Phenotypic plasticity, the ability of a single genotype to be expressed as a range of phenotypes in response to environmental variation, is a widespread phenomenon. Documented increasingly among the larval stages of marine organisms, phenotypic plasticity in the veliger larvae of the marine snail Littorina scutulata was investigated in response to predatory, nutritional, and gravitational stimuli. Veligers developed rounder shells, smaller apertures, and reinforced aperture margins in response to water-borne cues from predatory crab larvae. The nature and degree of the induced-morphologies depended on cue composition and conferred decreased vulnerability to predation. Food-limited veligers developed larger feeding and swimming structures (vela) with longer cilia relative to shell size compared to larvae raised with high food. This inducible offense corresponded with a decrease in vertical swimming speed, an unexpected result possibly reflecting behavioral manipulation of individual velar components. A cell proliferation assay indicated that growth of the larger structure was achieved partially by a steady rate of cell division over a longer period of time; an initially higher level of cell proliferation in veligers raised on high food dropped off sharply. Velar lobe asymmetry, where one lobe is larger than the other, may exist to offset an asymmetry in weight distribution due to how the larval shell is carried. The larger velar lobe overlies the protruding spire of the larval shell. Bi- and multi-lobed vela get bigger with shell size but follow different rules with regards to the relationship between velar asymmetry and shell asymmetry. Experimental alternations of mass distribution of the larval shell caused changes in the ratio of area between each side of the velum and total velar growth for larvae of L. scutulata. Following settlement and metamorphosis, juveniles of intertidal marine invertebrates are subject to additional stressors that can manifest as phenotypic variation. Color differences between juvenile and adult Strongylocentrotus purpuratus were shown to be caused by variation in light exposure. Green juveniles raised in sunlight turned purple (due to more pigment) and showed decreased susceptibility to artificial UVR than urchins kept in the dark, which remained green (due to less pigment). This dissertation includes previously unpublished co-authored material.

Virtually all multicellular organisms are capable of developing differently in response to environmental variation. At the molecular level, such developmental plasticity requires interpretation and perpetuation of environmental signals without changing the underlying genotype. Such non-genetic, heritable information is known as epigenetic information. This dissertation examines epigenetic information among social insects, and how differences in such information relate to phenotypic caste differences. The studies included herein primarily focus on one form of epigenetic information: DNA methylation. In particular, these studies explore DNA methylation as it relates to and impacts (i) alternative phenotype and particular gene expression differences in two social insect species, (ii) histone modifications, another important form of epigenetic information, in insect genomes, and (iii) molecular evolutionary rate of underlying actively transcribed gene sequences. We find that DNA methylation exhibits marked epigenetic and evolutionary associations, and is associated with alternative phenotype in multiple insect species. Thus, DNA methylation is emerging as one important epigenetic mediator of phenotypic plasticity in social insects.

Forensic entomology is a well-established science linking arthropod biology and ecology to legal investigations. Specifically, immature development on a decomposing corpse may give insight into the minimum time elapsed since death. Until recently, biological variation within a single species has been overlooked when estimating colonization events. Variation in the form of phenotypic plasticity, or the ability of a single genotype to produce multiple phenotypes under alternative stresses, has been documented in genetic and ecological literature and spans across all phyla. Taking this into account, different subpopulations of forensically pertinent insect species should also possess the ability to adapt to changing environments as geographic distribution increases. Thus, plastic responses of a species to alternative stresses may be measured in biological parameters, such as development time. In this research, three geographically distinct strains of the blow fly Cochliomyia macellaria Fabricius (Diptera Calliphoridae) were reared in two distinct environments in order to measure development time, as well as pupal and adult masses. Strains exhibited genetic variance when compared to each other, and each strain exhibited variable responses across environments (phenotypic plasticity). Plasticity in the form of genotype by environment (GxE) interactions was also exhibited by C. macellaria, although consistent adherence to any single rule explaining ontogenetic trends was not apparent. This research supports the existence of intraspecific variation in a common blow fly of forensic importance. Results of this study will impact the forensic entomology community by encouraging the generation of either strain;specific developmental datasets or statistical models to minimize variation caused by genetic, environment, or GxE effects in order to compare developmental data across strains.

