Plant-Herbivore Interaction in a Fragmented Landscape: Local Adaptation and Inbreeding

Reciprocal selection between interacting species is a major driver of biodiversity at both the genetic and the species level. This reciprocal selection, or coevolution, has led to the diversification of two highly diverse and abundant groups of organisms, flowering plants and their insect herbivores. In heterogeneous environments, the outcome of coevolved species interactions is influenced by the surrounding community and/or the abiotic environment. The process of adaptation allows species to adapt to their local conditions and to local populations of interacting species. However, adaptation can be disrupted or slowed down by an absence of genetic variation or by increased inbreeding, together with the following inbreeding depression, both of which are common in small and isolated populations that occur in fragmented environments. I studied the interaction between a long-lived plant Vincetoxicum hirundinaria and its specialist herbivore Abrostola asclepiadis in the southwestern archipelago of Finland. I focused on mutual local adaptation of plants and herbivores, which is a demonstration of reciprocal selection between species, a prerequisite for coevolution. I then proceeded to investigate the processes that could potentially hamper local adaptation, or species interaction in general, when the population size is small. I did this by examining how inbreeding of both plants and herbivores affects traits that are important for interaction, as well as among-population variation in the effects of inbreeding. In addition to bi-parental inbreeding, in plants inbreeding can arise from self-fertilization which has important implications for mating system evolution. I found that local adaptation of the plant to its herbivores varied among populations. Local adaptation of the herbivore varied among populations and years, being weaker in populations that were most connected. Inbreeding caused inbreeding depression in both plants and herbivores. In some populations inbreeding depression in herbivore biomass was stronger in herbivores feeding on inbred plants than in those feeding on outbred ones. For plants it was the other way around: inbreeding depression in anti-herbivore resistance decreased when the herbivores were inbred. Underlying some of the among-population variation in the effects of inbreeding is variation in plant phenolic compounds. However, variation in the modification of phenolic compounds in the digestive tract of the herbivore did not explain the inbreeding depression in herbivore biomass. Finally, adult herbivores had a preference for outbred host plants for egg deposition, and herbivore inbreeding had a positive effect on egg survival when the eggs were exposed to predators and parasitoids. These results suggest that plants and herbivores indeed exert reciprocal selection, as demonstrated by the significant local adaptation of V. hirundinaria and A. asclepiadis to one another. The most significant cause of disruption of the local adaptation of herbivore populations was population connectivity, and thus probably gene flow. In plants local adaptation tended to increase with increasing genetic variation. Whether or not inbreeding depression occurred varied according to the life-history stage of the herbivore and/or the plant trait in question. In addition, the effects of inbreeding strongly depended on the population. Taken together, inbreeding modified plant-herbivore interactions at several different levels, and can thus affect the strength of reciprocal selection between species. Thus inbreeding has the potential to affect the outcome of coevolution.

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.

Implementing an Evidence-based Hypertension Guideline into Finnish Primary Care Nursing

Kohonneen verenpaineen hoitosuosituksen käyttöönottosuomen perusterveydenhiollon hoitotyössä Tutkimuksen tavoitteena oli tuottaa suosituksia näyttöön perustuvien Käypä hoito -suositusten käytön edistämiseksi perusterveydenhuollon hoitotyössä. Tutkimuksen ensimmäisessä vaiheessa arvioitiin Kohonneen verenpaineen hoitosuosituksen käyttöönottoa terveyskeskuksissa. Toisessa vaiheessa selvitettiin hoitajien hoitosuositusasenteita ja kokemuksia hoitosuosituksen käyttöönotosta. Kolmannessa vaiheessa selvitettiin hoitohenkilöstön näkemyksiä hoitosuosituksen käyttöä edistävistä tekijöistä. Kohonneen verenpaineen hoitosuositus oli ylilääkäreiden ja ylihoitajien mukaan otettu käyttöön lähes kaikissa terveyskeskuksissa, mutta heidän näkemyksensä suositusten käyttöönottoa koskevista terveyskeskuksissa tehdyistä sopimuksista erosivat toisistaan monilta osin. Myös käyttöönoton toteutuksessa oli suurta vaihtelua terveyskeskusten välillä. Toteutustavan perusteella ääripäissä sijaitsevat terveyskeskukset luokiteltiin yksittäisin ja monin keinoin käyttöönottoa tukeneiksi. Hoitajien hoitosuositusasenteet olivat hyvin myönteisiä ja hoitosuosituksia pidettiin luotettavina tiedonlähteinä, ja niiden uskottiin parantavan hoidon laatua. Hoitosuositusten paikallinen soveltaminen sekä johdon ja lääkäreiden tuki olivat hoitajien mielestä keskeisiä käyttöönotossa, vaikkakin tulosten mukaan kaikki käytetyt keinot olivat yhteydessä positiivisempiin hoitosuositusasenteisiin sekä aktiivisempaan hoitajien itsensä ilmaisemaan hoitosuositusten käyttöön. Yhteenvetona voidaan todeta, että Käypä Hoito -suositukset on hyväksytty osaksi kliinistä hoitotyön käytäntöä. Niiden käytön tehostamiseksi tulisi kiinnittää huomiota suositusten paikalliseen soveltamiseen ja eri ammattiryhmien tehtäväkuvien määrittelyyn. Tähän tarvitaan terveyskeskusten johdon ja lääkäreiden selkeää tukea.

