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Title: IDENTIFICATION OF LTR RETROTRANSPOSONS, EVALUATION OF GENOME ASSEMBLY, AND MODELING RICE DOMESTICATION
Creator: Ou, Shujun
Date: 2018
Collection: Electronic Theses & Dissertations
Description: The majority of fundamental theories in genetics and evolution were proposed prior to the discovery of DNA as the genetic material in 1952. Those include Darwin’s theory of evolution (1859), Mendelian genetics (1865), Wright and Fisher’s population genetics (1918), and McClintock’s transposition of genetic elements (1951). Nevertheless, the underlining mechanisms of those theories were not fully elucidated till the appearance of DNA sequencing technology. At present, technological advances...
Show moreThe majority of fundamental theories in genetics and evolution were proposed prior to the discovery of DNA as the genetic material in 1952. Those include Darwin’s theory of evolution (1859), Mendelian genetics (1865), Wright and Fisher’s population genetics (1918), and McClintock’s transposition of genetic elements (1951). Nevertheless, the underlining mechanisms of those theories were not fully elucidated till the appearance of DNA sequencing technology. At present, technological advances have minimized the cost for sequencing genomes. The real bottleneck to establish genomic resources is the annotation of genomic sequences. Long Terminal Repeat (LTR) retrotransposon is a major type of transposable genetic elements and dominating plant genomes. We developed a new method called LTR_retriever for accurate annotation of LTR retrotransposons. Further, we studied genome dynamics, genome size variation, and polyploidy origin using LTR retrotransposons. The presence of LTR retrotransposons challenges current sequencing and assembly techniques due to their size and repetitiveness. We proposed an unbiased metric called LTR Assembly Index (LAI) which utilizes the assembled LTR retrotransposons to evaluate continuity of genome assembly. We revealed the massive gain of continuity for assembly sequenced based on long-read techniques over short-read methods, and further proposed a standardized classification system for genome quality based on LAI. With high-quality genomes, we can extend our knowledge about microevolution events using a population of genomes. The domestication history of rice is still unresolved due to its complicated demographic history. We collected, re-mapped, and re-analyzed 3,485 cultivated and wild rice resequencing accessions. With data imputation, a total of 17.7 million high-quality single-nucleotide polymorphisms (SNPs) were identified. Our dataset is highly accurate as verified by cross-platform Affymetrix Microarray data, with a pairwise concordance rate of 99%. Combining phylogeny, PCA, and ADMIXTURE analyses, we present profound diversification among rice ecotypes.
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Title: Understanding the role of standing genetic variation in functional genetics and compensatory evolution
Creator: Chari, Sudarshan R.
Date: 2014
Collection: Electronic Theses & Dissertations
Description: Conventionally the phenotypic outcome of a mutation is considered to be due to a specific DNA lesion. But it has long been known that mutational effects can be conditional on environment (GxE) and genetic background (GxG). Thus it is standard practice to perform experiments by controlling for rearing environment and using co-isogenic strains. Though such a controlled approach has been very successful in enabling many discoveries, by not considering conditional effects our understanding of...
Show moreConventionally the phenotypic outcome of a mutation is considered to be due to a specific DNA lesion. But it has long been known that mutational effects can be conditional on environment (GxE) and genetic background (GxG). Thus it is standard practice to perform experiments by controlling for rearing environment and using co-isogenic strains. Though such a controlled approach has been very successful in enabling many discoveries, by not considering conditional effects our understanding of biological systems is incomplete. My research utilized conditionality in terms of genetic background and standing genetic variation therein to understand whether mutational interactions can themselves be background dependent. I demonstrated that a majority of mutational interactions identified via a dominant modifier screen are background dependent. Extending this idea of contingency in terms of standing genetic variation to the phenomenon of compensatory evolution in the presence of deleterious mutations, I demonstrated that natural populations of Drosophila melanogaster possess standing genetic variation for compensatory alleles to ameliorate even severe phenotypic defects. I further demonstrated that, despite considerable standing variation to ameliorate the focal phenotype perturbed by the mutation, natural selection exploits alternative evolutionary trajectories to recover fitness. Additionally this model system also allowed me to understand that loss of sexual signaling can be compensated by modulating behavioural and life history traits.
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Title: Balancing convergence and diversity in evolutionary single, multi and many objectives
Creator: Seada, Haitham
Date: 2017
Collection: Electronic Theses & Dissertations
Description: "Single objective optimization targets only one solution, that is usually the global optimum. On the other hand, the goal of multiobjective optimization is to represent the whole set of trade-off Pareto-optimal solutions to a problem. For over thirty years, researchers have been developing Evolutionary Multiobjective Optimization (EMO) algorithms for solving multiobjective optimization problems. Unfortunately, each of these algorithms were found to work well on a specific range of objective...
Show more"Single objective optimization targets only one solution, that is usually the global optimum. On the other hand, the goal of multiobjective optimization is to represent the whole set of trade-off Pareto-optimal solutions to a problem. For over thirty years, researchers have been developing Evolutionary Multiobjective Optimization (EMO) algorithms for solving multiobjective optimization problems. Unfortunately, each of these algorithms were found to work well on a specific range of objective dimensionality, i.e. number of objectives. Most researchers overlooked the idea of creating a cross-dimensional algorithm that can adapt its operation from one level of objective dimensionality to the other. One important aspect of creating such algorithm is achieving a careful balance between convergence and diversity. Researchers proposed several techniques aiming at dividing computational resources uniformly between these two goals. However, in many situations, only either of them is difficult to attain. Also for a new problem, it is difficult to tell beforehand if it will be challenging in terms of convergence, diversity or both. In this study, we propose several extensions to a state-of-the-art evolutionary many-objective optimization algorithm - NSGA-III. Our extensions collectively aim at (i) creating a unified optimization algorithm that dynamically adapts itself to single, multi- and many objectives, and (ii) enabling this algorithm to automatically focus on either convergence, diversity or both, according to the problem being considered. Our approach augments the already existing algorithm with a niching-based selection operator. It also utilizes the recently proposed Karush Kuhn Tucker Proximity Measure to identify ill-converged solutions, and finally, uses several combinations of point-to-point single objective local search procedures to remedy these solutions and enhance both convergence and diversity. Our extensions are shown to produce better results than state-of-the-art algorithms over a set of single, multi- and many-objective problems."--Pages ii-iii.
