Graduate Theses & Dissertations

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Factors Controlling Peat Chemistry and Vegetation Composition in Sudbury Peatlands after 30 Years of Emission Reductions
Peatlands are prevalent in the Sudbury, Ontario region. Compared with the well documented devastation to the terrestrial and aquatic ecosystems in this region, relatively little work has been conducted on the peatlands. The objective of this research was to assess factors controlling peat and plant chemistry, and vegetation composition in 18 peatlands in Sudbury after over 30 years of emission reductions. Peatland chemistry and the degree of humification varies considerably, but sites closer to the main smelter had more humified peat and the surface horizons were enriched in copper (Cu) and nickel (Ni). Copper and Ni concentrations in peat were significantly correlated with Cu and Ni in the plant tissue of leatherleaf, although the increased foliar metal content did not obviously impact secondary chemistry stress indicators. The pH and mineral content of peat were the strongest determining factors for species richness, diversity and community composition. The bryophyte communities appear to be acid and metal tolerant, although Sphagnum mosses are showing limited recovery. Author Keywords: anthropogenic emissions, bryophytes, community comspoition, heavy metals, peatlands, wetland vegetation
Purification and Identification of Selenium-containing C-phycocyanin from Spirulina
Selenium is an essential trace nutrient to many organisms, yet in high concentrations it is toxic. Organic selenium is more bioavailable to aquatic biota than inorganic selenium, but is usually found in much lower concentrations. Algae are known to biotransform inorganic selenium into several organo-selenium compounds, but it is unknown whether any of these bioaccumulate in the food chain. In this study, selenium was incorporated into the methionine residues of an algal photosynthetic protein, c-phycocyanin from Spirulina spp. The extent of selenium incorporation was quantified by inductively coupled plasma-mass spectrometry (ICP-MS), and the protein was identified using electrospray mass spectrometry (ES-MS). C-phycocyanin was isolated and purified from Spirulina with a final recovery of 20-30 % of the total c-phycocyanin present. Selenomethionine replaced 92.8% ± 1.22 of the methionine residues in c-phycocyanin when grown in 2.5 ppm sodium selenite. ES-MS was used to obtain protein spectra, and pure c-phycocyanin was identified. Data of full scans provided estimated masses of both protein subunits--α-chain measured at 18,036 Da; β-chain measured at 19,250 Da--close to the theoretical masses. Protein fragmentation by collision-induced dissociation and electron capture dissociation provided approximately 52 % amino acid sequence match with c-phycocyanin from Spirulina platensis. This study demonstrates the incorporation of selenium into an algal protein, and the identification of c-phycocyanin using electrospray ionization-mass spectrometry. Author Keywords:
Cytokinin Oxidase/Dehydrogenase (CKX) Gene Family in Soybeans (Glycine max)
Glycine max (soybean) is an economically important plant species that registers a relatively low yield/seed weight compared to other food and oil seed crops due to higher rates of flower and pod abortion. Alleviation of this abortion rate can be achieved by altering the sink strength of the reproductive organs of soybeans. Cytokinin (CK) plays a fundamental role in promoting growth of sink organ (flowers and seeds) by increasing the assimilate demand. Cytokinin oxidase/dehydrogenase (CKX) is an enzyme that catalyses the irreversible breakdown of active CKs and hence reduce the cytokinin content. The current thesis uncovers the members of CKX gene family in soybeans and the natural variations among CKX genes within soybean varieties with different yield characteristics. The identification of null variants of OsCKX2 that resulted in large yield increases by Ashikari et al. (2005) provided a rationale for current thesis. The soybean CKX genes along with the ones from Arabidopsis, Rice and Maize were used to construct a phylogenetic tree. Using comparative phylogeny, protein properties and bioinformatic programs, the potential effect of the identified natural variations on soybean yield was predicted. Five genes among the seventeen soybean CKXs identified, showed polymorphisms. One of the natural variations, A159G, in the gene GmCKX16 occurred close to the active site of the protein and was predicted to affect the activity of enzyme leading to higher accumulation of CKs and hence increased seed weight. Use of such natural variations in marker assisted breeding could lead to the development of higher yielding soybean varieties. Author Keywords: CKX, Cytokinins, Seed weight, Seed Yield, SNPs, Soybeans
