus sylvestris L.) fine woody debris (FWD) decomposition were examined in three forestry-drained peatlands representing different site types along a climatic gradient from the north boreal (Northern Finland) to south (Southern Finland) and hemiboreal (Central Estonia) conditions. Decomposition (percent mass loss) of FWD with diameter <= 10 mm (twigs) and FWD with diameter > 10 mm (branches) was measured using the litter bag method over 1-4-year periods. Overall, decomposition rates increased from north to south, the rate constants (k values) varying from 0.128 to 0.188 year(-1) and from 0.066 to 0.127 year(-1) for twigs and branches, respectively. On average, twigs had lost 34%, 19% and 19%, and branches 25%, 17% and 11% of their initial mass after 2 years of decomposition at the hemiboreal, south boreal and north boreal sites, respectively. After 4 years at the south boreal site the values were 48% for twigs and 42% for branches. Based on earlier studies, we suggest that the decomposition rates that we determined may be used for estimating Scots pine FWD decomposition in the boreal zone, also in upland forests. Explanatory models accounted for 50.4% and 71.2% of the total variation in FWD decomposition rates when the first two and all years were considered, respectively. The variables most related to FWD decomposition included the initial ash, water extractives and Klason lignin content of litter, and cumulative site precipitation minus potential evapotranspiration. Simulations of inputs and decomposition of Scots pine FWD and needle litter in south boreal conditions over a 60-year period showed that 72 g m(-2) of organic matter from FWD vs. 365 g m(-2) from needles accumulated in the forest floor. The annual inputs varied from 5.7 to 15.6 g m(-2) and from 92 to 152 g m(-2) for FWD and needles, respectively. Each thinning caused an increase in FWD inputs, Up to 510 g m(-2), while the needle inputs did not change dramatically. Because the annual FWD inputs were lowered following the thinnings, the overall effect of thinnings on C accumulation from FWD was slightly negative. The contribution of FWD to soil C accumulation, relative to needle litter, seems to be rather minor in boreal Scots pine forests. (C) 2008 Elsevier B.V. All rights reserved."

Kemihaaran allasalueen luonto

Nature in the Kemihaara reservoir area, Northern Finland.