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Title: PARTNERSHIP AND PREDATION IN PLANT-BIOTIC INTERACTIONS : THEORY, METHODS, AND EVIDENCE
Creator: Rowe, Shawna L.
Date: 2022
Collection: Electronic Theses & Dissertations
Description: As sessile organisms, plants face an endless series of encounters with other living organisms in their environment. Ranging from the beneficial to the harmful, the pressures of these biotic interactions force plants to rapidly adapt to survive and thrive. This dissertation is aimed at addressing questions about the metabolism of plant interactions with herbivores and nutritional symbionts in legumes. Despite spanning different systems and methods, these questions reflect my interest in...
Show moreAs sessile organisms, plants face an endless series of encounters with other living organisms in their environment. Ranging from the beneficial to the harmful, the pressures of these biotic interactions force plants to rapidly adapt to survive and thrive. This dissertation is aimed at addressing questions about the metabolism of plant interactions with herbivores and nutritional symbionts in legumes. Despite spanning different systems and methods, these questions reflect my interest in understanding the biochemistry underlying ecological and evolutionary function and my desire to develop tools that enable the investigation of such questions. To this end, the work in this dissertation is aimed at building conceptual and methodological tools to enable more thorough investigations of plant symbioses and, more broadly, plant-biotic interactions across levels of biological organization.First, I present a framework for making predictions on evolutionary trajectories and origins of plant--microbe communication systems. By highlighting the prevalence of coercive interactions in plant--microbe interactions, I demonstrate the plausibility of such interactions types to be an evolutionary precursor to seemingly stable signaling mechanisms. This work aims to provide useful evolutionary context for investigations concerning the evolutionary stability and exploitation of signaling mechanisms in established biological relationships. Second, I present collaborative work in which we developed and applied a cost-effective, high-throughput protocol for quantifying multiple biochemical defense responses from small quantities of plant tissue using spectrophotometric techniques. This protocol was then applied on two distinct populations of the legume Medicago polymorpha to investigate how changes defensive traits in responses to altered selective pressures have manifested over the course of novel range expansion. Our work demonstrated the feasibility and potential of assessing defense responses across plant populations. Further, the work documents a shift in herbivore preference of plants from familiar and unfamiliar ranges by demonstrating a herbivore preference for plant tissues with pre-induced defenses over uninduced tissues from an unfamiliar geographic range. Finally, I demonstrate the creation, construction, and validity of a novel microcosm system for assessing nutrient exchange in the symbiotic mutualism between plants and arbuscular mycorrhizal fungi (AMF). The novel system is reliable, biologically-relevant, durable, and sufficiently simple and cost-effective to deploy. I demonstrate the validity of the microcosm system and discuss in-progress work which demonstrates its potential to rigorously investigate unknown aspects of the plant-AMF mutualism. Taken together, these developments and suggestions contribute to the growing set of methods and frameworks developed for improving our understanding a various plant-biotic interactions.
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Title: Subgenome dominance and genome evolution in allopolyploids
Creator: Bird, Kevin Andrew
Date: 2022
Collection: Electronic Theses & Dissertations
Description: The merger of divergent genomes, via hybridization or allopolyploidization, frequently results in a ‘genomic shock’ that induces a series of rapid genetic and epigenetic modifications as a result of conflicts between parental genomes. This conflict among the subgenomes routinely leads one subgenome to become dominant over the other subgenome(s), resulting in subgenome biases in gene content and expression. Recent advances in methods to analyze hybrid and polyploid genomes with comparisons to...
Show moreThe merger of divergent genomes, via hybridization or allopolyploidization, frequently results in a ‘genomic shock’ that induces a series of rapid genetic and epigenetic modifications as a result of conflicts between parental genomes. This conflict among the subgenomes routinely leads one subgenome to become dominant over the other subgenome(s), resulting in subgenome biases in gene content and expression. Recent advances in methods to analyze hybrid and polyploid genomes with comparisons to extant parental progenitors have allowed for major strides in understanding the mechanistic basis for subgenome dominance. In particular, our understanding of the role that homoeologous exchange might play in subgenome dominance and genome evolution is quickly growing. Here I present novel work in several polyploid species investigating the biological and evolution impact of polyploidy and the evolution of these polyploid species. The first chapter introduces concepts like whole-genome duplication and describes advances in genomic sequencing technology that have accelerated the study of polyploid genomes. The second chapter reviews subgenome dominance and recent breakthroughs in understanding its causes and implications for genome evolution. The third chapter explores the repeatability of subgenome dominance in independently resynthesized Brassica napus. The fourth chapter investigates the extent to which genomic rearrangements from chromosomal duplications and deletions and homoeologous exchange can bias the analysis of subgenome expression dominance from RNAseq data. The fifth chapter explores the prevalence and impact of homoeologous exchange on independently resynthesized Brassica napus, providing novel evidence that gene dosage changes from homoeologous exchange are constrained by the need to maintain dosage balance of gene products. The sixth chapter explores the origins and admixture of wild octoploid strawberries Fragaria virgniana and Fragaria chiloensis with newly generated genomic resources applied to global collections.
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Title: The Evolutionary Origins of Cognition : Understanding the early evolution of biological control systems and general intelligence
Creator: Carvalho Pontes, Anselmo
Date: 2021
Collection: Electronic Theses & Dissertations
Description: In the last century, we have made great strides towards understanding natural cognition and recreating it artificially. However, most cognitive research is still guided by an inadequate theoretical framework that equates cognition to a computer system executing a data processing task. Cognition, whether natural or artificial, is not a data processing system; it is a control system.At cognition's core is a value system that allows it to evaluate current conditions and decide among two or more...