Early Responses of Understory Vegetation to Above Canopy Nitrogen Additions in a Jack Pine Stand in Northern Alberta
Abstract Early Responses of Understory Vegetation After One Year of Above Canopy Nitrogen Additions in a Jack Pine Stand in Northern Alberta Nicole Melong Nitrogen (N) emissions are expected to increase in western Canada due to oil and gas extraction operations. An increase in N exposure could potentially impact the surrounding boreal forest, which has adapted and thrived under traditionally low N deposition. The majority of N addition studies on forest ecosystems apply N to the forest floor and often exclude the important interaction of the tree canopy. This research consisted of aerial NH4NO3 spray applications (5, 10, 15, 20, 25 kg N ha-1yr-1) by helicopter to a jack pine (Pinus banksiana Lamb.) stand in the Athabasca Oil Sands Region (AOSR) in northern Alberta, Canada. The main objective was to assess the impacts of elevated N after one year of treatment on the chemistry of understory vegetation, which included vascular plants, terricolous lichens, epiphytic lichens and a terricolous moss species. Changes in vegetation chemistry are expected to be early signs of stress and possible N saturation. Increased N availability is also thought to decrease plant secondary compound production because of a tradeoff that exists between growth and plant defense compounds when resources become available. Approximately 60% of applied N reached the ground vegetation in throughfall (TF) and stemflow (SF). Nitrate was the dominant form of N in TF in all treated plots and organic N (ON) was the dominant form of N in SF in all plots. The terricolous non-vascular species were the only understory vegetation that responded to the N treatments as N concentration increased with increased treatment. Foliar chemistry of the measured epiphytic lichens, vascular species, and jack pine was unaffected by the N treatments. Based on biomass measurements and N concentration increases, the non-vascular terricolous species appear to be assimilating the majority of TF N after one year. Vegetation from the high treatment plot (25 kg N ha-1yr-1) was compared to a jack pine forest receiving ambient high levels of N (21 kg N ha-1yr-1) due to its proximity to Syncrude mining activities. Nitrogen concentrations in plant tissues did not differ between the two sites; however, other elements and compounds differed significantly (Ca, Mg, Al, Fe). After one year of experimental N application, there were no environmental impacts consistent with the original N saturation hypothesis. Author Keywords: Athabasca Oil Sands Region, Canopy Interactions, Jack Pine, Nitrogen, Secondary Chemistry, Understory Vegetation
Fungi and Cytokinins
Cytokinin biosynthesis in organisms aside from plant species has often been viewed as a byproduct of tRNA degradation. Recent evidence suggests that these tRNA degradation products may actually have a role in the development of these organisms, particularly fungi. This thesis examines the importance of cytokinins, a group of phytohormones involved in plant cell division and differentiation as well as the phytohormone abscisic acid, involved in plant response to environmental factors, and their presence and role in fungi. An initial survey was conducted on 20 temperate forest fungi of differing nutritional modes. Using HPLC-ESI MS/MS, cytokinin and abscisic acid were detected in all fungi regardless of their mode of nutrition or phylogeny. The detection of the same seven CKs across all fungi suggested the existence of a common CK biosynthetic pathway and dominance of the tRNA pathway in fungi. Further, the corn smut fungus Ustilago maydis is capable of producing CKs separate from its host and different U. maydis strains induce disease symptoms of differing severity. To determine if CK production during infection alters disease development a disease time course was conducted on cob tissue infected with U. maydis dikaryotic and solopathogenic strains. Dramatic changes in phytohormones including an increase in ABA followed by increases in cisZCKs were detected in tumour tissue particularity in the more virulent dikaryon infection, suggesting a role for CKs in strain virulence. Mining of the U. maydis genome identified