From genes to communities : stress tolerance in eelgrass (Zostera marina)

Shallow coastal areas are dynamic habitats that are affected by a variety of abiotic and biotic factors. In addition to the natural environmental stress, estuarine and coastal seagrass ecosystems are exposed to effects of climate change and other anthropogenic impacts. In this thesis the effect of different abiotic (shading stress, salinity and temperature) and biotic stressors (presence of co-occurring species) and different levels and combinations of stressors on the performance and survival of eelgrass (Zostera marina) was assessed. To investigate the importance of scale for stress responses, varying levels of biological organization (genotype, life stage, population and plant community) were studied in field and aquarium experiments. Light limitation, decreased salinity and increased temperature affected eelgrass performance negatively in papers I, II and III, respectively. While co-occurring plant species had no notable effect on eelgrass in paper IV, the presence of eelgrass increased the biomass of Potamogeton perfoliatus. The findings in papers II and III confirmed that more extreme levels of salinity and temperature had stronger impacts on plant performance compared to intermediate levels, but intermediate levels also had more severe effects on plants when they were exposed to several stressors, as illustrated in paper II. Thus, multiple stressors had negative synergetic effects. The results in papers I, II and III indicate that future changes in light climate, salinity and temperature can have serious impacts on eelgrass performance and survival. Stress responses were found to vary among genotypes, life stages and populations in papers I, II and III, respectively, emphasizing the importance of study scale. The results demonstrate that while stress in general affects seagrass productivity negatively, the severity of effects can vary substantially depending on the studied scale or level of biological organization. Eelgrass genotypes can differ in their stress and recovery processes, as observed in paper I. In paper II, eelgrass seedlings were less prone to abiotic stress compared to adult plants, but stress also decreased their survival considerably. This indicates that recruitment and re-colonization through seeds might be threatened in the future. Variation among population responses observed in paper III indicates that long-term local adaptation under differing selection pressures has caused divergence in salinity tolerance between Baltic eelgrass populations. This variability in stress tolerance observed in papers I and III suggests that some eelgrass genotypes and populations have a better capacity to adapt to changes and survive in a changing environment. Multiple stressors and biological level-specific responses demonstrate the uncertainty in predicting eelgrass responses in a changing environment. As eelgrass populations may differ in their stress tolerance both within and across regions, conservation strategies at both local and regional scales are urgently needed in order to ensure the survival of these important ecosystems.

Adaptive Dynamics of Resource Specialization

Ecological specialization in resource utilization has various facades ranging from nutritional resources via host use of parasites or phytophagous insects to local adaptation in different habitats. Therefore, the evolution of specialization affects the evolution of most other traits, which makes it one of the core issues in the theory of evolution. Hence, the evolution of specialization has gained enormous amounts of research interest, starting already from Darwin’s Origin of species in 1859. Vast majority of the theoretical studies has, however, focused on the mathematically most simple case with well-mixed populations and equilibrium dynamics. This thesis explores the possibilities to extend the evolutionary analysis of resource usage to spatially heterogeneous metapopulation models and to models with non-equilibrium dynamics. These extensions are enabled by the recent advances in the field of adaptive dynamics, which allows for a mechanistic derivation of the invasion-fitness function based on the ecological dynamics. In the evolutionary analyses, special focus is set to the case with two substitutable renewable resources. In this case, the most striking questions are, whether a generalist species is able to coexist with the two specialist species, and can such trimorphic coexistence be attained through natural selection starting from a monomorphic population. This is shown possible both due to spatial heterogeneity and due to non-equilibrium dynamics. In addition, it is shown that chaotic dynamics may sometimes inflict evolutionary suicide or cyclic evolutionary dynamics. Moreover, the relations between various ecological parameters and evolutionary dynamics are investigated. Especially, the relation between specialization and dispersal propensity turns out to be counter-intuitively non-monotonous. This observation served as inspiration to the analysis of joint evolution of dispersal and specialization, which may provide the most natural explanation to the observed coexistence of specialist and generalist species.

