Novel Multiple Sclerosis Predisposing Genetic Variants Outside the HLA Region

Multiple sclerosis (MS) is a chronic inflammatory disease of the central nervous system (CNS). Both environmental factors and several predisposing genes are required to generate MS. Despite intensive research these risk factors are still largely unknown, the pathogenesis of MS demyelination is poorly understood, and no curative treatment exists. Both prevalence and familial occurrence of MS are exceptionally high in a Finnish population subisolate, Southern Ostrobothnia, presumably due to enrichment of predisposing genetic variants within this region. Previous linkage scan on MS pedigrees from Southern Ostrobothnia detected three main MS loci on chromosomes 5p, 6p (HLA) and 17q. Linkage studies in other populations have also provided independent evidence for the location of MS susceptibility genes in these regions. Further, these loci are syntenic to the experimental autoimmune encephalomyelitis (EAE) susceptibility loci of rodents. In this thesis work an effort was made to localize MS predisposing alleles of the linked loci outside the HLA region by studying familial MS cases from the Southern Ostrobothnia isolate. Analysis of the 5p locus revealed one region, flanking the complement component 7 (C7) gene. The identified relatively rare haplotype seems to have a fairly large effect on genetic susceptibility of MS (frequency MS 12%, controls 4%; p=0.000003, OR=2.73). Evidence for association with alleles of the region and MS was seen also in more heterogeneous populations. Convincingly, plasma C7 protein levels and complement activity correlated with the risk haplotype identified. The finding stimulated us to study other complement cascade genes in MS. No evidence for association could be observed with the complement component coding genes outside 5p. A scan of the 17q locus provided evidence for association with variants of the protein kinase C alpha (PRKCA) gene (p=0.0001). Modest evidence for association with PRKCA was observed also in Canadian MS families. Finally we used a candidate gene based approach to identify potential MS loci. Mutations of DAP12 and TREM2 cause a recessively inherited CNS white matter disease PLOSL. Interestingly, DAP12 and TREM2 are located in MS regions on 6p and 19q, and we tested them as potential candidate genes in the Finnish MS sample. No evidence for association with MS was observed. This thesis provides an example of how extended families from special populations can be utilized in fine-mapping of the linked loci. A first relatively rare MS variant was identified utilizing the strength of a Finnish population subisolate. This variant seems to have an effect on activity of the complement system, which has previously been suggested to have an important role in the pathogenesis of MS.

Sindbis virus and Pogosta disease in Finland

Sindbis virus (SINV) (genus Alphavirus, family Togaviridae) is an enveloped virus with a genome of single-stranded, positive-polarity RNA of 11.7 kilobases. SINV is widespread in Eurasia, Africa, and Australia, but clinical infection only occurs in a few geographically restricted areas, mainly in Northern Europe. In Europe, antibodies to SINV were detected from patients with fever, rash, and arthritis for the first time in the early 1980s in Finland. It became evident that the causative agent of this syndrome, named Pogosta disease, was closely related to SINV. The disease is also found in Sweden (Ockelbo disease) and in Russia (Karelian fever). Since 1974, for unknown reason, the disease has occurred as large outbreaks every seven years in Finland. This study is to a large degree based on the material collected during the 2002 Pogosta disease outbreak in Finland. We first developed SINV IgM and IgG enzyme immunoassays (EIA), based on highly purified SINV, to be used in serodiagnostics. The EIAs correlated well with the hemagglutination inhibition (HI) test, and all individuals showed neutralizing antibodies. The sensitivities of the IgM and IgG EIAs were 97.6% and 100%, and specificities 95.2% and 97.6%, respectively. E1 and E2 envelope glycoproteins of SINV were shown to be recognized by IgM and IgG in the immunoblot early in infection. We isolated SINV from five patients with acute Pogosta disease; one virus strain was recovered from whole blood, and four other strains from skin lesions. The etiology of Pogosta disease was confirmed by these first Finnish SINV strains, also representing the first human SINV isolates from Europe. Phylogenetic analysis indicated that the Finnish SINV strains clustered with the strains previously isolated from mosquitoes in Sweden and Russia, and seemed to have a common ancestor with South-African strains. Northern European SINV strains could be maintained locally in disease-endemic regions, but the phylogenetic analysis also suggests that redistribution of SINV tends to occur in a longitudinal direction, possibly with migratory birds. We searched for SINV antibodies in resident grouse (N=621), whose population crashes have previously coincided with human SINV outbreaks, and in migratory birds (N=836). SINV HI antibodies were found for the first time in birds during their spring migration to Northern Europe, from three individuals: red-backed shrike, robin, and song thrush. Of the grouse, 27.4% were seropositive in 2003, one year after a human outbreak, but only 1.4% of the grouse were seropositive in 2004. Thus, grouse might contribute to the human epidemiology of SINV. A total of 86 patients with verified SINV infection were recruited to the study in 2002. SINV RNA detection or virus isolation from blood and/or skin lesions was successful in eight patients. IgM antibodies became detectable within the first eight days of illness, and IgG within 11 days. The acute phase of Pogosta disease was characterized by arthritis, itching rash, fatigue, mild fever, headache, and muscle pain. Half of the patients reported in self-administered questionnaires joint symptoms to last > 12 months. Physical examination in 49 of these patients three years after infection revealed persistent joint manifestations. Arthritis (swelling and tenderness in physical examination) was diagnosed in 4.1% (2/49) of the patients. Tenderness in palpation or in movement of a joint was found in 14.3% of the patients in the rheumatologic examination, and additional 10.2% complained persisting arthralgia at the interview. Thus, 24.5% of the patients had joint manifestations attributable to the infection three years earlier. A positive IgM antibody response persisted in 3/49 of the patients; both two patients with arthritis were in this group. Persistent symptoms of SINV infection might have considerable public health implications in areas with high seroprevalence. The age-standardized seroprevalence of SINV (1999-2003, N=2529) in the human population in Finland was 5.2%. The seroprevalence was high in North Karelia, Kainuu, and Central Ostrobothnia. The incidence was highest in North Karelia. Seroprevalence in men (6.0%) was significantly higher than in women (4.1%), however, the average annualized incidence in the non-epidemic years was higher in women than in men, possibly indicating that infected men are more frequently asymptomatic. The seroprevalence increased with age, reaching 15.4% in persons aged 60-69 years. The incidence was highest in persons aged 50-59 years.

