Abstract The main objective of this thesis was to investigate tree - cover crop interactions and soil response in an intercropping system, in which leguminous and grass cover crops were used with birch (red-leaved Downy birch Betula pubescens L.). The investigation was conducted in two field experiments in a nursery and in a greenhouse experiment. In the latter the effect of the water supply in interspesific competition was also investigated. The cover crops in the field experiments were perennial clovers Trifolium pratense L., T. repens L. and T. hybridum L. and annual clovers T. incarnatum L., T. resupinatum L. and T. subterraneum L. and perennial Festuca rubra L. The height, stem diameter, leaf area and nutrient status of the birch were determined, as well as soil nutrient status and microbial characteristics. The cover crops in the greenhouse experiment were T. repens, T. subterraeum and F. rubra. The biomass, height, leaf area, leaf area index, specific leaf area and N concentration of the birch, the biomass and N concentration of the cover crops were measured, and soil N and microbial characteristics, as well. Bare ground was the control in all the experiments. The perennial clovers and Festuca strongly decreased the birch growth and nutrient status, but the annual clovers sown in midsummer in the field experiments provided about as good growth as bare ground. In the greenhouse experiment all cover crops were effective competitors with the birch. The soil NO3-N was, in general, the highest on bare ground and second highest in the annual clover plots. Though there were, in general, only minor differences in the soil nutrient concentrations between the treatments, there were significant differences in the tree growth and nutrient concentrations. The interspecific competition in this kind of intercropping system is mainly belowground. The growth reduction in the birch was mainly due to competition for nitrogen but water seems to play an important role in regulating the competitive interaction between the birch and cover crops. The competition for these resources seems to be most crucial at the beginning of the growing season. The microbial biomasses and soil respiration were greatest in the Festuca and perennial clover treatments, which may indicate that microorganisms together with these cover crops may seriously compete with birch for nutrients. Intercropping system is complex and comprises both negative and positive influences. In order to minimize negative competition effects, the cover crop should be non-competitive or the ground should be kept free of vegetation at the beginning of the growing season. By improving soil microbial characteristics, the vegetative ground covers make this cropping system one possiblity towards sustainability in the long-term.

Abstract Mycorrhizal fungi are important contributors to the functioning of boreal forests, since they act in the bilateral carbon and nutrient transport between above- and belowground parts of the ecosystem. In ectomycorrhizal (ECM) symbiosis of woody host plants, both fungal and plant partners depend on resources provided by the other. A single tree may simultaneously host several ECM fungal partners, which greatly enhance the host's nutrient uptake. At the same time nearly 20% of host primary production is allocated to mycorrhizal fungi. Although fungi depend on host-derived carbon, it is poorly understood how reduced carbon availability, e.g., due to herbivory, affects the ECM fungal symbionts. In this thesis I studied the impact of simulated insect defoliation or mammal browsing on mycorrhizal fungi of boreal woody hosts. Quantitative and qualitative changes in biomass partitioning in different fungal compartments were detected. None of the experiments showed that defoliation or shoot clipping treatments reduced the intensity of ECM colonisation, while treatments often shifted fungal composition towards less biomass producing ECM morphotypes. Above- and belowground diversity in ECM symbionts tended to decrease due to shoot or foliar damage. In addition, in some cases defoliation also reduced fungal biomass in fine roots and decreased ECM sexual reproduction by reducing the number of sporocarps produced. Defoliation induced a similar response pattern in the host and in ECM fungi with a stronger response to increasing severity of treatment (e.g. degree of removed foliage or repeated years of defoliation). This was also confirmed when relating the effects of host and ECM fungal symbionts to defoliation using present and previously published data. The present results suggest that belowground adaptation of boreal trees to the changing environment is mediated by changes in fungal community or biomass partitioning. The lack of response in the intensity of ECM colonisation further emphasises the importance of the symbiosis to boreal trees.

Thesis (doctoral)--Universiteit van Amsterdam, 1985. / Five charts in back cover pocket. Text in English, with foreword and summary in Dutch. Includes bibliographical references (p. 147-152).

