Evolutionary Ecology of Mountain Birch in Subarctic Stress Gradients: Interplay of Biotic and Abiotic Factors in Plant-Plant Interactions and Evolutionary Processes

In nature, variation for example in herbivory, wind exposure, moisture and pollution impact often creates variation in physiological stress and plant productivity. This variation is seldom clear-cut, but rather results in clines of decreasing growth and productivity towards the high-stress end. These clines of unidirectionally changing stress are generally known as ‘stress gradients’. Through its effect on plant performance, stress has the capacity to fundamentally alter the ecological relationships between individuals, and through variation in survival and reproduction it also causes evolutionary change, i.e. local adaptations to stress and eventually speciation. In certain conditions local adaptations to environmental stress have been documented in a matter of just a few generations. In plant-plant interactions, intensities of both negative interactions (competition) and positive ones (facilitation) are expected to vary along stress gradients. The stress-gradient hypothesis (SGH) suggests that net facilitation will be strongest in conditions of high biotic and abiotic stress, while a more recent ‘humpback’ model predicts strongest net facilitation at intermediate levels of stress. Plant interactions on stress gradients, however, are affected by a multitude of confounding factors, making studies of facilitation-related theories challenging. Among these factors are plant ontogeny, spatial scale, and local adaptation to stress. The last of these has very rarely been included in facilitation studies, despite the potential co-occurrence of local adaptations and changes in net facilitation in stress gradients. Current theory would predict both competitive effects and facilitative responses to be weakest in populations locally adapted to withstand high abiotic stress. This thesis is based on six experiments, conducted both in greenhouses and in the field in Russia, Norway and Finland, with mountain birch (Betula pubescens subsp. czerepanovii) as the model species. The aims were to study potential local adaptations in multiple stress gradients (both natural and anthropogenic), changes in plant-plant interactions under conditions of varying stress (as predicted by SGH), potential mechanisms behind intraspecific facilitation, and factors confounding plant-plant facilitation, such as spatiotemporal, ontogenetic, and genetic differences. I found rapid evolutionary adaptations (occurring within a time-span of 60 to 70 years) towards heavy-metal resistance around two copper-nickel smelters, a phenomenon that has resulted in a trade-off of decreased performance in pristine conditions. Heavy-metal-adapted individuals had lowered nickel uptake, indicating a possible mechanism behind the detected resistance. Seedlings adapted to heavy-metal toxicity were not co-resistant to others forms of abiotic stress, but showed co-resistance to biotic stress by being consumed to a lesser extent by insect herbivores. Conversely, populations from conditions of high natural stress (wind, drought etc.) showed no local adaptations, despite much longer evolutionary time scales. Due to decreasing emissions, I was unable to test SGH in the pollution gradients. In natural stress gradients, however, plant performance was in accordance with SGH, with the strongest host-seedling facilitation found at the high-stress sites in two different stress gradients. Factors confounding this pattern included (1) plant size / ontogenetic status, with seedling-seedling interactions being competition dominated and host-seedling interactions potentially switching towards competition with seedling growth, and (2) spatial distance, with competition dominating at very short planting distances, and facilitation being strongest at a distance of circa ¼ benefactor height. I found no evidence for changes in facilitation with respect to the evolutionary histories of plant populations. Despite the support for SGH, it may be that the ‘humpback’ model is more relevant when the main stressor is resource-related, while what I studied were the effects of ‘non-resource’ stressors (i.e. heavy-metal pollution and wind). The results have potential practical applications: the utilisation of locally adapted seedlings and plant facilitation may increase the success of future restoration efforts in industrial barrens as well as in other wind-exposed sites. The findings also have implications with regard to the effects of global change in subarctic environments: the documented potential by mountain birch for rapid evolutionary change, together with the general lack of evolutionary ‘dead ends’, due to not (over)specialising to current natural conditions, increase the chances of this crucial forest-forming tree persisting even under the anticipated climate change.