a sole tRNA-isopentenyltransferase, a key enzyme in CK biosynthesis. Targeted gene deletion mutants were created in U. maydis which halted U. maydis CK production and decreased pathogenesis and virulence in seedling and cob infections. CK and ABA profiling carried out during disease development found that key changes in these hormones were not found in deletion mutant infections and cob tumour development was severely impaired. These findings suggested that U. maydis CK production is necessary for tumour development in this pathosystem. The research presented in this thesis highlights the importance of fungal CKs, outlines the dominant CK pathway in fungi, identifies a key enzyme in U. maydis CK biosynthesis and reveals the necessity of CK production by U. maydis in the development of cob tumours. Author Keywords: abscisic acid, cytokinins, high performance liquid chromatography-electrospray ionization tandem mass spectrometry, tRNA degradation pathway, Ustilago maydis, Zea mays
Bioremoval of copper and nickel on living and non-living Eugelna gracilis
This study assesses the ability of a unicellular protist, Euglena gracilis, to remove Cu and Ni from solution in mono- and bi-metallic systems. Living Euglena cells and non-living Euglena biomass were examined for their capacity to sorb metal ions. Adsorption isotherms were used in batch systems to describe the kinetic and equilibrium characteristics of metal removal. In living systems results indicate that the sorption reaction occurs quickly (<30 min) in both Cu (II) and Ni (II) mono-metallic systems and adsorption follows a pseudo-second order kinetics model for both metals. Sorption capacity and intensity was greater for Cu than Ni (p < 0.05) and were described by the Freundlich model. In bi-metallic systems sorption of both metals appears equivalent. In non-living systems sorption occurred quickly (10-30 min) and both Cu and Ni equilibrium uptake increased with a concurrent increase of initial metal concentrations. The pseudo-first-order model was applied to the kinetic data and the Langmuir and Freundlich models effectively described single-metal systems. The biosorption capacity of Cu (II) and) was 3x times greater than that of Ni (II). Sorption of one metal in the presence of relatively high concentrations of the other metal was supressed. Generally, it was found that living Euglena remove Cu and Ni more efficiently than non-living Euglena biomass in both mono- and bi-metallic systems. It is anticipated that this work should contribute to the identification of baseline uptake parameters and capacities for Cu and Ni by Euglena as well as to the increasing amount of research investigating sustainable bioremediation. Author Keywords: accumulation, biosorption, Cu, Euglena gracilis, kinetics, Ni
Hormonal Algae
Based on an endogenous hormone study, three cytokinin type phytohormones; benzyladenine (BA), trans-zeatin (tZ) and methylthiol trans-zeatin (MeSZ), as well as abscisic acid (ABA) were exogenously added at three concentrations (10-7, 10-6 and 10-5 M) to cultures of Chlorella vulgaris in an attempt to alter growth rate, total lipid and fatty acid yields and fatty acid profile. Growth stimulation was highest at 10-6 M for BA, MeSZ and ABA and 10-5 M for tZ. All treatments caused changes in total lipid and fatty acid content, with BA causing an increase to lipid content. The most significant change in the fatty acid profile was observed with the addition of MeSZ at 10-7 and 10-6 M causing increases of 204% and 457% in linolenic acid respectively above the control. These results are novel and potentially highly impactful, as MeSZ has never been added exogenously to algae and may be used to stimulate overproduction of linolenic acid for pharmaceutical or industrial purposes. Author Keywords: Abscisic Acid, Chlorella vulgaris, Cytokinin, Fatty acid, Linolenic Acid, Methylthiol trans-Zeatin
Carbon Exchange along a Natural Gradient of Deciduous Shrub Coverage in the Low-Arctic