Dengue virus infection : Diagnostics and molecular epidemiology

Dengue is a mosquito-borne viral disease caused by the four dengue virus serotypes (DENV-1-4) and is currently considered as the most important arthropod-borne viral disease in the world. Nearly half of the human population lives in risk areas, and 50-100 million infections occur yearly according to World Health Organization. The disease can vary from a mild febrile disease to severe haemorrhagic fever and shock. A secondary infection with heterologous serotype increases the risk for severe disease outcome. During the last three decades the impact of dengue has dramatically increased in the endemic areas including the tropics and subtropics of the world. The current situation with massive epidemics of severe disease forms has been associated with socio-ecological changes that have increased the transmission and enabled the co-circulation of different serotypes. Consequently, an increase of dengue has also been observed in travelers visiting these areas. Currently approximately 30 cases are diagnosed yearly in Finnish travelers. In travelers dengue is rarely a life-threatening disease, however in the current study, a fatality was documented in a young Finnish patient who experienced a prolonged primary dengue infection. To improve particularly early laboratory diagnostics, a novel real-time RT-PCR method was developed for the detection of DENV-1-4 RNA based on TaqMan chemistry. The method was shown to be sensitive and specific for detecting DENV RNA and suitable for diagnostic use. The newly developed real-time RT-PCR was compared to other available early diagnostic methods including IgM and NS1 antigen detection using a panel of selected patient samples. The results suggest that the best diagnostic rates are achieved by a combination of IgM with RNA or NS1 detection. The dengue virus strains studied here included the first DENV strains isolated from serum samples of Finnish travelers collected in 2000-2005. The results of sequence analysis demonstrated that the 11 isolates included all four DENV serotypes and presented a global sample of DENV strains from different geographical areas including Asia, Africa and South America. In the present study sequence analysis was also carried out for a collection of 23 novel DENV-2 isolates from Venezuelan patients collected in 1999-2005. The Venezuelan DENV-2 exclusively represented the American-Asian genotype, suggesting that no foreign DENV-2 lineages have recently been introduced to the country. The results also suggest that the DENV-2 viruses detected earlier from Venezuela have been maintained in the area where they have evolved into several lineages. This is in contrast to the pattern observed in some other dengue endemic areas, where introductions of novel virus types and lineages are frequently detected.

Is prosopis a curse or a blessing? : An ecological economic analysis of an invasive alien tree species in Sudan.

