The encroachment of woody species into grassland and savanna ecosystems has been well document since the early 1800s. Within the parkland ecoregion of western Canada, trembling aspen (Populus tremuloides Michx.) has been one of the key tree species increasing in dominance. Aspen encroachment is best explained not by single mechanism, but rather by a number of interacting ecological factors. In this study I examined the ecological consequences of the clonal biology in aspen as a means to explain persistence and observed tree-grass ratios within grassland savannas of western Canada. Results suggest that aspen stands cycle between a “stable” phase characterized by a dense mature canopy, and an “unstable” phase characterized by canopy breakup and increased regeneration from root suckers. It is during this unstable phase that clonal encroachment is likely to occur. Within these mature stands, different-aged ramets promote persistence by maximizing developmental variation. Such a mechanism overcomes the functional phenotypic uniformity of ramets within a single age-structured stand. Results from my study indicate that physiological integration is beneficial to the growth and survivorship of regenerating and encroaching aspen ramets. My results also suggest that the benefits of physiological integration are greatest in more stressful environments, and in recently established post-fire ramets. Finally, the consequences of variation in adaptive ecological relevant traits among individuals was examined by determining differences in vigor among aspen clones in relation to the production of secondary compounds (phenolic glycosides). My results demonstrate a high degree of variation in leaf phenolic glycosides production among clones. A significant amount of this variation was accounted for by differences in clone vigor (within population: individual susceptibility hypothesis), with a smaller amount related to environmental differences (among populations). In all instances, vigorous clones were significantly higher in levels of phenolic glycosides compared with dieback clones, suggesting that some individuals may be predisposed to undergo density-independent mortality. This has important ecological implications, because it implies that one of the key mechanisms regulating population dynamics, community interactions and biodiversity may be related to intrinsic adaptive differences in susceptibility among individuals.

To understand the temporal dynamics of a forest, long-term direct observations are required. My study examined the long-term persistence of trembling aspen (Populus tremuloides Michx) and white spruce (Picea glauca (Moench) Voss) in the boreal mixed-wood forests of Riding Mountain National Park. A set of 266, disturbance-free, permanent sample plots were established in 1947 (stand age = 120 years) and followed through time for 55 years. My results indicate that although the density and basal areas of aspen do decline over the 55-year period, a successful regeneration and establishment occurs around 140 years. The long-term persistence of aspen is a result of clonal reproduction following the canopy breakup beginning around 130 years or earlier. This implies that the long-term persistence of both aspen and spruce occur and the expected succession to softwood dominance does not occur.

The encroachment of woody species into grassland and savanna ecosystems has been well document since the early 1800s. Within the parkland ecoregion of western Canada, trembling aspen (Populus tremuloides Michx.) has been one of the key tree species increasing in dominance. Aspen encroachment is best explained not by single mechanism, but rather by a number of interacting ecological factors. In this study I examined the ecological consequences of the clonal biology in aspen as a means to explain persistence and observed tree-grass ratios within grassland savannas of western Canada. Results suggest that aspen stands cycle between a “stable” phase characterized by a dense mature canopy, and an “unstable” phase characterized by canopy breakup and increased regeneration from root suckers. It is during this unstable phase that clonal encroachment is likely to occur. Within these mature stands, different-aged ramets promote persistence by maximizing developmental variation. Such a mechanism overcomes the functional phenotypic uniformity of ramets within a single age-structured stand. Results from my study indicate that physiological integration is beneficial to the growth and survivorship of regenerating and encroaching aspen ramets. My results also suggest that the benefits of physiological integration are greatest in more stressful environments, and in recently established post-fire ramets. Finally, the consequences of variation in adaptive ecological relevant traits among individuals was examined by determining differences in vigor among aspen clones in relation to the production of secondary compounds (phenolic glycosides). My results demonstrate a high degree of variation in leaf phenolic glycosides production among clones. A significant amount of this variation was accounted for by differences in clone vigor (within population: individual susceptibility hypothesis), with a smaller amount related to environmental differences (among populations). In all instances, vigorous clones were significantly higher in levels of phenolic glycosides compared with dieback clones, suggesting that some individuals may be predisposed to undergo density-independent mortality. This has important ecological implications, because it implies that one of the key mechanisms regulating population dynamics, community interactions and biodiversity may be related to intrinsic adaptive differences in susceptibility among individuals.