Show moreIn the last century, we have made great strides towards understanding natural cognition and recreating it artificially. However, most cognitive research is still guided by an inadequate theoretical framework that equates cognition to a computer system executing a data processing task. Cognition, whether natural or artificial, is not a data processing system; it is a control system.At cognition's core is a value system that allows it to evaluate current conditions and decide among two or more courses of action. Memory, learning, planning, and deliberation, rather than being essential cognitive abilities, are features that evolved over time to support the primary task of deciding “what to do next”. I used digital evolution to recreate the early stages in the evolution of natural cognition, including the ability to learn. Interestingly, I found cognition evolves in a predictable manner, with more complex abilities evolving in stages, by building upon previous simpler ones. I initially investigated the evolution of dynamic foraging behaviors among the first animals known to have a central nervous system, Ediacaran microbial mat miners. I then followed this up by evolving more complex forms of learning. I soon encountered practical limitations of the current methods, including exponential demand of computational resources and genetic representations that were not conducive to further scaling. This type of complexity barrier has been a recurrent issue in digital evolution. Nature, however, is not limited in the same ways; through evolution, it has created a language to express robust, modular, and flexible control systems of arbitrary complexity and apparently open-ended evolvability. The essential features of this language can be captured in a digital evolution platform. As an early demonstration of this, I evolved biologically plausible regulatory systems for virtual cyanobacteria. These systems regulate the cells' growth, photosynthesis and replication given the daily light cycle, the cell's energy reserves, and levels of stress. Although simple, this experimental system displays dynamics and decision-making mechanisms akin to biology, with promising potential for open-ended evolution of cognition towards general intelligence.
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Title: Contextual influences on undergraduate biology students' reasoning and representations of evolutionary concepts
Creator: de Lima, Joelyn
Date: 2021
Collection: Electronic Theses & Dissertations
Description: Context is the background or the settings of an event or idea. It is only when events or ideas are considered within the context in which they occur that they can be fully understood. In education, the application of knowledge communicated in one context to a different one is a central feature of learning. However, knowledge transfer can be affected by multiple factors including contexts used. Context plays a vital role in both shaping students’ learning and in eliciting their knowledge....
Show moreContext is the background or the settings of an event or idea. It is only when events or ideas are considered within the context in which they occur that they can be fully understood. In education, the application of knowledge communicated in one context to a different one is a central feature of learning. However, knowledge transfer can be affected by multiple factors including contexts used. Context plays a vital role in both shaping students’ learning and in eliciting their knowledge. Therefore, understanding how context can help or hinder learning and how context impacts knowledge assessment is important for improving science learning outcomes.For my dissertation, I studied contextual influences on the ways students reason and represent their knowledge. My studies explored two types of contexts: surface features of prompts provided to students (e.g., organism used) and the mode of response requested (e.g., written narratives vs constructed models). I analysed the effect of prompt surface features on the content of students’ written responses and on the architecture of models they constructed to explain evolution by natural selection. I also analysed the effect of mode on the content and level of scientific plausibility of students’ responses. In addition, I explored the association between instruction and prior achievement and susceptibility to contextual influences.My results indicate that prompt contextual features and mode of response are eliciting differences in the content of students’ representations. Contextual susceptibility decreased with instruction and higher prior academic achievement. This could indicate that they are novice learners and have a fragile understanding of either the subject matter (evolution), the alternative representation that was required (constructing models), or of both the subject matter and the representation. Incorporating multiple contexts and modes of assessment has potential to generate a more holistic view of students’ understanding and may promote greater transfer by requiring students to think and reason across contexts.
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Title: Social Modulation of Individual Decision-Making in Foraging Bumblebees : Mechanisms and Evolution
Creator: Incorvaia, Darren
Date: 2021
Collection: Electronic Theses & Dissertations
Description: How and why animals choose to do what they do at any given moment is one of the fundamental questions in animal behavior. For social animals, influences on decision-making can come from both personal and social sources, and in eusocial insects like ants, bees, and wasps, the reliance on social information is taken to the extreme. Foraging bumblebees offer the perfect model in which to examine the social influences on individual decision-making because they are presented with extensive...
Show moreHow and why animals choose to do what they do at any given moment is one of the fundamental questions in animal behavior. For social animals, influences on decision-making can come from both personal and social sources, and in eusocial insects like ants, bees, and wasps, the reliance on social information is taken to the extreme. Foraging bumblebees offer the perfect model in which to examine the social influences on individual decision-making because they are presented with extensive personal and social information, and when foraging they are solely focused on the task at hand. Chapter 1 reviews information use by foraging bumblebees, setting the stage for the subsequent data chapters. Chapter 2 examines how the motivation for bumblebees to feed from a known feeder is modified by the nutritive state of the colony, such that individuals in colonies with full food stores show lower motivation to feed. In addition to this behavioral result, a biochemical analysis reveals that lipid levels may be involved in the mechanism underlying this social effect. Eusocial insects are famous for collective behaviors, such as the swarming behavior of honeybees, the foraging trails of termites, and the bridge-building of ants. While the collective foraging strategy of other eusocial insects has been well-studied, it has not received attention in bumblebees. In Chapter 3 I use a behavioral experiment to reveal that bumblebees use a strategy of informed individual initiative to collectively ensure they are foraging from the best resources in the environment. In this strategy, individual bees adjust their reward expectations based on the quality of nectar stored in the nest. I followed up this experiment with a computational model to reveal that this strategy is adaptive, as it results in higher fitness than does individual search alone. This strategy is markedly different from the spatial communication of the dance language used by honeybees, who are close relatives of bumblebees. This prompted me to extend the computational model to examine the selective pressures that shape foraging strategies in social insects, including the honeybee dance language and bumblebee strategy of informed individual initiative. In Chapter 4, I present the results of simulations of this extended model, demonstrating that, although resource density influences fitness for both the dance language and informed individual initiative, colony size only matters for the dance language. This suggests that the large colony sizes of honeybees may have been important for the dance language to evolve, whereas a similar spatial communication system would not be adaptive in bumblebees, which have smaller colony sizes. Taken all together, the results in this dissertation explore how individual decision-making is shaped by the social environment in bumblebees, and the potential selective pressures that led to these behavioral strategies over evolutionary time. Bumblebees are important pollinators in both agricultural and natural ecosystems, but many species are facing declines; a more thorough understanding of their behavior is imperative to help us conserve them as the planet continues to change due to climate change and other anthropogenic influences.