The issue of the usefulness of different prosopis species versus their status as weeds is a matter of hot debate around the world. The tree Prosopis juliflora had until 2000 been proclaimed weedy in its native range in South America and elsewhere in the dry tropics. P. juliflora or mesquite has a 90-year history in Sudan. During the early 1990s a popular opinion in central Sudan and the Sudanese Government had begun to consider prosopis a noxious weed and a problematic tree species due to its aggressive ability to invade farmlands and pastures, especially in and around irrigated agricultural lands. As a consequence prosopis was officially declared an invasive alien species also in Sudan, and in 1995 a presidential decree for its eradication was issued. Using a total economic valuation (TEV) approach, this study analysed the impacts of prosopis on the local livelihoods in two contrasting irrigated agricultural schemes. Primarily a problem-based approach was used in which the derivation of non-market values was captured using ecological economic tools. In the New Halfa Irrigation Scheme in Kassala State, four separate household surveys were conducted due to diversity between the respective population groups. The main aim was here to study the magnitude of environmental economic benefits and costs derived from the invasion of prosopis in a large agricultural irrigation scheme on clay soil. Another study site, the Gandato Irrigation Scheme in River Nile State represented impacts from prosopis that an irrigation scheme was confronted with on sandy soil in the arid and semi-arid ecozones along the main River Nile. The two cases showed distinctly different effects of prosopis but both indicated the benefits to exceed the costs. The valuation on clay soil in New Halfa identified a benefit/cost ratio of 2.1, while this indicator equalled 46 on the sandy soils of Gandato. The valuation results were site-specific and based on local market prices. The most important beneficial impacts of prosopis on local livelihoods were derived from free-grazing forage for livestock, environmental conservation of the native vegetation, wood and non-wood forest products, as well as shelterbelt effects. The main social costs from prosopis were derived from weeding and clearing it from farm lands and from canalsides, from thorn injuries to humans and livestock, as well as from repair expenses vehicle tyre punctures. Of the population groups, the tenants faced most of the detrimental impacts, while the landless population groups (originating from western and eastern Sudan) as well as the nomads were highly dependent on this tree resource. For the Gandato site the monetized benefit-cost ratio of 46 still excluded several additional beneficial impacts of prosopis in the area that were difficult to quantify and monetize credibly. In River Nile State the beneficial impact could thus be seen as completely outweighing the costs of prosopis. The results can contributed to the formulation of national and local forest and agricultural policies related to prosopis in Sudan and also be used in other countries faced with similar impacts caused by this tree.

From a tree to a stand in Finnish boreal forests: biomass estimation and comparison of methods

There is an increasing need to compare the results obtained with different methods of estimation of tree biomass in order to reduce the uncertainty in the assessment of forest biomass carbon. In this study, tree biomass was investigated in a 30-year-old Scots pine (Pinus sylvestris) (Young-Stand) and a 130-year-old mixed Norway spruce (Picea abies)-Scots pine stand (Mature-Stand) located in southern Finland (61º50' N, 24º22' E). In particular, a comparison of the results of different estimation methods was conducted to assess the reliability and suitability of their applications. For the trees in Mature-Stand, annual stem biomass increment fluctuated following a sigmoid equation, and the fitting curves reached a maximum level (from about 1 kg/yr for understorey spruce to 7 kg/yr for dominant pine) when the trees were 100 years old. Tree biomass was estimated to be about 70 Mg/ha in Young-Stand and about 220 Mg/ha in Mature-Stand. In the region (58.00-62.13 ºN, 14-34 ºE, ≤ 300 m a.s.l.) surrounding the study stands, the tree biomass accumulation in Norway spruce and Scots pine stands followed a sigmoid equation with stand age, with a maximum of 230 Mg/ha at the age of 140 years. In Mature-Stand, lichen biomass on the trees was 1.63 Mg/ha with more than half of the biomass occurring on dead branches, and the standing crop of litter lichen on the ground was about 0.09 Mg/ha. There were substantial differences among the results estimated by different methods in the stands. These results imply that a possible estimation error should be taken into account when calculating tree biomass in a stand with an indirect approach.