Voimalaitos- ja jätevesitutkimus

Tässä työssä tutkittiin Stora Enso Oyj:n Heinolan Flutingtehtaan voimalaitos- ja jätevesien seuranta-analyysimenetelmien kehittämistä. Käytössä olevia menetelmiä vertailtiin vaihtoehtoisiin uusiin menetelmiin, jotka perustuvat erotustekniikoihin ja automaatioon. Flutingtehtaalla nykyisin käytössä olevat analyysimenetelmät perustuvat suurelta osin standardimäärityksiin, joissa käytetään pääasiassa titrausta. Määritykset vievät paljon aikaa, koska titraukset toteutetaan manuaalisesti. Titrausten päätepisteet tulkitaan esim. indikaattorin värinmuutoksella ja saostamalla, joten määritysten tarkkuus vaihtelee. Kokeellisessa osassa Flutingtehtaan puhtaista voimalaitosvesistä yhdistetty sekoitenäyte analysoitiin kahdella ionikromatografilla, liekkiatomiabsorptiospektrometrillä ja kapillaarielektroforeesilla. Yksittäisiä näytteitä ei tutkittu. Lisäksi vesilaboratoriossa määritettävistä jätevesistä yhdistettiin sekoitenäyte, joka analysoitiin kapillaarielektroforeesilla. Samat sekoitenäytteet analysoitiin myös nykyisillä menetelmillä tehtaan vesilaboratoriossa. Tulokset osoittivat, että kokeellisessa osassa tutkitut menetelmät soveltuvat sekoitenäytteen perusteella hyvin vesilaboratoriossa käytössä oleviin kuukausianalyyseihin. Automaattisella näytteensyötöllä varustettuna kaikki kolme kokeellisessa osassa tutkittua menetelmää ovat yksikertaisia käyttää ja ne nopeuttavat analyysejä. Päivittäisiä titrausanalyysejä voidaan tehostaa ja nopeuttaa automaation avulla. Erotustekniikoilla esimerkiksi typpi voidaan määrittää kokonaistyppenä, mutta myös komponentteinaan eli ammoniakkina, nitraattina ja nitriittinä. Lisäksi samalla erotuksella voidaan määrittää useita alkali- ja maa-alkalimetalleja sekä raskas-metalleja toistomittauksilla. Menetelmän käyttöalue on laajempi erotustekniikoilla kuin perinteisillä liuoskemian mittauksilla. Erotustekniikoilla tunnistetaan helposti määritysten oikeellisuus detektointimahdollisuuksien monipuolisuuden vuoksi.