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Title: Digital Evolution in Experimental Phylogenetics and Evolution Education
Creator: Kohn, Cory
Date: 2021
Collection: Electronic Theses & Dissertations
Description: The creation and evaluation of known evolutionary histories and the implementation of student investigatory experiences on evolution are difficult endeavors that have only recently been feasible. The research presented in this dissertation is related in their shared use of digital evolution with Avidians as a model study system, both to conduct science research in experimental phylogenetics and to conduct education research in curricular intervention to aid student understanding.I first...
Show moreThe creation and evaluation of known evolutionary histories and the implementation of student investigatory experiences on evolution are difficult endeavors that have only recently been feasible. The research presented in this dissertation is related in their shared use of digital evolution with Avidians as a model study system, both to conduct science research in experimental phylogenetics and to conduct education research in curricular intervention to aid student understanding.I first present background discussions on the Avidian digital evolution study system—as implemented in Avida and Avida-ED—and its favorable use in experimental phylogenetics and biology education owing to its greater biological realism than computational simulations, and greater utility and generality than biological systems. Prior work on conducting experimental evolution for use in phylogenetics and work on developing undergraduate lab curricula using experimental evolution are also reviewed. I establish digital evolution as an effective method for phylogenetic inference validation by demonstrating that results from a known Avidian evolutionary history are concordant, under similar conditions, to established biological experimental phylogenetics work. I then further demonstrate the greater utility and generality of digital evolution over biological systems by experimentally testing how phylogenetic accuracy may be reduced by complex evolutionary processes operating singly or in combination, including absolute and relative degrees of evolutionary change between lineages (i.e., inferred branch lengths), recombination, and natural selection. These results include that directional selection aids phylogenetic inference, while stabilizing selection impedes it. By evaluating clade accuracy and clade resolvability across treatments, I evaluate measures of tree support and its presentation in the form of consensus topologies and I offer several general recommendations for systematists. Using a larger and more biologically realistic experimental design, I systematically examine a few of the complex processes that are hypothesized to affect phylogenetic accuracy—natural selection, recombination, and deviations from the model of evolution. By analyzing the substitutions that occurred and calculating selection coefficients for derived alleles throughout their evolutionary trajectories to fixation, I show that molecular evolution in these experiments is complex and proceeding largely as would be expected for biological populations. Using these data to construct empirical substitution models, I demonstrate that phylogenetic inference is incredibly robust to significant molecular evolution model deviations. I show that neutral evolution in the presence of always-occurring population processes, such as clonal or Hill-Robertson interference and lineage sorting, result in reduced clade support, and that selection and especially recombination, including their joint occurrence, restore this otherwise-reduced phylogenetic accuracy. Finally, this work demonstrates that inferred branch lengths are often quite inaccurate despite clade support being accurate. While phylogenetic inference methods performed relatively well in both theoretically facile and challenging molecular evolution scenarios, their accuracy in clade support might be a remarkable case of being right for misguided reasons, since branch length inference were largely inaccurate, and drastically different models of evolution made little difference. This work highlights the need for further research that evaluates phylogenetic methods under experimental conditions and suggests that digital evolution has a role here. Finally, I examine student understanding of the importance of biological variation in the context of a course featuring a digital evolution lab. I first describe the Avida-ED lab curriculum and its fulfillment of calls for reform in education. Then I describe the specific education context and other course features that aim to address student conceptualization of variation. I present a modified published assessment on transformational and variational understanding and findings regarding student understanding of variation within an evolution education progression. Finally, I offer suggestions on incorporating course material to engage student understanding of variation.
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Title: Developing Reverse Genetic Tools in Weakly Electric Fish : Investigating Electric Organ in vivo scn4aa Function Through CRISPR Knockouts and Morpholino Knockdowns
Creator: Constantinou, Savvas James
Date: 2021
Collection: Electronic Theses & Dissertations
Description: The ability to determine gene function allows research to progress at one of the finest scales in biology and is a goal in electric fish research. Reverse genetics allows researchers to determine gene function and would aid the electric fish community in beginning to answer some of the broadest and most complicated questions in biology such as linking genotype to phenotype and understanding the processes that lead to biological diversity. In this dissertation, I describe the development of...
Show moreThe ability to determine gene function allows research to progress at one of the finest scales in biology and is a goal in electric fish research. Reverse genetics allows researchers to determine gene function and would aid the electric fish community in beginning to answer some of the broadest and most complicated questions in biology such as linking genotype to phenotype and understanding the processes that lead to biological diversity. In this dissertation, I describe the development of two major reverse genetic tools for use in the electric fish system: CRISPR/Cas9 genome editing and morpholinos. To develop these tools, I also produced protocols for in-vitro breeding, husbandry, and single-cell embryo microinjections. In the first chapter, I describe in-vitro breeding, husbandry, and single-cell embryo microinjections and demonstrate that CRISPR/Cas9 is a promising tool for future electric fish research by targeting nonsense mutations to scn4aa in the mormyrid Brienomyrus brachyistius and gymnotiform Brachyhypopomus gauderio, two independently evolved lineages of weakly electric fish, resulting in a reduction in the electric organ discharge amplitude. In the second chapter, I provide electric fish researchers with a detailed analysis of our many successes and failures applying CRISPR/Cas9 methods to this system and discuss future suggestions on how to best apply them to novel electric fish research. In the third chapter, I describe my efforts to utilize vivo-morpholinos in mormyrid electric fish. While a single early pilot study I performed demonstrated vivo-morpholinos can reduce target gene mRNA levels and cause a phenotypic effect, my efforts to replicate these findings demonstrate inconsistent performance: control vivo-morpholino and scn4aa targeting vivo-morpholino injected fish had indistinguishable effects on electric organ discharge amplitude. Due to additional concerns of toxicity, I suggest morpholinos are not an ideal reverse genetic tool in Brienomyrus brachyistius and should only be utilized for future research with caution.
