Growth of and partitioning between shoot and storage root of carrot in a northern climate

Selostus: Porkkanan kasvu ja biomassan jakautuminen varastojuuren ja verson välillä pohjoisissa oloissa

Biodiversity and ecosystem functioning in angiosperm communities in the Baltic Sea

The biological variation in nature is called biodiversity. Anthropogenic pressures have led to a loss of biodiversity, alarming scientists as to what consequences declining diversity has for ecosystem functioning. The general consensus is that diversity (e.g. species richness or identity) affects functioning and provides services from which humans benefit. The aim of this thesis was to investigate how aquatic plant species richness and identity affect ecosystem functioning in terms of processes such as primary production, nutrient availability, epifaunal colonization and properties e.g. stability of Zostera marina subjected to shading. The main work was carried out in the field and ranged temporally from weeklong to 3.5 months-long experiments. The experimental plants used frequently co-occur in submerged meadows in the northern Baltic Sea and consist of eelgrass (Z. marina), perfoliate pondweed (Potamogeton perfoliatus), sago pondweed (P. pectinatus), slender-leaved pondweed (P. filiformis) and horned pondweed (Zannichellia palustris). The results showed that plant richness affected epifaunal community variables weakly, but had a strong positive effect on infaunal species number and functional diversity, while plant identity had strong effects on amphipods (Gammarus spp.), of which abundances were higher in plant assemblages consisting of P. perfoliatus. Depending on the starting standardizing unit, plant richness showed varying effects on primary production. In shoot density-standardized plots, plant richness increased the shoot densities of three out of four species and enhanced the plant biomass production. Both positive complementarity and selection effects were found to underpin the positive biodiversity effects. In shoot biomass-standardized plots, richness effects only affected biomass production of one species. Negative selection was prevalent, counteracting positive complementarity, which resulted in no significant biodiversity effect. The stability of Z. marina was affected by plant richness in such that Z. marina growing in polycultures lost proportionally less biomass than Z. marina in monocultures and thus had a higher resistance to shading. Monoculture plants in turn gained biomass faster, and thereby had a faster recovery than Z. marina growing in polycultures. These results indicate that positive interspecific interactions occurred during shading, while the faster recovery of monocultures suggests that the change from shading stress to recovery resulted in a shift from positive interactions to resource competition between species. The results derived from this thesis show that plant diversity affects ecosystem functioning and contribute to the growing knowledge of plant diversity being an important component of aquatic ecosystems. Diverse plant communities sustain higher primary productivity than comparable monocultures, affect faunal communities positively and enhance stability. Richness and identity effects vary, and identity has generally stronger effects on more variables than richness. However, species-rich communities are likely to contain several species with differing effects on functions, which renders species richness important for functioning. Mixed meadows add to coastal ecosystem functioning in the northern Baltic Sea and may provide with services essential for human well-being.

„Und frei erklär' ich alle meine Knechte“ : Höflichkeit als Ausdruck sozialer Utopie in Schillers Drama Wilhelm Tell.

My dissertation is an interdisciplinary study of the relationship between the Swiss peasants and nobles in Friedrich Schiller’s play Wilhelm Tell (1804). Changes to this relationship are of particular interest. Communication in the play is examined via a micro-analysis based on Penelope Brown’s and Stephen C. Levinson’s theoretical framework Politeness: Some universals in language usage. (1978, 1987). Brown and Levinson distinguish between positivepoliteness and negative-politeness strategies and their distinction is useful for my argument, since they claim that the use of positive-politeness strategies reflects the speaker’s intention to minimize the distance between the interlocutors (Brown and Levinson 1987: 103). Negative-politeness strategies, by contrast, result in social distancing (Brown and Levinson 1987: 130). In accordance with Brown’s and Levinson’s theory, it can be argued that the distribution of positive-politeness and negative-politeness strategies reflects changes in the distance between the fictional interlocutors as representatives of their social classes in Wilhelm Tell. The analysis of the communication in the play highlights that existing conflicts within the social groups are resolved and replaced with solidarity (cf. ‘Claim ’common ground’ and ‘Claim in-group membership with H’) before the peasants and noblemen appear on stage simultaneously in III, 3. In the scene in question, Geßler forces Tell to shoot the apple off his son’s head. Although both nobles and peasants are present, they do not communicate with each other. Thus, communication between the social classes occurs for the first time in IV, 2. This scene is crucial with regard to the changes in the relationship between the social classes in the play. The younger generation, with Rudenz as a representative of the nobility and Melchthal as a representative of the peasants, break with the prevailing conventions and initiate a new type of cooperation based on mutual helping and equality in the right and the duty to protect the country from the violent oppressors representing Habsburg (cf. ‘Convey that S and H are cooperators’, ‘Claim reflexivity’ and ‘Claim reciprocity’). The linguistic analysis reveals the crucial role of the Swiss nobility in the development of the social utopia, as well as the non-contribution of Wilhelm Tell. In fact, Tell never communicates with the Swiss noblemen. The role of the nobility and the role of Wilhelm Tell are further investigated in a comparison between Schiller’s Wilhelm Tell and four of Schiller’s historical sources. These sources, which contain earlier, non-dramatic versions of the establishment of the Swiss Confederation, are the following: Kronica von der loblichen Eydtgnoschaft compiled by Petermann Etterlin (1507), Gemeiner loblicher Eydgnoschafft Stetten Landen vnd Völckeren Chronik wirdiger thaaten beschreybung (1548) written by Johann Stumpf, Chronicon Helveticum of Aegidius Tschudi (the publication of Iselin from 1734 and 1736) and Johannes von Müller’s Geschichten schweizerischer Eidgenossenschaft (1786). Thecomparison sheds light on the fact that both Tschudi and von Müller emphasize unity among the Swiss and cooperation in their attempt to defeat the enemy. However, Schiller has reinforced the role of the nobility in the cooperation. In addition, the comparison between Schiller’s play and the historical sources reveals profound differences with regard to the role of Wilhelm Tell. In all of Schiller’s sources, Tell is present at Rütli, whereas he is absent from Rütli in Schiller’s play. In the play in general, Tell is conspicuously separated from the other peasants. Explanations of my linguistic results, which correspond to the above mentioned differences between the roles of the figures in Wilhelm Tell and the depictions in Schiller’s sources, are found by comparing the path of Schiller’s Swiss towards an egalitarian perspective with the kind of social evolution depicted by the German philosopher Johann Benjamin Erhard in his essay Über das Recht des Volks zu einer Revolution (1795). Aiming at considering Schiller’s social utopia within the larger cultural framework of the German response to the French Revolution, the comparison suggests that both authors depict the need for social change in terms of a change in social hierarchies (Erhard 1970: 95-96, cf. Foi 2005: 225). Erhard’s essay thus helps explain the political intention of Schiller’s play to keep the ideals of the French Revolution as crucial aims but profoundly change the means towards freedom and equality. In his attempt to claim the political resistance of the nobles and peasants as just, however, Schiller sacrificed the figure of Wilhelm Tell. Guilty of the murder of Geßler, Tell was no longer suitable for the righteous revolution imagined by Schiller (cf. Bloch 2008: 215 and Schulz 2005: 228). This explains Tell’s absence in the Rütli scene, his isolation in the plot, as well as his non-contribution to the social utopia. Together, the linguistic analysis and contextualisation of Schiller’s play support my hypothesis that Wilhelm Tell describes a process of change in the relationship between peasants and nobles. The interdisciplinary approach to the topic proved to be fruitful for all areas of the research involved.