Stop bugging me! : diversity in appearance of warning coloration

In nature, many animals use body coloration to communicate with each other. For example, colorations can be used as signals between individuals of the same species, but also to recognise individuals of other species, and if they may comprise a threat or not. Many animals use protective coloration to avoid predation. The two most common strategies of protective coloration are camouflage and aposematism. Camouflaged animals have coloration that minimises detection, usually by matching colours or structures in the background. Aposematic animals, on the other hand, signal to predators that they are defended. The defence can be physical structures, such as spikes and hairs, or chemical compounds that make the animal distasteful or even deadly toxic. In order for the warning signal to be effective, the predator has to recognise it as such. Studies have shown that birds for example, that are important visual predators on insects, learn to recognise and avoid unpalatable prey faster if they contrast the background or have large internal contrasts. Typical examples of aposematic species have conspicuous colours like yellow, orange or red, often in combination with black. My thesis focuses on the appearance and function of aposematic colour patterns. Even though researchers have studied aposematism for over a century, there is still a lot we do not know about the phenomenon. For example, as it is crucial that the predators recognise a warning signal, aposematic colorations should assumingly evolve homogeneously and be selected for maximal conspicuousness. Instead, there is an extensive variation of colours and patterns among warning colorations, and it is not uncommon to find typical cryptic colours, such as green and brown in aposematic colour patterns. One hypothesis to this variation is that an aposematic coloration does not have to be maximally signalling in order to be effective, instead it is sufficient to have distinct features that can be easily distinguished from edible prey. To be maximally conspicuous is one way to achieve this, but not the only way. Another hypothesis is that aposematic prey that do not exhibit maximal conspicuousness can exploit both camouflage and aposematism in a distance-dependent fashion, by being signalling when seen close up but camouflaged at a distance. Many prey animals also make use of both strategies by shifting colour at different ecological conditions such as seasonal variations, fluctuations in food resources or between life stages. Yet another explanation for the variation may be that prey animals are usually exposed to several predator species that vary in visual perception and tolerance towards various toxins. The aim with this thesis is, by studying their functions, to understand why aposematic warning signals vary in appearance, specifically in the level of conspicuousness, and if warning coloration can be combined with camouflage. In paper I, I investigated if the colour pattern of the aposematic larva of the Apollo butterfly (Parnassius apollo) can switch function with viewing distance, and be signalling at close range but camouflaged at a distance, by comparing detection time between different colour variants and distances. The results show that the natural coloration has a dual distance-dependent function. Moreover, the study shows that an aposematic coloration does not have to be selected for maximal conspicuousness. A prey animal can optimise its coloration primarily by avoiding detection, but also by investing in a secondary defence, which presence can be signalled if detected. In paper II, I studied how easily detected the coloration of the firebug (Pyrrhocoris apterus), a typical aposematic species, is at different distances against different natural backgrounds, by comparing detection time between different colour variants. Here, I found no distance-dependent switch in function. Instead, the results show that the coloration of the firebug is selected for maximal conspicuousness. One explanation for this is that the firebug is more mobile than the butterfly larva in study I, and movement is often incompatible with efficient camouflage. In paper III, I investigated if a seasonal related colour change in the chemically defended striated shieldbug (Graphosoma lineatum) is an adaptation to optimise a protective coloration by shifting from camouflage to aposematism between two seasons. The results confirm the hypothesis that the coloration expressed in the late summer has a camouflage function, blending in with the background. Further, I investigated if the internal pattern as such increased the effectiveness of the camouflage. Again, the results are in accordance with the hypothesis, as the patterned coloration was more difficult to detect than colorations lacking an internal pattern. This study shows how an aposematic species can optimise its defence by shifting from camouflage to aposematism, but in a different fashion than studied in paper I. The aim with study IV was to study the selection on aposematic signals by identifying characteristics that are common for colorations of aposematic species, and that distinguish them from colorations of other species. I compared contrast, pattern element size and colour proportion between a group of defended species and a group of undefended species. In contrast to my prediction, the results show no significant differences between the two groups in any of the analyses. One explanation for the non-significant results could be that there are no universal characteristics common for aposematic species. Instead, the selection pressures acting on defended species vary, and therefore affect their appearance differently. Another explanation is that all defended species may not have been selected for a conspicuous aposematic warning coloration. Taken together, my thesis shows that having a conspicuous warning coloration is not the only way to be aposematic. Also, aposematism and camouflage is not two mutually exclusive opposites, as there are prey species that exploit both strategies. It is also important to understand that prey animals are exposed to various selection pressures and trade-offs that affect their appearance, and determines what an optimal coloration is for each species or environment. In conclusion, I hold that the variation among warning colorations is larger and coloration properties that have been considered as archetypically aposematic may not be as widespread and representative as previously assumed.

Spray coating technique asa surface treatment for woodcontaining paper grades

The objective of this work was to introduce the emerging non-contacting spray coating process and compare it to the existing coating techniques. Particular emphasis was given to the details of the spraying process of paper coating colour and the base paper requirements set by the new coating method. Spraying technology itself is nothing new, but the atomisation process of paper coating colour is quite unknown to the paper industry. The differences between the rheology of painting and coating colours make it very difficult to utilise the existing information from spray painting research. Based on the trials, some basic conclusion can be made:The results of this study suggest that the Brookfield viscosity of spray coating colour should be as low as possible, presently a 50 mPas level is regarded as an optimum. For the paper quality and coater runnability, the solids level should be as high as possible. However, the graininess of coated paper surface and the nozzle wear limits the maximum solids level to 60 % at the moment. Most likelydue to the low solids and low viscosity of the coating colour the low shear Brookfield viscosity correlates very well with the paper and spray fan qualities. High shear viscosity is also important, but yet less significant than the low shear viscosity. Droplet size should be minimized and besides keeping the brrokfield viscosity low that can be helped by using a surfactant or dispersing agent in the coating colour formula. Increasing the spraying pressure in the nozzle can also reduce the droplet size. The small droplet size also improves the coating coverage, since there is hardly any levelling taking place after the impact with the base paper. Because of the lack of shear forces after the application, the pigment particles do not orientate along the paper surface. Therefore the study indicates that based on the present know-how, no quality improvements can be obtained by the use of platy type of pigments. The other disadvantage of them is the rapid deterioration of the nozzle lifetime. Further research in both coating colour rheology and nozzle design may change this in the future, but so far only round shape pigments, like typically calcium carbonate is, can be used with spray coating. The low water retention characteristics of spray coating, enhanced by the low solids and low viscosity, challenge the base paper absorption properties.Filler level has to be low not to increase the number of small pores, which have a great influence on the absorption properties of the base paper. Hydrophobic sizing reduces this absorption and prevents binder migration efficiently. High surface roughness and especially poor formation of the base paper deteriorate thespray coated paper properties. However, pre-calendering of the base paper does not contribute anything to the finished paper quality, at least at the coating colour solids level below 60 %. When targeting a standard offset LWC grade, spraycoating produces similar quality to film coating, but yet blade coating being on a slightly better level. However, because of the savings in both investment and production costs, spray coating may have an excellent future ahead. The porousnature of the spray coated surface offers an optimum substrate for the coldset printing industry to utilise the potential of high quality papers in their business.