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Title: Diversity of South American ambrosia beetles (Curculionidae : Scolytinae: Xyleborini) and their fungal partners
Creator: Osborn, Rachel Kathryn
Date: 2022
Collection: Electronic Theses & Dissertations
Description: Ambrosia beetles from the tribe Xyleborini (Coleoptera: Curculionidae: Scolytinae) small, haplodiploid beetles that farm nutritional fungi on the walls of tunnels they excavate in the xylem of dead or nearly dead trees. These biological traits make them successful participants in worldwide wooded ecosystems and facilitate their human-mediated invasion beyond their native ranges. A minority of these introduced species are classified as pests because of the physical damage they cause to their...
Show moreAmbrosia beetles from the tribe Xyleborini (Coleoptera: Curculionidae: Scolytinae) small, haplodiploid beetles that farm nutritional fungi on the walls of tunnels they excavate in the xylem of dead or nearly dead trees. These biological traits make them successful participants in worldwide wooded ecosystems and facilitate their human-mediated invasion beyond their native ranges. A minority of these introduced species are classified as pests because of the physical damage they cause to their plant hosts, or because they vector pathogenic fungi that infect ornamental, lumber, and forest trees. Most of the current knowledge on the diversity of xyleborine beetles and their fungi centers around species found in North America, Asia, and Europe. Little is known about the ambrosia partnerships in the Neotropics, which is concerning because South America is a strong trading partner with the US and the potential for new invasive Xyleborini to be imported from this area is significant. Continuing forest damage caused by invasive Xyleborini/fungi inspires robust research efforts to describe these symbionts and document their biological traits. Considerable efforts are required to enhance such endeavors in underrepresented regions such as South America and Africa. To increase understanding of the South American Xyleborini and their associated fungi, I compiled current knowledge of their historical and contemporary taxonomic records, biological records, and ecological studies. I also completed surveys throughout Ecuador to collect beetles and fungi. Molecular analysis of fungi isolated from Ecuadorian beetles reveals that several Coptoborus species associate with Fusarium fungi, including the ambrosia Fusarium Clade (AFC) that has previously been recovered from Euwallacea spp. and Xyleborus ferrugineus in Central America, Florida, California, Israel, and Asia. Examination of the morphology of some South American xyleborine specimens previously classified as Coptoborus spp. suggests a high similarity to Xyleborus spp. from Africa. Phylogenetic analysis of these South American and African beetles as well as morphological assessment of additional specimens necessitates the designation of a new genus Xenoxylebora gen. nov. containing species endemic to both continents. This unusual distribution demonstrates the ability of these ambrosia beetles to survive long-distance trans-oceanic dispersal.
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Title: MECHANISMS UNDERLYING DESICCATION RESISTANCE IN DROSOPHILA SPECIES
Creator: Wang, Zinan
Date: 2021
Collection: Electronic Theses & Dissertations
Description: Adaptation to various and extreme environments is key to long-term species persistence. Reducing water loss is important for organisms adapting to different terrestrial environments. In Drosophila fruit flies and other terrestrial insects, their small body size and large surface areas to volume ratios make them vulnerable to desiccation stress. Their ability to prevent water loss is crucial for their survival. Previous studies have suggested that cuticular water loss accounts for the majority...
Show moreAdaptation to various and extreme environments is key to long-term species persistence. Reducing water loss is important for organisms adapting to different terrestrial environments. In Drosophila fruit flies and other terrestrial insects, their small body size and large surface areas to volume ratios make them vulnerable to desiccation stress. Their ability to prevent water loss is crucial for their survival. Previous studies have suggested that cuticular water loss accounts for the majority of water loss in insects and hypothesized that differences in cuticular hydrocarbon (CHC) content accounted for differences in desiccation resistance between mesic and desert species. However, the specific association between different CHC components and desiccation has not been established, and the genetic mechanisms underlying the evolution of these CHC components that confer high desiccation resistance have not been elucidated. This dissertation investigated how the evolution of CHCs in insects affected desiccation resistance and elucidates the genetic mechanisms underlying their evolution. With a comprehensive association study of desiccation resistance and CHCs in 46 Drosophila species and 4 species in closely-related genera, the analyses showed that mbCHC chain lengths were important predictors of desiccation resistance and longer mbCHCs contributed to higher desiccation resistance. This dissertation further investigated the genetic and molecular mechanisms underlying longer chain mbCHCs and higher desiccation resistance in a desert Drosophila species, Drosophila mojavensis. A fatty acyl-CoA elongase gene, mElo (methyl-branched CHC Elongase), was identified in Drosophila species for the elongation of mbCHCs. Overexpression experiments in D. melanogaster demonstrated that coding changes in mElo from D. mojavensis lead to longer mbCHCs and higher desiccation resistance. Further experiments using CRISPR-Cas9 to knock out mElo from D. mojavensis showed that knockout of this gene decreased the production of the longest mbCHCs and significantly reduced desiccation resistance at their ecological-relevant temperature. Results from this dissertation elucidate the molecular and evolutionary mechanisms that enable species to reduce water loss and maintain water balance as our planet gets warmer and more arid in the next few decades.
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Title: Adaptation to agriculture in a serious crop weed, weedy radish (raphanus raphanistrum)
Creator: Garrison, Ava
Date: 2022
Collection: Electronic Theses & Dissertations
Description: The colonization of novel environments requires organisms to shift their trait means in response to differing abiotic and biotic conditions in order to survive and persist. This response can be done via phenotypic plasticity (a trait shift in response to the environment), adaptation (a trait shift due to genetic change), or both strategies can be used together, with plasticity “buying time” for adaptation to occur. The colonization of novel environments is especially important to the...