Evolutionary processes in any population depend upon patterns of phenotypic variation available to selection and their underlying heritability. In this thesis, I used the filamentous red seaweed Asparagopsis armata, with particular focus on its modularity, to test several key questions underlying its growth-form evolution in heterogeneous environments. I established that experimental manipulations of light quantity and quality mimicking variation in underwater light due to shading or depth induce growthform plasticity in A. armata that may be evolutionarily significant given its variability among clones. Current patterns of plasticity displayed by A. armata appear adaptive, moreover, given that a reciprocal transplant of phenotypes between light environments found densely-branched (phalanx-like) phenotypes to have higher relative growth rates than sparsely-branched (guerrilla-like) phenotypes in well-lit patches, but lower relative growth rates than the latter in shaded patches. Using the capacity for rapid growth as a proxy for fitness, multivariate selection analyses identified environment-dependent patterns of directional selection on single traits coupled with linear and nonlinear selection on multi-trait combinations that shape growth-form variation within patches of differing light intensity, thereby reinforcing plasticity across light environments. Quantitative genetic analyses, however, suggest that the modular iteration of genes in morphogenesis may limit further growth-form evolution in A. armata populations exposed to spatial heterogeneity in light by constraining thallus responses to environment-dependent selection. Last, heritable responses to artificial selection on growth-form variation among clonal cell-lineages revealed the surprising capacity for A.armata to circumvent genetic constraints inherent to its development by adapting to environmental change in the absence of sexually-generated variance among clones.

Evolutionary processes in any population depend upon patterns of phenotypic variation available to selection and their underlying heritability. In this thesis, I used the filamentous red seaweed Asparagopsis armata, with particular focus on its modularity, to test several key questions underlying its growth-form evolution in heterogeneous environments. I established that experimental manipulations of light quantity and quality mimicking variation in underwater light due to shading or depth induce growthform plasticity in A. armata that may be evolutionarily significant given its variability among clones. Current patterns of plasticity displayed by A. armata appear adaptive, moreover, given that a reciprocal transplant of phenotypes between light environments found densely-branched (phalanx-like) phenotypes to have higher relative growth rates than sparsely-branched (guerrilla-like) phenotypes in well-lit patches, but lower relative growth rates than the latter in shaded patches. Using the capacity for rapid growth as a proxy for fitness, multivariate selection analyses identified environment-dependent patterns of directional selection on single traits coupled with linear and nonlinear selection on multi-trait combinations that shape growth-form variation within patches of differing light intensity, thereby reinforcing plasticity across light environments. Quantitative genetic analyses, however, suggest that the modular iteration of genes in morphogenesis may limit further growth-form evolution in A. armata populations exposed to spatial heterogeneity in light by constraining thallus responses to environment-dependent selection. Last, heritable responses to artificial selection on growth-form variation among clonal cell-lineages revealed the surprising capacity for A.armata to circumvent genetic constraints inherent to its development by adapting to environmental change in the absence of sexually-generated variance among clones.

Phenotypic plasticity is the ability of the single genotype to pruduce multiple phenotypes in response to evironmental conditions. There are many factors affecting phenotypic plasticity. The aim of this thesis is to summarize the current knowledge of phenotypic plasticity of aquatic insects with emphasis on the role of temperature and food availability. The review is complemented by a laboratory experiments designed to investigate the effect of temperature and food availability on growth and development rate of diving beetle Acilius canaliculatus (Coleoptera: Dytiscidae). The results are discussed in the light of the ecological concepts of temperature-size rule and developmental isomorphy.