Arctic terrestrial ecosystems have experienced substantial structural and compositional changes in response to warming climate in recent decades, especially the expansion of shrub species in Arctic tundra. Climatic and vegetation changes could feedback to the global climate by changing the carbon balance of Arctic tundra. The objective of this thesis was to investigate the influence of increased shrub coverage on carbon exchange processes between atmosphere and the Arctic tundra ecosystem. In this study a space-for-time substitution was used, referred to as a shrub expansion “chronosequence”, with three sites along a natural gradient of deciduous shrub coverage in the Canadian low Arctic. Leaf-level photosynthetic capacity (Amax) of dominating birch shrub Betula glandulosa (Michx.) was significantly higher (P<0.05) at the site where shrubs were more abundant and taller than at the other sites. For all sites, mean Amax in 2014 was significantly lower than in 2013, in part potentially due to differences in precipitation distribution. Bulk soil respiration (RS) rate was significantly higher (P<0.05) at the site with more shrubs compared with the other sites. The differences in RS across sites appeared to be driven by differences in soil physiochemical properties, such as soil nitrogen and soil bulk density rather than soil microclimate factors (e.g. soil temperature, moisture). The three sites were either annual CO2 sources (NEP<0) to the atmosphere or CO2 neutral, with strongest annual CO2 sources (-44.1±7.0 g C m-2) at the site with most shrubs. Overall this study suggests that shrubs tundra carbon balance will change with shrub expansion and that shrub ecosystems in the Arctic currently act as annual carbon sources or neutral to the atmospheric CO2 and further shrub expansion might strengthen the CO2 emissions, causing a positive feedback to the warming climate. Author Keywords: arctic tundra, carbon exchange, climate change, photosynthetic capacity, shrub expansion, soil respiration
Time-dependent effects of predation risk on stressor reactivity and growth in developing larval anurans (Lithobates pipiens)
The predator vs. prey dynamic is an omnipresent factor in ecological systems that may drive changes in life history patterns in prey animals through behavioural, morphological, and physiological changes. Predation risk can have profound effects on the life history events of an animal, and is influenced by the neuroendocrine stress response. Activation of the hypothalamic-pituitary-adrenal/interrenal axis, and the induction of stress hormones (e.g., corticosterone (CORT)) have been shown to mediate the onset of inducible anti-predator defensive traits including increased tail-depth, and reduced activity. The predator-prey relationship between dragonfly nymphs and tadpoles can be a powerful model system for understanding mechanisms that facilitate changes in the stress response in accordance with altered severity of risk. It has been well demonstrated early in tadpole ontogeny that increased corticosterone (CORT) levels, observed within three weeks of predator exposure, are correlated with increased tail depth morphology. However, the reactivity of the stress response in relation to the growth modulation in developing prey has yet to be fully explored. Accordingly, this thesis assessed the stress and growth response processes in tadpoles that were continuously exposed to perceived predation risk later in ontogeny. Continuous exposure of prey to predation risk for three weeks significantly increased CORT levels, and tail depth. However, tadpoles exposed to six weeks of predation risk acclimated to the presence of the predator, which was observed as a significant reduction of stressor-induced CORT levels. In addition, although increased tail depth has been attributed to predator defense, predator-naïve tadpoles began to display similar tail depth morphology as treated tadpoles at the six week time point. Thus, this thesis suggests that the stress response in lower vertebrate systems (e.g., tadpoles) may operate in a similarly complex manner to that observed in higher vertebrates (e.g., rats), for which severity of risk associated with the stressor aids in defining activity of the stress response. Moreover, the lack of morphological difference between treatments among tadpoles exposed later in ontogeny suggests that the mechanisms for inducing defenses are normal morphological traits in the development of the animal. This thesis paves the way for future research to elucidate the relationship between the neuroendocrine stress response and hormonal pathways involved in growth modulation in the presence of environmental pressures. Author Keywords: Acclimation, Corticosterone, Growth Modulation, Predation Risk, R. pipiens, Tadpole
Regulation of Cytokinins During Kernel Development in High and Low Yielding Oat and Barley Lines