To understand the temporal dynamics of a forest, long-term direct observations are required. My study examined the long-term persistence of trembling aspen (Populus tremuloides Michx) and white spruce (Picea glauca (Moench) Voss) in the boreal mixed-wood forests of Riding Mountain National Park. A set of 266, disturbance-free, permanent sample plots were established in 1947 (stand age = 120 years) and followed through time for 55 years. My results indicate that although the density and basal areas of aspen do decline over the 55-year period, a successful regeneration and establishment occurs around 140 years. The long-term persistence of aspen is a result of clonal reproduction following the canopy breakup beginning around 130 years or earlier. This implies that the long-term persistence of both aspen and spruce occur and the expected succession to softwood dominance does not occur.

Στην παρούσα διπλωματική εργασία εφαρμόζουμε τις αρχές της Θεωρίας Παιγνίων και πιο συγκεκριμένα τις έννοιες των Ισορροπιών Nash και των Παιγνίων Εξισορρόπησης Φορτίου, με σκοπό να αναλύσουμε την επίδραση που έχει στην απόδοση των δικτύων και των κατανεμημένων συστημάτων η εγωιστική και ανταγωνιστική συμπεριφορά των χρηστών τους. Πρώτα εξετάζουμε το παίγνιο της εξισορρόπησης φορτίου με τρεμάμενο χέρι σε ταυτόσημες μηχανές ως προς την ύπαρξη αγνών ισορροπιών Nash. Δείχνουμε πως υπάρχει πάντα μία αγνή ισορροπία Nash με αναγωγή από τα αποτελέσματα για τα παίγνια εξισορρόπησης φορτίου. Έπειτα, δίνουμε αλγόριθμο πολυωνυμικού χρόνου για τον υπολογισμό της ισορροπίας αυτής. Τέλος, εξετάζουμε το κόστος της Αναρχίας του παιγνίου. Το κόστος της Αναρχίας εκφράζει την απόκλιση της απόδοσης της χειρότερης Ισορροπίας Nash από την βέλτιστη απόδοση. Αποδεικνύουμε πως το κόστος της Αναρχίας του παιχνιδιού φράσσεται εκ των άνω από μία μικρή σταθερά. / In the present diploma thesis we will be using basic concepts of Game Theory, more specifically the concepts of Nash Equilibrium and Load Balancing Games, in order to analyse the effect of egoistic and competitive user's behaviour on the efficiency of networks and distributed systems. Firstly, we prove that the trembling hand load balancing game on identical machines always admits a pure Nash equilibrium. Secondly, we find an algorithm that computes this Nash equilibrium in polynomial time. Finally, we compare the social cost of pure equilibria with optimal solutions. This ratio is called pure price of Anarchy. We prove that the pure price of anarchy is bounded by a small constant factor.

Most clonal plants exhibit a modular structure at multiple levels. At the level of the organs, they are characterized by functional modules, such as, internodes, leaves, branches. At the level of the genetic individual (clone or genet), they possess independent evolutionary and physiological units (ramets). These evolutionary units arise through the widespread phenomenon of clonal reproduction, achieved in a variety of ways including rhizomes, stolons, bulbils, or lateral roots. The focus of this study was Populus tremuloides, trembling aspen, a dioecious tree that reproduces sexually by seed and asexually through lateral roots. Local forest patches in western populations of Populus tremuloides consisted largely of multiple genotypes. Multi-clonal patches were dominated by a single genotype, and in one population (Riske Creek) we found several patches (five out of 17) consisting of a single genotype. A second consequence of modularity is that during the repeated cycle of ramet birth, development and death, somatic mutations have the opportunity to occur. Eventually, the clone becomes a mosaic of mutant and non-mutant cell lineages. We found that neutral somatic mutations accumulated across 14 microsatellite loci at a rate of between 10^-6 and 10^-5 per locus per year. We suggest that neutral genetic divergence, under a star phylogeny model of clonal growth, is an alternative way to estimate clone age. Previous estimates of clone age couple the mean growth rate per year of shoots with the area covered by the clone. This assumes a positive linear relationship between clone age and clone size. We found, however, no repeatable pattern across our populations in terms of the relationship of either shape or size to the number of somatic changes. A final consequence of modularity is that during clonal growth, natural selection is relaxed for traits involving sexual function. This means that mutations deleterious to sexual function can accumulate, reducing the overall sexual fitness of a clone. We coupled neutral genetic divergence within clones with pollen fitness data to infer the rate and effect of mildly deleterious mutations. Mutations reduced relative sexual fitness in clonal aspen populations by about 0.12x10^-3 to 1.01x10^-3 per year. Furthermore, the decline in sexual function with clone age is evidence that clonal organisms are vulnerable to the effects of senescence. / Medicine, Faculty of / Medical Genetics, Department of / Graduate