Show moreThe colonization of novel environments requires organisms to shift their trait means in response to differing abiotic and biotic conditions in order to survive and persist. This response can be done via phenotypic plasticity (a trait shift in response to the environment), adaptation (a trait shift due to genetic change), or both strategies can be used together, with plasticity “buying time” for adaptation to occur. The colonization of novel environments is especially important to the establishment of agricultural weeds worldwide, which thrive in these extreme environments of intense competition and frequent disturbance. In this dissertation, I address the establishment and evolution of a harmful agricultural weed, weedy radish (Raphanus raphanistrum), as well as its divergence from a wild relative of the same species, the native radish ecotype. I first investigated the hypothesis of phenotypic plasticity “buying time” for adaptation to agricultural fields in weedy radish. Using growth chambers to simulate the ancestral (native) and derived (weedy) environments of weedy radish, I performed a reciprocal transplant with the weedy and native radish ecotypes. I found phenotypic plasticity between environments and genetic divergence between ecotypes to be equally common among traits, suggesting similar importance of plasticity and adaptation in weedy radish establishment. Further, in the majority of traits that were both plastic and differentiated between ecotypes, the direction of change matched, with the weedy environment producing phenotypic shifts in the direction of the weedy ecotype mean. This suggests plasticity in these traits may have enabled the subsequent adaptation and ecotype differentiation, supporting the buying-time hypothesis. Next, I explored the role of the plant hormone Gibberellic Acid (GA) in the evolution of weedy radish. Using exogenous application of GA both in the greenhouse and in weedy and native growth chamber environments, I found evidence that there has been an evolutionary change in the role of GA in trait expression between the two ecotypes. Namely, weedy radish is less responsive to GA application than native radish, suggesting either upregulation in GA production in weeds, or a lower level of GA required to enable gene expression in the weedy ecotype. This change in gene regulation by GA may have been important in the evolution of weedy radish in the agricultural field. Finally, I assessed the likelihood of weedy radish diverging from a native ancestor via adaptive evolution. I found that adaptive evolution was likely in the establishment of weedy radish due to increased fitness of the weedy ecotype compared to the native ecotype in the agricultural field. I also found traits under directional selection in the native ecotype, with the key takeaway that faster flowering is adaptive in the agricultural fields. I finally looked at the ability of weedy radish to evolve advanced flowering in the agricultural field via standing genetic variance by artificially selecting for early flowering in native radish. I found that in only two generations of selection, native populations significantly advanced their flowering time, supporting the notion of weedy radish rapidly adapting to agricultural conditions via standing genetic variation alone. Taken together, these findings work to piece together the evolutionary history of weedy radish, providing insight into its mechanisms of establishment. This work also contributes to our overall understanding of rapid evolution and phenotypic plasticity in the colonization of novel environments, in agricultural weeds and beyond.
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Title: MODELING AND PREDICTION OF GENETIC REDUNDANCY IN ARABIDOPSIS THALIANA AND SACCHAROMYCES CEREVISIAE
Creator: Cusack, Siobhan Anne
Date: 2020
Collection: Electronic Theses & Dissertations
Description: Genetic redundancy is a phenomenon where more than one gene encodes products that perform the same function. This frequently manifests experimentally as a single gene knockout mutant which does not demonstrate a phenotypic change compared to the wild type due to the presence of a paralogous gene performing the same function; a phenotype is only observed when one or more paralogs are knocked out in combination. This presents a challenge in a fundamental goal of genetics, linking genotypes to...
Show moreGenetic redundancy is a phenomenon where more than one gene encodes products that perform the same function. This frequently manifests experimentally as a single gene knockout mutant which does not demonstrate a phenotypic change compared to the wild type due to the presence of a paralogous gene performing the same function; a phenotype is only observed when one or more paralogs are knocked out in combination. This presents a challenge in a fundamental goal of genetics, linking genotypes to phenotypes, especially because it is difficult to determine a priori which gene pairs are redundant. Furthermore, while some factors that are associated with redundant genes have been identified, little is known about factors contributing to long-term maintenance of genetic redundancy. Here, we applied a machine learning approach to predict redundancy among benchmark redundant and nonredundant gene pairs in the model plant Arabidopsis thaliana. Predictions were validated using well-characterized redundant and nonredundant gene pairs. Additionally, we leveraged the availability of fitness and multi-omics data in the budding yeast Saccharomyces cerevisiae to build machine learning models for predicting genetic redundancy and related phenotypic outcomes (single and double mutant fitness) among paralogs, and to identify features important in generating these predictions. Collectively, our models of genetic redundancy provide quantitative assessments of how well existing data allow predictions of fitness and genetic redundancy, shed light on characteristics that may contribute to long-term maintenance of paralogs that are seemingly functionally redundant, and will ultimately allow for more targeted generation of phenotypically informative mutants, advancing functional genomic studies.
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Title: Evolving Phenotypically Plastic Digital Organisms
Creator: Lalejini, Alexander
Date: 2021
Collection: Electronic Theses & Dissertations
Description: The ability to dynamically respond to cues from the environment is a fundamental feature of most adaptive systems. In biological systems, changes to an organism based on environmental cues is called phenotypic plasticity. Indeed, phenotypic plasticity underlies many of the adaptive traits and developmental patterns found in nature and serves as a key mechanism for responding to spatially or temporally variable environments. Most computer programs require phenotypic plasticity, as they must...
Show moreThe ability to dynamically respond to cues from the environment is a fundamental feature of most adaptive systems. In biological systems, changes to an organism based on environmental cues is called phenotypic plasticity. Indeed, phenotypic plasticity underlies many of the adaptive traits and developmental patterns found in nature and serves as a key mechanism for responding to spatially or temporally variable environments. Most computer programs require phenotypic plasticity, as they must respond dynamically to stimuli such as user input, sensor data, et cetera. As such, phenotypic plasticity also has practical applications in genetic programming, wherein we apply the natural principles of evolution to automatically synthesize computer programs rather than writing them by hand. In this dissertation, I achieve two synergistic aims: (1) I use populations of self-replicating computer programs (digital organisms) to empirically study the conditions under which adaptive phenotypic plasticity evolves and how its evolution shapes subsequent evolutionary outcomes; and (2) I transfer insights from biology to develop novel genetic programming techniques in order to evolve more responsive (i.e., phenotypically plastic) computer programs. First, I illustrate the importance of mutation rate, environmental change, and partially-plastic building blocks for the evolution of adaptive plasticity. Next, I show that adaptive phenotypic plasticity stabilizes populations against environmental change, allowing them to more easily retain novel adaptive traits. Finally, I improve our ability to evolve phenotypically plastic computer programs with three novel genetic programming techniques: (1) SignalGP, which provides mechanisms to control code expression based on environmental cues, (2) tag-based genetic regulation to adjust code expression based on current context, and (3) tag-accessed memory to provide more dynamic mechanisms for storing data.