Fen ecosystem carbon gas dynamics in changing hydrological conditions

Northern peatlands are thought to store one third of all soil carbon (C). Besides the C sink function, peatlands are one of the largest natural sources of methane (CH4) to the atmosphere. Climate change may affect the C gas dynamics as well as the labile C pool. Because the peatland C sequestration and CH4 emissions are governed by high water levels, changes in hydrology are seen as the driving factor in peatland ecosystem change. This study aimed to quantify the carbon dioxide (CO2) and CH4 dynamics of a fen ecosystem at different spatial scales: plant community components scale, plant community scale and ecosystem scale, under hydrologically normal and water level drawdown conditions. C gas exchange was measured in two fens in southern Finland applying static chamber and eddy covariance techniques. During hydrologically normal conditions, the ecosystem was a CO2 sink and CH4 source to the atmosphere. Sphagnum mosses and sedges were the most important contributors to the community photosynthesis. The presence of sedges had a major positive impact on CH4 emissions while dwarf shrubs had a slightly attenuating impact. C fluxes varied considerably between the plant communities. Therefore, their proportions determined the ecosystem scale fluxes. An experimental water level drawdown markedly reduced the photosynthesis and respiration of sedges and Sphagnum mosses and benefited shrubs. Consequently, changes were smaller at the ecosystem scale than at the plant group scale. The decrease in photosynthesis and the increase in respiration, mostly peat respiration, made the fen a smaller CO2 sink. CH4 fluxes were significantly lowered, close to zero. The impact of natural droughts was similar to, although more modest than, the impact of the experimental water level drawdown. The results are applicable to the short term impacts of the water level drawdown and to climatic conditions in which droughts become more frequent.

Temperature sensitivity of soil organic matter decomposition in boreal soils

The temperature sensitivity of decomposition of different soil organic matter (SOM) fractions was studied with laboratory incubations using 13C and 14C isotopes to differentiate between SOM of different age. The quality of SOM and the functionality and composition of microbial communities in soils formed under different climatic conditions were also studied. Transferring of organic layers from a colder to a warmer climate was used to assess how changing climate, litter input and soil biology will affect soil respiration and its temperature sensitivity. Together, these studies gave a consistent picture on how warming climate will affect the decomposition of different SOM fractions in Finnish forest soils: the most labile C was least temperature sensitive, indicating that it is utilized irrespective of temperature. The decomposition of intermediate C, with mean residence times from some years to decades, was found to be highly temperature sensitive. Even older, centennially cycling C was again less temperature sensitive, indicating that different stabilizing mechanisms were limiting its decomposition even at higher temperatures. Because the highly temperature sensitive, decadally cycling C, forms a major part of SOM stock in the organic layers of the studied forest soils, these results mean that these soils could lose more carbon during the coming years and decades than estimated earlier. SOM decomposition in boreal forest soils is likely to increase more in response to climate warming, compared to temperate or tropical soils, also because the Q10 is temperature dependent. In the northern soils the warming will occur at a lower temperature range, where Q10 is higher, and a similar increase in temperature causes a higher relative increase in respiration rates. The Q10 at low temperatures was found to be inversely related to SOM quality. At higher temperatures respiration was increasingly limited by low substrate availability.

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.

Owner-driven decision support in holding-specific forest planning

Socio-economic and demographic changes among family forest owners and demands for versatile forestry decision aid motivated this study, which sought grounds for owner-driven forest planning. Finnish family forest owners’ forest-related decision making was analyzed in two interview-based qualitative studies, the main findings of which were surveyed quantitatively. Thereafter, a scheme for adaptively mixing methods in individually tailored decision support processes was constructed. The first study assessed owners’ decision-making strategies by examining varying levels of the sharing of decision-making power and the desire to learn. Five decision-making modes – trusting, learning, managing, pondering, and decisive – were discerned and discussed against conformable decision-aid approaches. The second study conceptualized smooth communication and assessed emotional, practical, and institutional boosters of and barriers to such smoothness in communicative decision support. The results emphasize the roles of trust, comprehension, and contextual services in owners’ communicative decision making. In the third study, a questionnaire tool to measure owners’ attitudes towards communicative planning was constructed by using trusting, learning, and decisive dimensions. Through a multivariate analysis of survey data, three owner groups were identified as fusions of the original decision-making modes: trusting learners (53%), decisive learners (27%), and decisive managers (20%). Differently weighted communicative services are recommended for these compound wishes. The findings of the studies above were synthesized in a form of adaptive decision analysis (ADA), which allows and encourages the decision-maker (owner) to make deliberate choices concerning the phases of a decision aid (planning) process. The ADA model relies on adaptability and feedback management, which foster smooth communication with the owner and (inter-)organizational learning of the planning institution(s). The summarized results indicate that recognizing the communication-related amenity values of family forest owners may be crucial in developing planning and extension services. It is therefore recommended that owners, root-level planners, consultation professionals, and pragmatic researchers collaboratively continue to seek stable change.