Changes in forest ecosystem vegetation also bring about changes to the associated soil. In order to maintain forest productivity, it is important to know the effects of tree species upon the soil, especially the influence of deciduous versus coniferous tree species. Many deciduous species increase pH, nitrogen, base saturation and/or accumulation of organic matter in the forest floor. The chemical properties of the forest floor may, in turn, influence the chemical properties of the underlying mineral soil. If a tree species significantly alters the soil, then silviculturists may consider crop rotation between deciduous and coniferous trees or growing mixed-species stands to maintain greater nutrient availability and maintain site productivity. Trembling aspen (Populus tremuloides) and black spruce (Picea mariana) may occupy similar sites in the North American boreal forest. Shade-intolerant aspen is generally a seral species while shade-tolerant black spruce can be a seral species but also forms a major component in late successional stages. This study investigated differences in nitrogen-related soil properties between trembling aspen and black spruce stands on upland sites in the BWBS zone of northeastern BC. We asked two questions: (1) are the differences in soil nutrient properties manifested in both forest floor and mineral soil? (2) To what extent are these differences reflected in the floristic composition of understory vegetation?

No description available.

Accurate and reliable predictions of site index (height of dominant trees at a reference age, usually 50 years at breast-height) for timber crop species is essential for silvicultural site-specific decision making. Site index can be predicted from site quality once the relationship between site index and site quality has been quantified. Site quality is defined as the sum of all environmental factors affecting the biotic community, such as the factors directly influencing the growth of vascular plants (light, heat, soil moisture, soil nutrients, and soil aeration). Since these factors vary greatly in time, indirect estimates of site quality have widely been used as predictors for site index in various multiple regression models. Trembling aspen (Populus tremuloides Michx.) is the most widely distributed broadleaf species in British Columbia, especially in the Boreal White and Black Spruce (BWBS) biogeoclimatic zone. Growing this species for sustainable timber production requires a good understanding of its productivity attributes and accurate predictions of its growth. This extension note presents (1) relationships between trembling aspen site index and some indirect measures of site quality, and (2) site index prediction models using the indirect measures of site quality as predictors.

The boreal forest is confined to the Northern Hemisphere and is the most continuous and extensive forest in the world. In North America boreal forest extends from the Pacific to Atlantic coast spanning over 10° latitude. White spruce (Picea glauca (Moench) Voss), black spruce (P. mariana (Mill.) B.S.P.), and trembling aspen (Populus tremuloides Michx.) are among the dominant tree species. Black spruce and trembling aspen may form pure stands and occupy similar sites as their edaphic amplitudes overlap; however, spruce is rare on water-deficient sites and aspen does not tolerate excess water. Despite many studies conducted in the North American boreal forest, little is known about relationships between the boreal understory vegetation and softwood or hardwood canopy species in different climate regions. Furthermore, the variation in species diversity and succession between the stands dominated by coniferous trees and those dominated by broadleaved trees within the same region is unknown. The objectives of this study are to determine (1) the difference in the species diversity and floristic composition of understory vegetation between black spruce and trembling aspen stands within the same climatic region, and (2) how the species diversity and floristic composition of understory vegetation in each stand type vary with climate, and soil moisture and soil nutrient conditions.