Grass-endophyte coevolution and ploidy levels in fescues

ABSTRACT Fescues consist of wild and cultivated grasses that have adapted to a wide range of environmental conditions. They are an excellent model species for evolutionary ecology studies that investigate symbiosis and polyploidization and their effects on plant performance. First, they are frequently infected with symbiotic endophytic fungi known to affect a plant’s ability to cope with biotic and abiotic environmental factors. Second, fescue species have been reported to have substantial intraspecific variation in their ploidy level and morphology. In my thesis, I examined large-scale generalizations for frequency of polyploidy and endophyte infections and their effects on plant morphology. As a model species, I selected red (Festuca rubra) and viviparous sheep’s (F. vivipara) fescues. They are closely related, but they differ in terms of distribution and endophyte infection frequency. I investigated the biogeographic pattern and population biology of 29 red and 12 viviparous sheep’s fescue populations across ≈300 latitudes in Europe (400-690 N). To examine plant ploidy levels, I implemented time- and cost-efficient plate-based high throughput flow cytometric analysis. This efficient procedure enabled me to analyze over 1000 red fescue individuals. I found three ploidy levels among them: overall 84 %, 9 % and 7 % of the red fescue plants were hexaploid, tetraploid and octoploid, respectively. However, all viviparous sheep’s fescue plants were tetraploid. Ploidy level of red fescue appeared to some extent follow gradients in latitude and primary production as suggested by previous studies, but these results could be explained better by taking the sampling design and local adaptation into account. Three Spanish populations were mostly tetraploids and one high elevation population in northernmost Finland (Halti) was octoploid, while most other populations (25 sites) were dominated by hexaploids. Endophyte infection frequencies of wild fescue populations varied from 0 to 81 % in red fescue populations and from 0 to 30 % in viviparous sheep’s fescue populations. No gradients with latitude or primary production of the sites were detected. As taxonomy of red fescues is somewhat unclear, I also studied morphology, ploidy variation and endophyte status of proposed subspecies of European red fescues. Contrary to previous literature, different ploidy levels occurred in the same subspecies. In addition to wild fescues, I also used two agronomically important cultivars of meadow and tall fescue (Schedonorus phoenix and S. pratensis). As grass-legume mixtures have an agronomic advantage over monocultures in meadows, I carried out a mixture/competition experiment with fescues and red clover to find that species composition, nutrient availability and endophyte status together determined the total biomass yield that was higher in mixtures compared to monocultures. The results of this thesis demonstrate the importance of local biotic and abiotic factors such as grazing gradients and habitat types, rather than suggested general global geographical or environmental factors on grass polyploidization or its association with symbiotic endophytic fungi. I conclude that variation in endophyte infection frequencies and ploidy levels of wild fescues support the geographic mosaic theory of coevolution. Historical incidents, e.g., glaciation and present local factors, rather than ploidy or endophyte status, determine fescue morphology.

Adaptability, Transformation and Complex Changes in Namibia and Tanzania: Resilience and Innovation System Development in Local Communities