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Title: Replaying Life's Virtual Tape : Examining the Role of History in Experiments with Digital Organisms
Creator: Bundy, Jason Nyerere
Date: 2021
Collection: Electronic Theses & Dissertations
Description: Evolution is a complex process with a simple recipe. Evolutionary change involves three essential “ingredients” interacting over many generations: adaptation (selection), chance (random variation), and history (inheritance). In 1989’s Wonderful Life, the late paleontologist Stephen Jay Gould advocated for the importance of historical contingency—the way unique events throughout history influence future possibilities—using a clever thought experiment of “replaying life’s tape”. But not...
Show moreEvolution is a complex process with a simple recipe. Evolutionary change involves three essential “ingredients” interacting over many generations: adaptation (selection), chance (random variation), and history (inheritance). In 1989’s Wonderful Life, the late paleontologist Stephen Jay Gould advocated for the importance of historical contingency—the way unique events throughout history influence future possibilities—using a clever thought experiment of “replaying life’s tape”. But not everyone was convinced. Some believed that chance was the primary driver of evolutionary change, while others insisted that natural selection was the most powerful influence. Since then, “replaying life’s tape” has become a core method in experimental evolution for measuring the relative contributions of adaptation, chance, and history. In this dissertation, I focus on the effects associated with history in evolving populations of digital organisms—computer programs that self-replicate, mutate, compete, and evolve in virtual environments. In Chapter 1, I discuss the philosophical significance of Gould’s thought experiment and its influence on experimental methods. I argue that his thought experiment was a challenge to anthropocentric reasoning about natural history that is still popular, particularly outside of the scientific community. In this regard, it was his way of advocating for a “radical” view of evolution. In Chapter 2—Richard Lenski, Charles Ofria, and I describe a two-phase, virtual, “long-term” evolution experiment with digital organisms using the Avida software. In Phase I, we evolved 10 replicate populations, in parallel, from a single genotype for around 65,000 generations. This part of the experiment is similar to the design of Lenski’s E. coli Long-term Evolution Experiment (LTEE). We isolated the dominant genotype from each population around 3,000 generations (shallow history) into Phase I and then again at the end of Phase I (deep history). In Phase II, we evolved 10 populations from each of the genotypes we isolated from Phase I in two new environments, one similar and one dissimilar to the old environment used for Phase I. Following Phase II, we estimated the contributions of adaptation, chance, and history to the evolution of fitness and genome length in each new environment. This unique experimental design allowed us to see how the contributions of adaptation, chance, and history changed as we extended the depth of history from Phase I. We were also able to determine whether the results depended on the extent of environmental change (similar or dissimilar new environment). In Chapter 3, we report an extended analysis of the experiment from the previous chapter to further examine how extensive adaptation to the Phase I environment shaped the evolution of replicates during Phase II. We show how the form of pleiotropy (antagonistic or synergistic) between the old (Phase I) and new (Phase II) habitats was influenced by the depth of history from Phase I (shallow or deep) and the extent of environmental change (similar or dissimilar new environment). In the final chapter Zachary Blount, Richard Lenski, and I describe an exercise we developed using the educational version of Avida (Avida-ED). The exercise features a two-phase, “replaying life’s tape” activity. Students are able to explore how the unique history of founders that we pre-evolved during Phase I influences the acquisition of new functions by descendent populations during Phase II, which the students perform during the activity.
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Title: Sex and Individual Differences in Agonistic Behavior of Spotted Hyenas (Crocuta Crocuta) : Effects on Fitness and Dominance
Creator: McCormick, S. Kevin
Date: 2022
Collection: Electronic Theses & Dissertations
Description: Agonistic behavior can be observed across all taxa arising from a common need to compete over limiting resources. Within species, individual variation of agonistic behavior can allow individuals to acquire and maintain limiting resources leading to higher reproductive success or fitness. However, what is often overlooked in studies of agonistic behavior is submissiveness, and how this aspect of agonistic behavior relates to aggressiveness. Further, historical studies of agonistic behavior...
Show moreAgonistic behavior can be observed across all taxa arising from a common need to compete over limiting resources. Within species, individual variation of agonistic behavior can allow individuals to acquire and maintain limiting resources leading to higher reproductive success or fitness. However, what is often overlooked in studies of agonistic behavior is submissiveness, and how this aspect of agonistic behavior relates to aggressiveness. Further, historical studies of agonistic behavior among social mammals are biased towards studies of male agonistic behavior, often ignoring aspects and effects of female agonistic behavior. Here, I address these knowledge gaps through a long-term study of a free living highly gregarious mammal, the spotted hyena (Crocuta crocuta). Spotted hyenas offer an excellent model system for studying variation in aggressive and submissive behavior within individuals and between sexes, as they live in complex societies formed around a female dominated, or matrilineal, hierarchy that is enforced through constant agonistic interactions. For this dissertation, I utilized 30 years’ worth of consistently recorded behavioral data collected by Dr. Kay E. Holekamp and her team from free living hyenas residing within the Masai Mara National Reserve, Keyna. Because this dissertation involved many collaborations with other scientists, I use “we” throughout this abstract to describe participation in each chapter. In Chapter 1, we describe sexually dimorphic traits within spotted hyenas that fit common mammalian patterns, as well as numerous traits that violate mammalian norms, including sex differences in agonistic behavior. In particular, adult female spotted hyenas are significantly more likely to emit unsolicited acts of aggression down the hierarchy than adult breeding males, and females do so significantly more ferociously, or intensely. For Chapter 2, we analyzed rates and intensities of unprovoked aggressive and submissive acts emitted by adult females to determine if these two behaviors were individually consistent, as well as testing the hypothesis that these two behaviors may represent separate traits within individuals. Here we found that the intensity at which females emit aggressive and submissive behaviors are consistent, and that these traits were not correlated within individuals. Further, both consistent aggressive intensity and submissive intensity were correlated to adult female fitness, such that individuals expressing high or low extremes of these behaviors had lower annual offspring survival. Then in Chapter 3, we assessed drivers of female dominance within spotted hyenas. Within this chapter we tested two hypotheses 1) that intrinsic sex differences in agonistic behavior drives female dominance and/or 2) social support facilitates female dominance in this species. Further, we assessed these hypotheses among juvenile age classes to determine if drivers of female dominance occurred prior to sexual maturity and subsequent male dispersal. We found that females are intrinsically more aggressive both as cubs and adults, and adult males more submissive whether provoked or not. Further, social support during agonistic encounters is more likely to occur when acting against a female than a male, and adult females can dominate males with or without support. In completion, my dissertation provides interesting insights to sexual and individual variation on agonistic behavior among a social mammal.
