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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: Evolutionary dynamics of 3D digital constructs
Creator: Stredwick, Jason Michael
Date: 2005
Collection: Electronic Theses & Dissertations

Title: Genome-wide association study reveals genes associated with mite recruitment phenotypes in the domesticated grapevine (vitis vinifera
Creator: LaPlante, Erika R.
Date: 2020
Collection: Electronic Theses & Dissertations
Description: Plants in the grape genus Vitis have varying densities of trichomes and mite-domatia on their leaves, which facilitate the recruitment, retention, and reproduction of beneficial mites. By increasing the abundance of mites on grape leaves, these phenotypes promote a defense mutualism contributing to the control of grape pests and pathogens. Identification of the genes controlling these phenotypes would inform our understanding of the genetics underlying mite-plant mutualistic interactions and...
Show morePlants in the grape genus Vitis have varying densities of trichomes and mite-domatia on their leaves, which facilitate the recruitment, retention, and reproduction of beneficial mites. By increasing the abundance of mites on grape leaves, these phenotypes promote a defense mutualism contributing to the control of grape pests and pathogens. Identification of the genes controlling these phenotypes would inform our understanding of the genetics underlying mite-plant mutualistic interactions and could lead to breeding domesticated Vitis vinifera L. varieties that are naturally defended against pathogens. Little is known about the genetics underlying mite recruitment phenotypes in Vitis. We conducted a GWAS to determine the genetic architecture of mite recruitment traits in V. vinifera using 399 cultivars from a common garden diversity panel. We investigated eight traits previously established in the literature associated with an increase in beneficial mite abundance. We found single nucleotide polymorphisms (SNPs) significantly associated with each mite recruitment trait investigated. Corresponding gene annotations of SNP genetic coordinates revealed notable gene associations, including a trichome development gene, and a physiological defense response gene, suggesting these genetic regions may have a large impact on mediating mite-plant interactions in this species. Our findings are among the first to investigate which genes underly ecologically important mutualisms between plants and beneficial mites and suggest promising candidate genes for breeding and genetic editing to increase naturally occurring predator-based defenses in grapes.
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Title: THE TRANSCRIPTOMIC AND EPIGENOMIC RESPONSE OF KOCHIA SCOPARIA TO SUBLETHAL GLYPHOSATE
Creator: Claucherty, Carly Abbegail
Date: 2022
Collection: Electronic Theses & Dissertations
Description: Weed populations respond and adapt to herbicide stress by evolving resistance. Glyphosate resistance is primarily caused by the amplification of the target site gene, EPSPS, where multiple copies produce a large enough protein pool so that field rates do not kill the plant. This mechanism has evolved independently in at least nine divergent weed species. It has been demonstrated that EPSPS gene duplication may be transposon mediated in Kochia scoparia. A key regulator of transposable element ...
Show moreWeed populations respond and adapt to herbicide stress by evolving resistance. Glyphosate resistance is primarily caused by the amplification of the target site gene, EPSPS, where multiple copies produce a large enough protein pool so that field rates do not kill the plant. This mechanism has evolved independently in at least nine divergent weed species. It has been demonstrated that EPSPS gene duplication may be transposon mediated in Kochia scoparia. A key regulator of transposable element (TE) activity is DNA methylation. The role of the epigenome and subsequent transcriptome in transient responses to herbicides of their primary target, weeds, is not well understood In this study, we performed RNA-Seq and bisulfite sequencing on leaf tissue from glyphosate-sensitive kochia before and three weeks after treatment with two sublethal doses to determine if glyphosate causes hypomethylation of the genome, allowing for the activation of transposons and upregulation of stress-related genes. Our results shows that overall gene expression was suppressed by glyphosate and increases in CHH methylation through development were also ceased. We did not observe significant global changes in cytosine methylation, and overall responses were stochastic. When combining the two datasets together, there was no direct correlation between changes in methylation and changes in gene expression suggesting that DNA methylation is not the primary cause of differential expression in our study. Our results broaden the knowledge pool of weedy species epigenomics and aid in understanding the contribution of DNA methylation to plant resilience in response to herbicide stress.