Läksi puut ylenemähän : Metsien hoidon historia Suomessa keskiajalta metsäteollisuuden läpimurtoon 1870-luvulla

Tutkimuksen tavoitteena on historiantutkimuksen menetelmiä käyttäen selvittää metsänhoidon teorian sekä käytännön metsänhoidon kehitysvaiheet ja näihin vaikuttaneet tekijät, keskiajalta lähtien 1870-luvulla tapahtuneeseen metsäteollisuuden läpimurtoon saakka. Tutkimus tarkastelee Suomen metsiä, niiden käyttöä ja metsänhoidon alkuvaiheita Ruotsin ja Venäjän vallan aikoina. Vastauksia haetaan erityisesti seuraaviin pääkysymyksiin: - miten eri metsänkäyttömuodot ja -käyttäjät vaikuttivat metsiin ja metsänhoidon edistymiseen? - millä tavoin maanomistuksen kehitys vaikutti metsien käyttöön ja hoitoon? - millaisiin päämääriin ja yhteiskunnallisiin taustatekijöihin metsien käytön julkinen ohjaus perustui? - mitä käytännön vaikutuksia valtion metsäpolitiikalla ja ohjauksella oli metsänhoidon kehitykseen? - missä ja miten kehittyivät Suomessa sovellettu metsänhoidon teoria ja käytännön menetelmät? - mitkä tekijät säätelivät metsänhoidon teorioiden soveltamista käytäntöön? - mikä oli naapurimaiden metsänhoidon sekä kansainvälisten yhteyksien merkitys metsänhoidon kehitykselle Suomessa? - miten vuosisatainen pelko metsien ja puun loppumisesta vaikutti metsänhoidon kehitykseen? - millainen merkitys puun arvon kehityksellä oli metsänhoidon alkuun saattamiselle ja edistymiselle? Suomessa harjoitettiin 1870-luvulle saakka pääasiassa talonpoikaista metsänkäyttöä. Maaseudun väestö hankki toimeentulonsa metsistä eränkäynnin, kaskiviljelyn, laiduntamisen, rakennushirsien valmistamisen, tervantuotannon ja paikoin myös potaskan tai sysien valmistamisen avulla. Erityisesti rannikkoseuduilla tuotettiin "isorakennuksen puita", lehtereitä, mastopuita ja muuta erikoispuutavaraa. Lautojen ja lankkujen sahaus laajeni vähitellen, saavuttaen 1800-luvun lopulla hallitsevan aseman myyntiin tarkoitettujen metsäntuotteiden tuotannossa. Polttopuun sekä muun kotitarvepuun kulutus säilyi suurimpana puunkäytön ryhmänä pitkälle 1900-luvulle saakka. Mainituista metsänkäyttömuodoista erityisesti kaskeaminen ja sitä seuraava laiduntamisvaihe sekä tervaspuiden koloaminen "autioittivat" laajoja metsäalueita. Tiheimmin asutuilla seuduilla esiintyi pulaa poltto- ja rakennuspuusta myöhäiskeskiajalta alkaen. Nämä ongelmat sekä laivanrakennuksen ja vuoriteollisuuden puunsaannin turvaamisen tarve johtivat 1600-luvun puolivälissä pysyvään metsänkäytön julkiseen ohjaukseen. Tuolloin Ruotsin valtakunnan metsälainsäädännön kivijalaksi tuli kestävyyden periaate, josta kruunu kylläkin joutui tinkimään moneen otteeseen. Valtion jatkuva rahantarve oli käytännössä metsäpolitiikan tärkein taustavoima sekä Ruotsin vallan että autonomian aikana. Jo 1600-luvulla ruvettiin vaatimaan talonpoikien yhteismaiden jakamista omistajilleen vastuullisemman metsänkäytön nimissä. Isoajakoa saatiin Suomessa odottaa 1770-luvulle saakka. Etelä-Suomessa se valmistui melko nopeasti, 1800-luvun puoliväliin mennessä. Sillä olikin myönteinen, metsien säästävämpään käsittelyyn johtava vaikutus. Valtiosta tuli isonjaon myötä erityisesti Pohjois-Suomessa merkittävä metsänomistaja 1800-luvun jälkipuoliskolla. Valtion metsähallinto, jota maaherrat ja sivistyneistö vaativat perustettavaksi jo 1700-luvun puolivälissä, aloitti toimintansa maanlaajuisesti 1860-luvulla. Se oli ensimmäinen merkittävä metsänhoidon organisaatio, ja vasta sen myötä metsänkäyttöä ohjaavilla säädöksillä ja ohjeilla alkoi olla käytännön merkitystä. Yksityismetsiä varten ei tällaista organisaatiota vielä perustettu, niitä rasittivat pahoin nousevan sahateollisuuden määrämittahakkuut pitkälle 1900-luvun puolelle. Turun Akatemiassa tehtiin mittavaa metsänhoidon menetelmiä koskevaa sekä myös metsäpoliittista tutkimustyötä 1700-luvun jälkipuoliskolla. Tulokset eivät vielä sanottavasti siirtyneet käytäntöön, lähinnä puun alhaisen arvon ja tarvittavien organisaatioiden puuttumisen takia. Kun valtion metsähallintoa ja Suomen omaa metsäopetusta ryhdyttiin perustamaan 1800-luvun puolivälissä, haettiin metsänhoidon mallia alan johtavaksi maaksi kehittyneestä Saksasta. Tultaessa 1870-luvulle, oli Evolla jo käynnissä voimakas kehitystyö maamme olosuhteisiin soveltuvien menetelmien luomiseksi saksalaisen teorian pohjalta. Metsänhoidon tiedot ja taidot olisivat jo tässä vaiheessa riittäneet kestävän metsätalouden harjoittamiseen kaikkien omistajaryhmien metsissä, jos tarvittavat organisaatiot olisi kyetty perustamaan ja metsäammattilaisia olisi koulutettu tarpeeksi. Metsänhoidon kehitystä hidastivat 1800-luvun lopulla lähinnä valtion heikko talous ja poliittiset näkemyserot. Metsäteollisuuden 1870-luvulta alkanut voimakas kasvu ja lisääntyvä puuntarve pakottivat kuitenkin valtiovallan pitämään huolta puuntuotannon jatkuvuudesta. Metsäteollisuuden kasvavan viennin kautta lisääntyvät verotulot ja kan-santalouden myönteinen kehitys antoivat vähitellen mahdollisuuden metsänhoidon edistämiseen ammattilaisten koulutuksen, kansalaisten neuvonnan, lainsäädännön ja viranomaisten toiminnan kautta. Tämä tutkimus lähestyy aihettaan metsähistorian, taloushistorian, yhteiskuntahistorian ja ympäristöhistorian näkökulmista. Ajankohtaista merkitystä sillä on kehitysmaiden sekä Itä-Euroopan siirtymätalouksien metsänhoidon edistämiselle, missä suomalaiset metsäammattilaiset ovat mukana lukuisten kehityshankkeiden asiantuntijoina. Kymmenissä maissa metsätalous kamppailee samanlaisten ongelmien kanssa kuin Suomessa ja naapurimaissa 100 - 300 vuotta sitten. Meidän kokemuksistamme on näille kansantalouksille hyötyä valtion- ja yksityismetsätalouden metsänhoito-organisaatioita sekä metsälainsäädäntöä kehitettäessä.