Effects of algal turbidity on foraging and antipredator behaviour of the three-spined stickleback (Gasterosteus aculeatus)

The aim of this thesis was to examine how aquatic organisms, such as fish, behave in an altered environmental condition. Many species of fish use vision as their primary tool to gain information about their surrounding environment. The visual conditions of aquatic habitats are often altered as a result of anthropogenic disturbance, such as eutrophication that initiates algal turbidity. In general, turbidity reduces the visibility and can be hypothesized to have an influence on the behaviour of fish. I used the three-spined stickleback (Gasterosteus aculeatus) as a model species and conducted four studies in the laboratory to test how algal turbidity affects its behaviour. In this thesis, two major behavioural aspects are discussed. The first is antipredator behaviour. In study I, the combined effects of turbidity and shoot density on habitat choice (shelter vs open) behaviour was tested on a group of sticklebacks (20 fish) in the presence and absence of piscivorous perch (Perca fluviatilis). In study II, I examined the behavioural responses of feeding sticklebacks when they were exposed to the sudden appearance of an avian predator (the silhouette of a common tern, Sterna hirundo). The study was done in turbid and clear water using three different groups sizes (1, 3 and 6 fish). The second aspect is foraging behaviour. Study III & IV focused on the effects of algal turbidity on the foraging performance of sticklebacks. In study III, I conducted two separate experiments to examine the effects of turbidity on prey consumption and prey choice of sticklebacks. In this experiment turbidity levels and the proportion of large and small prey (Daphnia spp.) were manipulated. In study IV, I studied whether a group of six sticklebacks can distribute themselves according to food input at two feeding stations in a way that provided each fish with the same amount of food in clear and turbid water. I also observed whether the fish can follow changes in resource distribution between the foraging patches. My results indicate an overall influence of algal turbidity on the antipredator and foraging behaviour of sticklebacks. In the presence of a potential predator, the use of the sheltered habitat was more pronounced at higher turbidity. Besides this, sticklebacks reduced their activity levels with predator presence at higher turbidity and shoot density levels, suggesting a possible antipredator adaptation to avoid a predator. When exposed to a sudden appearance of an avian predator, sticklebacks showed a weaker antipredator response in turbid water, which suggests that turbidity degrades the risk assessment capabilities of sticklebacks. I found an effect of group size but not turbidity in the proportion of sticklebacks that fled to the shelter area, which indicates that sticklebacks are able to communicate among group members at the experimental turbidity levels. I found an overall negative effect of turbidity on food intake. Both turbidity and changes in the proportion of prey sizes played a significant role in a stickleback’s prey selection. At lower turbidity levels (clear <1 and 5 NTU) sticklebacks showed preferences for large prey, whereas in more turbid conditions and when the proportion of large to small prey increased sticklebacks became increasingly random in their prey selection. Finally, my results showed that groups of sticklebacks disperse themselves between feeding stations according to the reward ratios following the predictions of the ideal free distribution theory. However, they took a significantly longer time to reach the equilibrium distribution in turbid water than in clear water. In addition, they showed a slower response to changes in resource distribution in a turbid environment. These findings suggest that turbidity interferes with the information transfer among group foragers. It is important to understand that aquatic animals are often exposed to a degraded environment. The findings of this thesis suggest that algal turbidity negatively affects their behavioural performance. The results also shed light on the underlying behavioural strategies of sticklebacks in turbid conditions that might help them adapt to an altered environmental situation and increase their survival. In conclusion, I hold that although algal turbidity has detrimental effects on the antipredator and foraging behaviour of sticklebacks, their behavioural adjustment might help them adapt to a changing environment.