This thesis is a research about the recent complex spatial changes in Namibia and Tanzania and local communities’ capacity to cope with, adapt to and transform the unpredictability engaged to these processes. I scrutinise the concept of resilience and its potential application to explaining the development of local communities in Southern Africa when facing various social, economic and environmental changes. My research is based on three distinct but overlapping research questions: what are the main spatial changes and their impact on the study areas in Namibia and Tanzania? What are the adaptation, transformation and resilience processes of the studied local communities in Namibia and Tanzania? How are innovation systems developed, and what is their impact on the resilience of the studied local communities in Namibia and Tanzania? I use four ethnographic case studies concerning environmental change, global tourism and innovation system development in Namibia and Tanzania, as well as mixed-methodological approaches, to study these issues. The results of my empirical investigation demonstrate that the spatial changes in the localities within Namibia and Tanzania are unique, loose assemblages, a result of the complex, multisided, relational and evolutional development of human and non-human elements that do not necessarily have linear causalities. Several changes co-exist and are interconnected though uncertain and unstructured and, together with the multiple stressors related to poverty, have made communities more vulnerable to different changes. The communities’ adaptation and transformation measures have been mostly reactive, based on contingency and post hoc learning. Despite various anticipation techniques, coping measures, adaptive learning and self-organisation processes occurring in the localities, the local communities are constrained by their uneven power relationships within the larger assemblages. Thus, communities’ own opportunities to increase their resilience are limited without changing the relations in these multiform entities. Therefore, larger cooperation models are needed, like an innovation system, based on the interactions of different actors to foster cooperation, which require collaboration among and input from a diverse set of stakeholders to combine different sources of knowledge, innovation and learning. Accordingly, both Namibia and Tanzania are developing an innovation system as their key policy to foster transformation towards knowledge-based societies. Finally, the development of an innovation system needs novel bottom-up approaches to increase the resilience of local communities and embed it into local communities. Therefore, innovation policies in Namibia have emphasised the role of indigenous knowledge, and Tanzania has established the Living Lab network.

Kaupungin ja maaseudun vuorovaikutus alueellisessa kehittämispolitiikassa

In Finland, European Union membership and economic globalisation have changed the position of regions from closed territorial systems to nodes of open international networks. The increasing complexity of cities as globalised knowledge centres and functionally specialised and diversified rural areas, and on the other hand growing disparities between prosperous urban cores and lagging peripheral areas are also essential features in contemporary regional development. These trends have produced new needs to promote mutual dialogue between cities and the countryside in western market economies. Urban-rural interaction is an idea which was developed in the late 1990s within regional policy to pull together these new challenges to regional development and handle cities and the countryside as a whole. The aim of my study is to conceptualise the idea of urban-rural interaction, explain the phenomenon theoretically, clarify past and present urban and rural development and analyse regional policies from the interaction angle. The ultimate purpose is to illustrate the existence and nature of particular interaction policy in a globalising society. The general method is discourse analysis, which I use in three cases: Central Finland, South-Ostrobothnia and South-West Finland. Theoretically I have a two-dimensional approach. On the first hand I use World-System theory to explain how the global economy is moulding urban and rural structures at the regional level. On the other hand I use regime theory to explain local political actions and practises between cities and the countryside under the overlapping pressures deriving from reformulated regional structures and policies.

Adaptation to globalisation in Finland has been carried out by strengthening urban centres. The stress in regional policy has been in urban development. The development of the countryside has mostly been implemented by a separate rural policy. At the end of the 1990s and early 2000s Finnish cities have actually shown themselves to be competitive in global markets. The drawbacks of the new growth centre policy have been the sparse network of prosperous cities and their weak spreading effects, which have hindered comprehensive regional development. Tensions between urban and rural areas have also deepened. In this situation the interaction policy is used as a way of balancing development and moderate conflicts within the regions. From this point of view urban-rural interaction can be seen as a way of tackling the challenges of globalisation.

On the other hand the results emphasise that actors involved in regional development still believe, although the hegemonic discourse is on urban policy, that there are opportunities to stimulate progress in the countryside as well. In the situation where regional authorities control development resources, rural development can be successful only if rural actors manage to establish fruitful relationships with their urban partners. This is also the weakness of the programme-based regional policy. If rural municipalities or other actors are for any reason incapable of building development regimes with cities, the offers of interaction policy will be useless.

The problem of the interaction policy is that the focus and methods of it have so far been rather underdeveloped. In order to improve the efficiency of the interaction policy, further research should concentrate on the social processes which define the position of cities and the countryside as partners of interaction, and practises which promote or prohibit the possibilities of developing the interaction policy. The efforts to define different contents of urban-rural interaction or promote interaction projects should not have such an important role in the future as they have had so far. Instead, the focus of interaction policy should be on questions such as how to manage the political tensions between town and country and how to create a positive atmosphere for regional policy where the needs of urban and rural development are promoted equally.