Plant acclimation to natural and intensified solar UV-B irradiance was investigated in three species, Oenothera stricta Ledeb., Rumex obtusifolius L., and R . patientia L. The objectives of this study were to determine: (1) the relationship between plant sensitivity and epidermal UV attenuation, (2) the effect of phenotypic changes in the leaf epidermis, resulting from UV-B exposure, on plant sensitivity to UV radiation, and (3) the plasticity of these changes in the epidermis leading to plant acclimation to UVB radiation. Epidermal UV transmittance was found to differ in magnitude and spectral distribution among the three species examined in this study. Epidermal tissue from the leaves of Oenothera stricta, Rumex obtusifolius, and R. patientiaattenuated up to 951, 90%, and 851 of the UV-B radiation incident on the leaf, respectively. The spectral distribution of transmittance appeared to be characteristic of each species. The capacity of the epidermis to attenuate UV-B radiation was found to have some degree of plasticity in Oenothera stricta and Rumex obtusifolius. After exposure to UV-B radiation for periods of 11 to 15 days, at a mean dose rate of approximately 2050 biologically effective J·m·-2d-1 , epidermal UV-B transmittance was significantly reduced by up to 36% in mature leaves of O. stricta. Increased capacity of the epidermis to attenuate UV-B radiation was not observed in young leaves of this species. These leaves only transmitted about 4% of the UV-8 radiation incident on the leaf, The transmittance of shorter wavelengths of visible radiation was reduced by 6 to 14% in young and mature leaves after UV-B irradiation. A similar reduction in epidermal UV-8 transmittance in the leaves of R. obtusifolius was also observed. Ultraviolet absorbance in leaf epidermal and mesophyll tissue of Oenothera stricta generally increased in response to UV-B irradiation. Absorbance increased more in young leaves than mature leaves after UV-B irradiation. This increase in UV absorbance was also found in mature leaves of Rumex obtusifolius and R. patientia after UV-B irradiation. The increase in absorbance was most likely caused by an increase in flavonoid and related phenolic compounds in leaf tissues. The rate of photosynthesis was used as an indicator of the degree of plant sensitivity to UV-B radiation. In general, photosynthesis was not significantly depressed in the leaves of any of the three species. A trend toward photosynthetic depression in response to UV-B irradiation was found, however, and thus some degree of UV-B sensitivity is suggested in these species. A limited degree of plant acclimation was suggested in plants that were exposed to a low UV-B dose rate prior to a higher dose rate. A mechanism of UV-B attenuation, possibly involving the biosynthesis of UV-ab sorbing flavonoid compounds in the epidermis and mesophyll under the stress of UV-B radiation, and a subsequent increase in the UV-B attenuation capacity of the epidermis, is suggested. The degree of plant sensitivity and acclimation to natural and intensified solar UV-B radiation may involve a dynamic balance between the capacity for UV-B attenuation and UV-radiation-repair mechanisms in the leaf.

The evolution of synapses from early proto-synaptic protein complexes in unicellular eukaryotes to sophisticated machines comprising thousands of proteins parallels the emergence of finely tuned synaptic plasticity, a molecular correlate for memory and learning. Phenotypic change in organisms is ultimately the result of evolution of their genotype at the molecular level. Selection pressure is a measure of how changes in genome sequence that arise though naturally occurring processes in populations are fixed or eliminated in subsequent generations. Inferring phylogenetic information about proteins such as the variation of selection pressure across coding sequences can provide valuable information not only about the origin of proteins, but also the contribution of specific sites within proteins to their current roles within an organism. Recent evolutionary studies of synaptic proteins have generated attractive hypotheses about the emergence of finely-tuned regulatory mechanisms in the post-synaptic proteome related to learning, however, these analyses are relatively superficial. In this thesis, I establish a scalable molecular phylogenetic modelling framework based on three new inference methodologies to investigate temporal and spatial aspects of selection pressure changes for the whole human proteome using protein orthologs from up to 68 taxa. Temporal modelling of evolutionary selection pressure reveals informative features and patterns for the entire human proteome and identifies groups of proteins that share distinct diversification timelines. Multi-ontology enrichment analysis of these gene cohorts was used to aid biological interpretation, but these approaches are statistically under powered and do not capture a clear picture of the emergence of synaptic plasticity. Subsequent pathway-centric analysis of key synaptic pathways extends the interpretation of temporal data and allows for revision of previous hypotheses about the evolution of complex synaptic function. I proceed to integrate inferred selection pressure timeline information in the context of static protein-protein interaction data. A network analysis of the full human proteome reveals systematic patterns linking the temporal profile of proteins’ evolution and their topological role in the interaction graph. These graphs were used to test a mechanistic hypothesis that proposed a propagating diversification signal between interactors using the temporal modelling data and network analysis tools. Finally, I analyse the data of amino-acid level spatial modelling of selection pressure events in Arc, one of the master regulators of synaptic plasticity, and its interactors for which detailed experimental data is available. I use the Arc interactome as an example to discuss episodic and localised diversifying selection pressure events in tightly coupled complexes of protein and showcase potential for a similar systematic analysis of larger complexes of proteins using a pathway-centric approach. Through my work I revised our understanding of temporal evolutionary patterns that shaped contemporary synaptic function through profiling of emergence and refinement of proteins in multiple pathways of the nervous system. I also uncovered systematic effects linking dependencies between proteins with their active diversification, and hypothesised about their extension to domain level selection pressure events.