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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: 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: Oceanic salt spray and herbivore pressure contribute to local adaptation of coastal perennial and inland annual ecotypes of the Seep Monkeyflower (Mimulus guttatus)
Creator: Popovic, Damian
Date: 2018
Collection: Electronic Theses & Dissertations
Description: In this study, we used the emergent model system Mimulus guttatus to explore the agents of selection that drive local adaptation across California’s coast-inland moisture gradient. We implemented a field reciprocal transplant experiment within agrofabric exclosures at sites in coastal and inland Sonoma County, California – minimizing the effects of aboveground stressors in an effort to elucidate their role in the evolution of local adaptation in this system. ASTER life-history modeling and...
Show moreIn this study, we used the emergent model system Mimulus guttatus to explore the agents of selection that drive local adaptation across California’s coast-inland moisture gradient. We implemented a field reciprocal transplant experiment within agrofabric exclosures at sites in coastal and inland Sonoma County, California – minimizing the effects of aboveground stressors in an effort to elucidate their role in the evolution of local adaptation in this system. ASTER life-history modeling and generalized linear mixed modeling approaches were used to analyze survival and dry aboveground biomass as fitness proxies. Despite altering no edaphic conditions, we found that among coastal exclosures, inland replicates were significantly more fit than their control counterparts, essentially rescuing inland fitness outside of their native range. Exclosures provided no fitness advantage for either ecotype at the inland site, aside from a moderate but statistically significant increase in biomass among exclosed coastal replicates. While it is unlikely to know all the agents of selection limited by our exclosures, we found that the number of replicates that experienced herbivory were significantly lower across all exclosures at the coast site. An elemental analysis of Sodium (Na) using salt traps installed at each site also demonstrated a reduction in salt exposure within exclosures. It is likely that some combination of aboveground stressors, likely driven primarily by herbivory and/or salt stress, plays a continuing role in the evolution of coastal M. guttatus populations, thus providing a new understanding of how local adaptation is maintained in this model system.
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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: Do latex and resin canals spur plant diversification? : re-examining a classic example of escape and radiate coevolution
Creator: Foisy, Michael
Date: 2020
Collection: Electronic Theses & Dissertations
Description: The association between increased lineage diversification rates and the evolution of latex and resin canals is widely cited as a paradigmatic example of Ehrlich and Raven's 'escape-and-radiate' hypothesis of co-evolution. However, it has been nearly three decades since these plant defenses have been examined as key innovations, and updates to phylogenetic comparative methods, plant molecular systematics, and phenotypic data warrant a reassessment of this classic finding. I gathered data on...
Show moreThe association between increased lineage diversification rates and the evolution of latex and resin canals is widely cited as a paradigmatic example of Ehrlich and Raven's 'escape-and-radiate' hypothesis of co-evolution. However, it has been nearly three decades since these plant defenses have been examined as key innovations, and updates to phylogenetic comparative methods, plant molecular systematics, and phenotypic data warrant a reassessment of this classic finding. I gathered data on latex and resin canals across 345 families and 986 genera of vascular plants and conducted a multi-scale test of the association between these traits and lineage diversification rates. At a broad scale (across clades), I used sister-clade comparisons to test whether 28 canal-bearing clades had higher net diversification rates than their canal-lacking sister clades. At a finer scale (within clades), I used ancestral state reconstructions and phylogenetic models of lineage diversification rates to examine the relationship between trait evolution and the timing of diversification rate shifts in two better-characterized clades - Araceae and Papaveraceae. At both scales of analyses, I found poor support for the predicted relationship between diversification and the evolution of latex and resin canals. This re-examination reveals that there is no longer strong evidence for latex or resin canals as general, consistently replicable drivers of species diversity across plants. However, I could not rule out a relationship in all groups, and therefore argue that theoretical and empirical work aimed at understanding ecological factors that condition 'escape-and-radiate' dynamics will allow for more nuanced tests of the hypothesis in the future.
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Title: The role of evolutionary history and niche differentiation in structuring species co-occurrence in New Zealand Pittosporum (Pittosporaceae)
Creator: Nolting, Kristen Marie
Date: 2014
Collection: Electronic Theses & Dissertations
Description: Understanding the processes that shape the spatial distribution of species, and their co-existence in communities, remains one of the biggest challenges that ecologists face today. Investigations of co-existence typically treat species as independent entities, thereby ignoring their shared evolutionary history, niche preferences and functional similarity. This limits the ability of ecologists to make strong inferences regarding co-existence mechanisms. It is more useful to employ a...