Rural Food System Change in Tanzania during the Post-Ujamaa Era. A Case Study from Western Bagamoyo District

Food systems in Sub-Saharan Africa have been rapidly transforming during the recent decades with diverse outcomes on human development and environment. This study explores the food system change in rural villages in eastern Tanzania where subsistence agriculture has traditionally been the main source of livelihood. The focus is on the salient changes in the spatial dimensions and structural composition of the food system in the context of economic liberalization that has taken place after the end of the socialist ujamaa era in the mid-1980s. In addition, the linkages of the changes are examined in relation to food security, socio-economic situation, livelihoods, and local environment. The approach of the study is geographical, but also involves various multi-disciplinary elements, particularly from development studies. The research methods included thematic and questionnaire interviews, participatory tools, and the analysis of land use/ cover data and official documents. Several earlier studies that were made in the area during the late 1970s and 1980s provided an important reference base. The study shows that subsistence farming has lost its dominant role in food provisioning due to the declining productivity of land, livestock losses, and the increasing shift of labour to non-farm sectors. Also rapid population growth has added to the pressure on land and other natural resources. Despite the increasing need for money for buying marketed foods and other necessities, the nutritional situation shows improvement and severe malnutrition has diminished. However, the long-term sustainability of this transformation raises concerns. Firstly, the food security situation continues to be fragile and prone to shocks such as adverse climatic conditions, crop failures and price hikes. Secondly, the commodification of the food system and livelihoods in general is linked to rapid environmental degradation in the area, particularly the loss of soil fertility and deforestation. The situation calls for efforts that take more determined and holistic approaches towards sustainable development of the rural food system with particular focus on the role and viability of small-scale farming.

Managing the change of organizational culture in cross-border mergers and acquisitions: The role of post-acquisition integration in organizational culture change

Over the years, cross-border mergers and acquisitions have become a popular strategic option for variety of firms. Companies often seek rapid growth through acquiring potentially valuable enterprises or attempting to enhance their organization’s profitability by merging with other firms. However, managing the change of organizational culture is a major managerial challenge as companies often confront difficulties when merging two previously autonomous organizational cultures into one, joint organizational culture. Therefore, the purpose of this study is to increase understanding related to the challenges and possibilities concerning the management of organizational culture change in cross-border mergers and acquisitions. The research question “How to manage the change of organizational culture in cross-border mergers and acquisitions?” is analysed in relation to the theories presented in this thesis regarding organizational culture, organizational change and acculturation as well as in relation with the collected empirical data. The research question is divided into three sub-questions according to the following: (1) “What is the role of organizational culture in organizations?”, (2) “How to manage organizational change in mergers and acquisitions?” and (3) “How to manage organizational culture change through acculturation?”. The thesis is conducted as a qualitative case study research including three personal interviews and one group interview. The interviews were conducted as a combination of semi-structured and unstructured interviews. Theories related to organizational culture, the management of change as well as acculturation are studied and further analysed in relation to empirical material collected by the researcher. Research findings indicate that that several factors can influence the success of managing the organizational culture change in cross-border mergers and acquisitions. Factors such as defining the preferred acculturation model prior the merger; managing the resistance of change; open communication; acknowledgement of local culture and cultural differences; involvement of personnel in change processes; as well as the formulation and implementation of comprehensive change plans proved to be important factors with relation to successful management of organizational culture change