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Title: Evolution of laboratory and natural populations of Escherichia coli
Creator: Maddamsetti, Rohan
Date: 2016
Collection: Electronic Theses & Dissertations
Description: My dissertation spans two dichotomies: evolution in the laboratory versus evolution in nature, and asexual versus sexual evolutionary dynamics. In Chapter 1 I describe asexual evolutionary dynamics in one population of Lenski’s long-term evolution experiment with Escherichia coli. I describe cohorts of mutations that sweep to fixation together as characteristic of clonal interference dynamics. I also describe an ecological interaction that evolved and then went extinct after thousands of...
Show moreMy dissertation spans two dichotomies: evolution in the laboratory versus evolution in nature, and asexual versus sexual evolutionary dynamics. In Chapter 1 I describe asexual evolutionary dynamics in one population of Lenski’s long-term evolution experiment with Escherichia coli. I describe cohorts of mutations that sweep to fixation together as characteristic of clonal interference dynamics. I also describe an ecological interaction that evolved and then went extinct after thousands of generations, and discuss how such interactions affect cohorts of mutations. In Chapter 2 I report that conserved core genes tend to be targets of selection in the long-term experiment. In Chapter 3, I investigate the surprising observation that synonymous genetic diversity is not uniform across the genomes of natural E. coli isolates. This observation is surprising because in clonal organisms with a constant point mutation rate, synonymous diversity should be constant across the genome. I use patterns of synonymous mutations in the long-term experiment to argue that genome-wide variation in the mutation rate does not adequately explain patterns of synonymous genetic diversity. In Chapter 4, I propose that recombination and gene flow could account for genome-wide variation in synonymous genetic diversity. In Chapter 5, I analyze E. coli genomes isolated from an evolution experiment with recombination in which E. coli K-12 with known growth defects could donate genetic material to recipient populations founded by long-term experiment clones. The degree of recombination varied dramatically across sequenced clones. The strongest predictor of successful transfer was proximity to the oriT origin of transfer in the K-12 donors. Donor alleles close to oriT replaced their recipient counterparts at a high rate, and in many of those cases, known beneficial mutations in the recipients were replaced by donor alleles.
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Title: Factors influencing morphological evolution of the limbs in fossorial mammals (sciuridae and soricidae)
Creator: Swiderski, Donald L.
Date: 1990
Collection: Electronic Theses & Dissertations

Title: Genome evolution of Campylobacter jejuni during experimental adaptation
Creator: Jerome, John Paul
Date: 2012
Collection: Electronic Theses & Dissertations
Description: Campylobacter jejuni is a leading cause of foodborne bacterial enteritis in humans. An important reservoir for C. jejuni is in chickens, but it has been shown to colonize a large host range. Passage through a mouse model of campylobacteriosis resulted in a hypervirulent phenotype in mice for C. jejuni strain NCTC11168. After analyzing the wild-type and mouse-adapted variants by phenotype assays, expression microarray, pulse-field gel...
Show moreCampylobacter jejuni is a leading cause of foodborne bacterial enteritis in humans. An important reservoir for C. jejuni is in chickens, but it has been shown to colonize a large host range. Passage through a mouse model of campylobacteriosis resulted in a hypervirulent phenotype in mice for C. jejuni strain NCTC11168. After analyzing the wild-type and mouse-adapted variants by phenotype assays, expression microarray, pulse-field gel electrophoresis and whole genome sequencing we discovered that the genetic changes in the mouse-adapted variant were confined to thirteen hypermutable regions of DNA in contingency loci. We also show that specific contingency loci changes occurred in parallel during mouse infection when reisolates from multiple mice were analyzed. Furthermore, a mathematical model that considers contingency loci mutation rates and patterns does not explain the observed changes. Taken together, this is the first experimental evidence that contingency loci play a role in the rapid genetic adaptation of C. jejuni to a host, which results in increased virulence. In contrast to the observed virulence increase by serial host passage, we showed that C. jejuni rapidly loses an essential host colonization determinant during adaptive laboratory evolution. Passage in broth culture selected for flagellar motility deficient C. jejuni cells in parallel for five independently evolved lines. Moreover, the loss of motility occurred by two genetic mechanisms: reversible and irreversible. Reversible loss of motility occurred early during broth adaptation, followed by irreversible motility loss in the majority of cells by the end of the experiment. Whole genome sequencing implicated diverse mutation events that resulted in the loss of gene expression necessary for flagellar biosynthesis. Furthermore, reversible mutations in homopolymeric DNA tracts of adenine/thymine residues, and irreversible types of mutation such as gene deletion, were discovered in the broth-evolved populations. In all evolved lines, an alternative sigma factor necessary for flagellar structural gene expression was removed from the genome. Overall, this dissertation contains the first accounts of C. jejuni experimental evolution. The results provide insight into the biological importance of reversible mutations in homopolymeric DNA tracts, and provide a basis for future studies of C. jejuni evolvability.
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