Cytokinins (CKs) are a family of plant phytohormones responsible for many areas of plant growth and development. There are four free base types of CKs found in higher plants, trans-zeatin (tZ), N6-(∆2-isopentenyl)adenine (iP), cis-Zeatin (cZ) and dihydrozeatin (DZ). CK biosynthesis is regulated by adenosine phosphate-isopentenyltransferase (IPT), which is encoded by a multi-gene family in many plant species. There are two types of IPT pathways responsible for CK production, the tRNA pathway and the AMP (ATP/ADP) pathway. The tRNA pathway putatively produces cZ and the latter predominantly produces iP type nucleotides. CKs have long been studied for their role in stress tolerance, signal transduction, and involvement in many areas of plant growth and development. This study focuses on the role of CKs and CK biosynthesis by IPT during kernel development and comparisons of its regulation in high and low yielding barley and oat lines. The sequence of a putative IPT encoding gene in barley and oat was identified by a blast search of other known IPT gene fragments in closely related species. Quantitative Real time PCR results based on primers designed for the putative barley and oat IPT gene revealed changes in expression of IPT during different stages of kernel development, but no significance difference was associated with yield. Correlation of IPT gene expression in barley with cZ CK profiles measured by HPLC-MS/MS, confirms a putative IPT gene is a tRNA- IPT. HPLC-MS/MS results reveal some CK types, such as benzyladenine, are more predominant in higher yielding lines. This suggests different types of CKs play a role in yield production. Future studies on more IPT genes in the barley and oat IPT gene family will outline a more clear representation of the role of IPT in barley kernel development. Author Keywords: Benzyladenine, Cereal grain, Cytokinin, Isopentenyl Transferase, Mass Spectrometry, Real Time PCR
Involvement of Endogenous Plant Hormones in The Regulatory Network of Fatty Acid Biosynthesis in Soybean Seed
The activities of phytohormones during the reproductive phase have been partially clarified in seed physiology while the biological role of plant hormones in oil accumulation during seed development has been investigated in part only. In this research, fatty acid (FA) contents and hormone profiles, including abscisic acid (ABA) and cytokinins (CKs) of seed samples in four different stages and comparing six soybean varieties have been investigated in order to examine the hypothesis that the endogenous plant hormones play important roles in FA production in soybean seeds. The FA contents increased significantly during this period while the hormone concentrations gradually declined towards the seed physical maturation. However, the interactions between FA contents and hormone profiles were complex and went beyond linear correlations. Hormone metabolism in the earlier stages of seed maturation period demonstrated numerous robust relationships with FA accumulations, as derived from several simple and multiple regression models in the determination of different FA contents. Evaluation of the effects of exogenous ABA and trans-Zeatin (tZ) on FA biosynthesis has revealed that ABA appears to be involved in the accumulations of unsaturated FAs while tZ participated in the synthesis of saturated and unsaturated FAs. Notably, the alterations of FA synthesis differ according to what exogenous hormone concentrations could be used. Author Keywords: Abscisic acid, Cytokinin, Fatty acid, Seed development, Soybean
Exploring reproduction in wild blue lupine (Lupinus perennis) in comparison to L. polyphyllus and L. albus
Wild lupine (Lupinus perennis) restoration efforts seek to increase and connect populations, using seeds, to facilitate the recovery of endangered butterflys in Ontario. This study observed plant growth and phytohormone levels of L. albus, L. polyphyllus, and L. perennis through stages of seed development, each with varying strategies in growth and reproductive investment. L. polyphyllus is similar to L. perennis in morphology, acting as similar comparable with L. albus, a well-studied annual, as an outgroup comparator. Wild lupines showed a lack of sexual reproductive effort as they did not put as much effort into above ground growth, and few in the population reproduces. They also showed cis-zeatin, a weaker cytokinin, throughout development and had higher amounts of abscisic acid at the end of seed maturity, impacting their ability to develop and germinate. These factors contribute to why wild lupines are difficult to restore using seeds, limiting expansion and challenging restoration. Author Keywords: L. albus, L. perennis, L. polyphyllus, plant physiology, seed development, Wild blue lupine

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