Phenotypic plasticity is the ability of the single genotype to pruduce multiple phenotypes in response to evironmental conditions. There are many factors affecting phenotypic plasticity. The aim of this thesis is to summarize the current knowledge of phenotypic plasticity of aquatic insects with emphasis on the role of temperature and food availability. The review is complemented by a laboratory experiments designed to investigate the effect of temperature and food availability on growth and development rate of diving beetle Acilius canaliculatus (Coleoptera: Dytiscidae). The results are discussed in the light of the ecological concepts of temperature-size rule and developmental isomorphy.

Abstract Plants have evolved various adaptive strategies for balancing the benefits and costs of having a high affinity for resources, plasticity of growth allocation and mycorrhizal symbiosis. The relative growth rates of mycorrhizal and non-mycorrhizal plants were modelled for stable and variable nutrient availability. Mycorrhizal plants had higher growth rates at low and non-mycorrhizal plants at high nutrient availability. Variation in nutrient availability reduced the growth rate of mycorrhizal plants due to a high affinity for nutrients. However, mycorrhizal plants may be able to buffer against external fluctuations and therefore experience less environmental variation than non-mycorrhizal plants. Non-mycorrhizal plants may even benefit from variation. The optimal allocation of growth between shoot and roots depends on the availability of energy and nutrients. The optimisation model predicted that the requirement for phenotypic plasticity of shoot/root allocation is greatest in environments with low resource availability. Plants with a high affinity for resources required more plasticity in order to tolerate variation than plants with a low affinity. The model predicted a trade-off between the ability to deplete resources and the ability to tolerate resource fluctuations. Changes in the availability and ratio of resources lead to changes in the structure and composition of vegetation during primary succession. The field study of the forested phases of the land uplift island Hailuoto showed a successional change in the vegetation from the dominance of bryophytes and deciduous dwarf shrubs to dominance by lichens and evergreen dwarf shrubs. The humus layer became thinner and the availability of nutrients declined, while the C/N ratio of soil organic matter increased during succession indicating a decline in the quality of organic matter. The increased soil respiration rate indicates a successional increase in the energetic costs of decomposing organic matter. Nutrients mediate both direct and indirect trophic interactions. Indirect interactions of nutrient cycling are not explicit in continuous time models. A transformation to a discrete time model was shown to make the indirect interactions explicit as transition probabilities and allowed their dynamic contribution to be evaluated with an elasticity analysis. The importance of indirect interactions was greater in tundra than temperate forest and increased with the rate of nutrient cycling.

No description available.

With global climates changing rapidly, animals must adapt to new environmental conditions with altered weather and phenology. Key to adapting to these new conditions is adjusting the timing of reproduction to have offspring when the conditions are best to maximize growth and survival. Using a long-term dataset on a wild population of yellow-bellied marmots (Marmota flaviventer) at the Rocky Mountain Biological Laboratory (RMBL), we investigated how the timing of reproduction changed with changing spring conditions over the past 50 years. Marmots are hibernators with a four-month active season. It is thus crucial to reproduce early enough in the season to have time to prepare for hibernation, but not too early so as snow cover prevents access to food. Importantly, climate change in this area has increased spring temperatures by 5 °C and decreased spring snowpack by 50 cm over the past 50 years. This directional change in climate may have caused adaptation. Given that adaptation to environmental conditions could arise from either microevolution or phenotypic plasticity, we evaluated how female marmots adjust the timing of their reproduction and estimated the importance of both genetic variance and plasticity in the variation in this timing. We show that, within a year, the timing of reproduction is not as tightly linked to the date a female emerges from hibernation as previously thought. We report a positive effect of spring snowpack but not of spring temperature on the timing of reproduction. There is inter-individual variation in the timing of reproduction but not in its response to changing spring conditions. Genetic variance in the timing of reproduction is low, and heritability was 8%. Earlier pup emergence date increases the number and weighted proportion of pups surviving their first winter, indicative of directional selection on this trait. The same pattern is not found for litter size with no effect of pup emergence date on the number of pups born. Further, all three of these traits are not under stabilizing selection. Taken together, it seems that we should expect some changes in this population with changing climatic conditions, but because of plasticity and not due to natural selection. Further, future studies on the marmots should not operate under the assumption that females reproduce immediately following their emergence.