Maiseman- ja luonnonhoidon taloudelliset vaikutukset Pihlajaveden metsänomistajille

Tutkimuksen tavoitteet Tutkimuksen teoreettinen osa esittelee yhteiskunnallisten hankkeiden arvottamiseen liittyvää koti- ja ulkomaista kirjallisuutta. Empiirinen osa tutkii Savonlinnasta poimitun metsätilaotoksen avulla maiseman- ja luonnonhoidon aiheuttamia taloudellisia vaikutuksia tilojen metsänomistajille sekä vertaa näitä tuloksia haastattelututkimuksesta saatuihin vastauksiin maksuhalukkuudesta monimuotoisuuden hyväksi. Lähdeaineisto Metsämaiseman- ja luonnonhoidon aiheuttamia taloudellisia vaikutuksia tutkittaessa case-aineistona käytettiin keskeisen Saimaan Pihlajaveden saaristoalueen 24 tilaa, joiden omistajat olivat tilanneet Etelä-Savon metsäkeskukselta uudentyyppisen maiseman- ja luonnonhoidon arvot huomioivan metsäsuunnitelman. Samalla aineistolla suoritettiin myös metsänomistajien haastattelu, jonka avulla selvitettiin metsänomistajien maksuhalukkuutta monimuotoisuudesta ja maisemamansuojelusta. Lisäksi kysyttiin mielipiteitä nykyisistä metsänhoitosuosituksista ja alueelle kaavailluista luonnonsuojeluhankkeista. Aineiston käsittely Taloudellisia vaikutuksia tutkittaessa maisema- ja luontoarvoja korostavaa metsäsuunnitelmalaskelmaa verrattiin maksimaaliseen puuntuotantoon tähtäävään metsäsuunnitelmaan. Näiden kahden vaihtoehtoisten laskelman välinen erotus tulkittiin monimuotoisuuden turvaamisen metsälötason puuntuotannolliseksi vaikutukseksi. Vastaavasti haastatteluosuudessa metsänomistajilta kysyttiin, kuinka paljon he olisivat valmiita luopumaan metsätuloistaan monimuotoisuuden edistämiseksi. Maksuhalukkuutta tutkittaessa käytettiin contingent valuation -menetelmää. Tulokset Siirtyminen ns. perinteisestä metsänkäsittelystä maiseman- ja luonnonhoidon huomioivaan metsänkäsittelyyn pienentää metsästä saatavia nettotuloja ensimmäisen kymmenvuotiskauden aikana 36 % eli 289 mk/ha vuosittain. Vastaavasti metsänomistajien maksuhalukkuus monimutoisuuden edistämisestä oli 5,3 % metsätuloista, joka on rahassa 18 mk/ha vuosittain. Nettotulojen merkittävä pieneneminen selittyy kohteen arvokkailla maisema-arvoilla ja ekologisten elinympäristöjen runsaalla lukumäärällä. Tulosten tilastollista luotettavuutta ei otoksen pienuuden vuoksi voitu tarkastella. Avainsanat: metsämaiseman- ja luonnonhoito, metsätulot, maksuhalukkuus, monimuotoisuus, Pihlajaveden saaristo