Abstract Mechanical harvesting of peat changes the original mire ecosystem completely, and without active measures these areas may remain non-vegetated even for decades. Afforestation is one of the most popular after-use options for cut-away peatlands in Finland since it has both economic and aesthetic values. Recycling of wood-ash as a fertilizer has been studied extensively in peatlands drained for forestry. Wood-ash is reported to promote tree growth in these areas without any significant negative impact to the environment and could, therefore, be a suitable option also on cut-away peatlands. However, the environmental effects of ash-fertilization on cut-away areas and on ground vegetation are not fully understood. The impact of wood- and peat-ash application on the early establishment of ground vegetation and downy birch (Betula pubescens) seedlings and on post-fertilization element concentrations in plants and peat substrate were studied in a cut-away peatland. Six treatments of wood-ash, peat-ash, biotite or Forest PK-fertilizer were replicated in three blocks in different mixtures and quantities corresponding to 50 kg ha-1 of phosphorus. All the fertilizers accelerated the revegetation of the bare peat surface significantly, whereas the establishment of plants in the unfertilized area was non-existent even several years after the peat harvesting had ceased. The most striking difference between the wood- and peat-ash-fertilizers and the commercial Forest PK-fertilizer was the extensive coverage of fire-loving moss species in all the areas where ash was spread. Wood- and peat-ash application also accelerated the germination and early establishment of downy birch seedlings more efficiently than the PK-fertilizer. Ground vegetation proved to be highly important in increasing the early biomass production and carbon sequestration on ash-fertilized cut-away peatland. In addition, the below-ground biomass was equal to the above-ground biomass, or even greater. Both wood- and peat-ash fertilization ensured an adequate level of nutrients for the early establishment of ground vegetation and birch seedlings in a cut-away peatland. The mosses and herbaceous plants proved to have a major role in retaining the nutrients and heavy metals that otherwise might have leached away from the ash-fertilized cut-away site during the early stages of the afforestation. Although both wood- and peat-ash proved to be suitable for the initial fertilization of afforested cut-away peatlands, a later application of nutrients may be needed to guarantee the growth in a energy-wood stand of downy birch over its entire rotation. / Tiivistelmä Turvetuotannon päätyttyä jäljelle jäävä suopohja on aluksi täysin paljas ja vailla maaperän siemenpankkia. Kasvipeitteen luontainen uudistuminen voi viedä jopa vuosikymmeniä. Ympäristönhoidollisesti onkin tärkeää, että suopohjat otetaan uuteen käyttöön mahdollisimman nopeasti tuotannon päätyttyä. Metsitys on tällä hetkellä suosituin suopohjien jälkikäyttömuoto Suomessa. Suopohjan turpeessa on tyypillisesti runsaasti typpeä, mutta niukasti muita kasvuun tarvittavia ravinteita. Puutuhka on osoittautunut pitkäaikaisissa metsäojitettujen turvemaiden tutkimuksissa kilpailukykyiseksi vaihtoehdoksi kaupallisille lannoitteille. Energiantuotannon sivutuotteena syntyvä puu- ja turvetuhka voisi soveltua hyvin myös suopohjien lannoitteeksi. Tuhkan käytöstä lannoitteena turvetuotannosta vapautuneilla suopohjilla ei kuitenkaan ole riittävästi tutkimustietoa. Tässä työssä tutkittiin puu- ja turvetuhkan vaikutuksia turvetuotannosta vapautuneen suopohjan kasvittumiseen ja puun taimien alkukehitykseen viiden ensimmäisen kasvukauden ajan. Lisäksi tutkittiin kasvillisuuden ravinne- ja raskasmetallipitoisuuksien muutoksia sekä turpeen ravinteisuutta lannoituksen jälkeen. Kaikki lannoitteet nopeuttivat merkittävästi kasvillisuuden muodostumista paljaalle suopohjalle, kun taas lannoittamaton alue pysyi kasvittomana. Tuhkalannoitetuille alueille syntyi nopeasti laajoja palopaikoilla viihtyvien pioneerisammalten kasvustoja, jotka peittivät ja samalla sitoivat paljaan ja irtonaisen turvemaan pinnan. Tuhkalannoitus edisti myös koivun taimien alkukehitystä tehokkaammin kuin kaupallinen Metsän PK-lannoite. Sammalista ja ruohovartisista kasveista muodostuva aluskasvillisuus ylitti puuntaimet selvästi biomassan määrässä ja toimi metsityksen alkuvaiheessa puuntaimia merkittävämpänä hiilensitojana. Lisäksi kasvien maanalainen biomassa oli maanpäällistä biomassaa suurempi. Sekä puu- että turvetuhka takasivat riittävän määrän ravinteita energiapuumetsikön alkukehitykselle. Aluskasvillisuus osoittautui tärkeäksi tuhkasta liukenevien ravinteiden ja raskasmetallien sitojaksi metsityksen alkuvaiheessa. Vaikka sammalten kadmiumpitoisuudet nousivat tuhkalannoituksen seurauksena, ne olivat kuitenkin alhaisia Suomessa aiemmin mitattuihin sammalten yleisiin pitoisuuksiin suhteutettuna. Tuhkalannoitus ei lisännyt haitallisten raskasmetallien pitoisuuksia koivun taimien ja ruohovartisten kasvien lehdissä ja varsissa. Tutkimuksen tulokset tukevat puu- ja turvetuhkan käyttöä energiapuumetsiköiden alkuvaiheen lannoitteena turvetuotannosta vapautuneilla suopohjilla.

Boreal forests are particularly vulnerable to climate change, experiencing a much more drastic increase in temperatures and having a limited amount of more northern refugia. The trees making up these vast and important ecosystems already had to adapt previously to environmental pressures brought about by the repeated glaciations during past ice ages. Studying the patterns of adaption of these trees can thus provide valuable insights on how to mitigate future damage. This thesis presents and analyses population structure, demo- graphic history and the distribution of fitness effects (DFE) of the diploid Betula pendula and tetraploid B. pubescens across Scandinavia. Birches–being widespread in boreal forests as well as having great economical importance–constitute superb model species. The analyses of this work confirm the expectations on postglacial population expansion and diploid-tetraploid introgression. They furthermore ascertain the presence of two genetic clusters and a remarkably similar DFE for the species. This work also contributes with a transparent, reproducible and reusable pipeline which facilitates running similar analyses for related species.