Metsän kiertoajan vaikutus hiilensidontaan ja metsänkasvatuksen kannattavuuteen

Cost-effective mitigation of climate change is essential for both climate and environmental policy. Forest rotation age is one of the silvicultural measures by which the forest carbon stocks can be influenced with in accordance with the Kyoto Protocol, Article 3.4. The purpose of this study is to evaluate how forest rotation age affects carbon sequestration and the profitability of forestry. The relation between the forest rotation period optimizing forest owners’ discounted net returns over time and rotations which are 10, 20 and 30 years longer than the optimal rotation is examined. In addition, the cost of lengthening the rotation period is studied as well as whether carbon sequestration revenues can improve the profitability of forestry. The data used in the study consist of 16 stands located in Southern Finland. The main tree species in these stands were Norway spruce and Scots pine. Forest simulation tool MOTTI was used in the analysis. The results indicate that by lengthening the rotation period forest carbon stocks increase. However, as the rotation period is lengthened by more than 10 years, as a result of the diminishing growth curve, the rate of carbon sequestration slows down. The average discounted cost of carbon sequestration varied between 2.4 – 14.1 €/tCO2. Carbon sequestration rates in spruce stands were higher and the costs lower than those obtained from pine stands. The absence of carbon trading schemes is an obstacle for the commercialization of forest carbon sinks. In the future, research should concentrate on analysing what kind of operational models of carbon trading could be feasible in Finland.

Energiapuuharvennus osana metsänkasvatusta ja ilmastonmuutoksen hillitsemistä

The aim of this study was to compare the differences between forest management incorporating energy wood thinning and forest management based on silvicultural recommendations (baseline). Energy wood thinning was substituted for young stand thinning and the first commercial thinning of industrial wood. The study was based on the forest stand data from Southern Finland, which were simulated by the MOTTI-simulator. The main interest was to find out the climatic benefits resulting from carbon sequestration and energy substitution. The value of energy wood was set to substitute it for coal as an alternative energy fuel (emission trade). Other political instruments (Kemera subsidies) were also analysed. The largest carbon dioxide emission reductions were achieved as a combination of carbon sequestration and energy substitution (on average, a 26-90 % increase in discounted present value in the beginning of rotation) compared to the baseline. Energy substitution increased emission reductions more effectively than carbon sequestration, when maintaining dense young stands. According to the study, energy wood thinning as a part of forest management was more profitable than the baseline when the value of carbon dioxide averaged more than 15 €/CO2 and other political subsidies were unchanged. Alternatively, the price of energy wood should on average exceed 21 €/m3 on the roadside in order to be profitable in the absence of political instruments. The most cost-efficient employment of energy wood thinning occured when the dominant height was 12 meters, when energy substitution was taken into account. According to alternative forest management, thinning of sapling stands could be done earlier or less intensely than thinning based on silvicultural recommendations and the present criteria of subsidies. Consequently, the first commercial thinning could be profitable to carry out either as harvesting of industrial wood or energy wood, or as integrated harvesting depending on the costs of the harvesting methods available and the price level of small-size industrial wood compared to energy wood.

Effects of seed origin latitude on the timing of height growth cessation and field performance of silver birch

The aim of this thesis was to increase our knowledge about the effects of seed origin on the timing of height growth cessation and field performance of silver birch from different latitudes, with special attention paid to the browsing damage by moose in young birch plantations. The effect of seed origin latitude and sowing time on timing of height growth cessation of first-year seedlings was studied in a greenhouse experiment with seven seed origins (lat. 58º - 67ºN). Variation in critical night length (CNL) for 50 % bud set within two latitudinally distant stands (60º and 67ºN) was studied in three phytotron experiments. Browsing by moose on 5-11 -year-old silver birch saplings from latitudinally different seed origins (53º - 67ºN) was studied in a field experiment in southern Finland. Yield and stem quality of 22-year-old silver birch trees of Baltic, Finnish and Russian origin (54º - 63ºN) and the effect of latitudinal seed transfers were studied in two provenance trials at Tuusula, southern and Viitasaari, central Finland. The timing of height growth cessation depended systematically on latitude of seed origin and sowing date. The more northern the seed origin, the earlier the growth cessation and the shorter the growth period. Later sowing dates delayed growth cessation but also shortened the growth period. The mean CNL of the southern ecotype was longer, 6.3 ± 0.2 h (95 % confidence interval), than that of the northern ecotype, 3.1 ± 0.3 h. Within-ecotype variance of the CNL was higher in the northern ecotype (0.484 h2) than in the southern ecotype (0.150 h2). Browsing by moose decreased with increasing latitude of seed origin and sapling height. Origins transferred from more southern latitudes were more heavily browsed than the more northern native ones. Southern Finnish seed origins produced the highest volume per unit area in central Finland (lat. 63º11'N). Estonian and north Latvian stand seed origins, and the southern Finnish plus tree origins, were the most productive ones in southern Finland (lat. 60º21'N). Latitudinal seed transfer distance had a significant effect on survival, stem volume/ha and proportion of trees with a stem defect. The relationship of both survival and stem volume/ha to the latitudinal seed transfer distance was curvilinear. Volume was increased by transferring seed from ca. 2 degrees of latitude from the south. A longer transfer from the south, and transfer from the north, decreased the yield. The proportion of trees with a stem defect increased linearly in relation to the latitudinal seed transfer distance from the south.

Undersowing in a northern climate: effects on spring cereal yield and risk of nitrate leaching

Disadvantages of invariable cereal cropping, concern of nutrient leaching and prices of nitrogen (N) fertilizer have all increased during last decades. An undersown crop, which grows together with a main crop and after harvest, could mitigate all those questions. The aim of this study was to develop undersowing in Finnish conditions, so that it suits for spring cereal farming as well as possible and enhances taking care of soil and environment, especially when control of N is concerned. In total, 17 plant species were undersown in spring cereals during the field experiments between 1991-1999 at four sites in South and Central Finland, but after selection, eight of them were studied more thoroughly. Two legumes, one grass species and one mixture of them were included in long-term trials in order to study annually repeated undersowing. Further, simultaneous broadcasting of seeds instead of separate undersowing was studied. Grain yield response and the capacity of the undersown crop to absorb soil N or fix N from atmosphere, and the release of N were of greatest interest. Seeding rates of undersown crops and N fertilization rates during annually repeated undersowing were also studied. Italian ryegrass (Lolium multiflorum Lam., IR) absorbed soil nitrate N (NO3-N) most efficiently in autumn and timothy (Phleum pratense L.) in spring. The capacity of other grass species to absorb N was low, or it was insufficient considering the negative effect on grain yield. Red clover (Trifolium pratense L.) and white clover (Trifolium repens L.) suited well in annually repeated undersowing, supplying fixed N for cereals without markedly increased risk of N leaching. Autumn oriented growth rhythm of the studied legumes was optimal for undersowing, whereas the growth rhythm of grasses was less suited but varied between species. A model of adaptive undersowing system was outlined in order to emphasize allocation of measures according needs. After defining the goal of undersowing, many decisions are to be done. When diminishing N leaching is primarily sought, a mixture of IR and timothy is advantageous. Clovers suit for replacing N fertilization, as the positive residual effect is greater than the negative effect caused by competition. A mixture of legume and non legume is a good choice when increased diversity is the main target. Seeding rate is an efficient means for adjusting competition and N effects. Broadcasting with soil covering equipment can be used to establish an undersown crop. In addition, timing and method of cover crop termination have an important role in the outcome. Continuous observing of the system is needed as for instance conditions significantly affect growth of undersown crop and on the other hand N release from crop residues may increase in long run.