Show moreUnderstanding the processes that shape the spatial distribution of species, and their co-existence in communities, remains one of the biggest challenges that ecologists face today. Investigations of co-existence typically treat species as independent entities, thereby ignoring their shared evolutionary history, niche preferences and functional similarity. This limits the ability of ecologists to make strong inferences regarding co-existence mechanisms. It is more useful to employ a pluralistic approach that integrates phylogenetic information and species-specific environmental and trait associations. In this study I evaluated the role of evolutionary history and environmental and functional trait differentiation in predicting species co-occurrence in the New Zealand plant genus Pittosporum. I hypothesized that co-occurring species would be more distantly related than non co-occurring species, given that closely related species are likely to be more ecologically similar as a result of their shared ancestry, and thus competitive interactions would preclude them from co-occurring. Similarly, I predicted that co-occurring species would be more divergent in their functional traits to enable co-existence. I found that co-occurring species were no different than non co-occurring species with respect to their phylogenetic dissimilarity, that co-occurring species had higher environmental niche overlap than non co-occurring species, and that for most traits measured there was no difference in trait dissimilarity among co-occurring and non co-occurring species. Approximate maximum vessel length and leaf nitrogen content, however, showed convergence among co-occurring species.
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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: Assessing the viability of the shifting balance process
Creator: Moore, Francis B.,-G
Date: 1996
Collection: Electronic Theses & Dissertations

Title: Adaptation and specialization in biological and digital organisms
Creator: Ostrowski, Elizabeth Anne
Date: 2005
Collection: Electronic Theses & Dissertations

Title: Local adaptation and fitness trade-offs
Creator: Dittmar, Emily Loring
Date: 2017
Collection: Electronic Theses & Dissertations
Description: Adaptation generates and maintains genetic and phenotypic diversity. This is thought to occur due to trade-offs, where adaptation to one environment comes at a cost in another. Although trade-offs are believed to play a prominent role in the generation and maintenance of genetic and phenotypic diversity, the mechanisms by which adaptation leads to trade-offs are not well understood.My research explores the forces that lead to adaptive trade-offs in two systems. First, using a RIL mapping...
Show moreAdaptation generates and maintains genetic and phenotypic diversity. This is thought to occur due to trade-offs, where adaptation to one environment comes at a cost in another. Although trade-offs are believed to play a prominent role in the generation and maintenance of genetic and phenotypic diversity, the mechanisms by which adaptation leads to trade-offs are not well understood.My research explores the forces that lead to adaptive trade-offs in two systems. First, using a RIL mapping population created from natural populations of Arabidopsis thaliana, I studied the genetic basis of flowering time, a putatively adaptive trait and one that differs between the parental populations. I identified flowering time QTL in growth chambers that mimicked the natural temperature and photoperiod variation across the growing season in each native environment and compared the genomic locations of flowering time QTL to those of fitness (total fruit number) QTL from a previous three-year field study.In addition, I studied two populations of Leptosiphon parviflorus, an annual wildflower native to California. At Jasper Ridge biological preserve, populations of L. parviflorus grow on and off serpentine soil in close proximity. Due to its harsh growing conditions, serpentine soil exerts strong selective pressures on plants. Despite the close proximity of study populations (<100 m) and ongoing gene flow, reciprocal transplant studies demonstrate that these populations are locally adapted to their native soil types. To determine the selective agents operating in both habitats and the forces underlying fitness trade-offs, I performed manipulative experiments in the field and greenhouse. Results from these studies show that both soil moisture and competitive interactions are important for mediating fitness differences among the populations, and adaptation to serpentine soil might result in a cost to competitive ability.I also addressed the causes of flowering-time differences in these populations. Field reciprocal-transplant studies and watering manipulations in the greenhouse demonstrate the contribution of both the genotype and the environment to observed flowering-time differences. The plasticity of flowering time in response to soil type appears to be driven by differences in soil moisture. In addition, selection on flowering time was measured in both soil types across four years of study using a set of F5 advanced generation hybrids and found to differ among the habitats. Therefore, both selection and plasticity contribute to flowering-time differences between these populations and thus have likely played an important role in the initiation and/or maintenance of adaptive divergence in this system.Finally, the two populations differ in their flower color, a Mendelian trait. Pollinators do not discriminate among flower colors and are unlikely to exert selection on this trait. Instead, flower color may be related to stress tolerance if the causal gene has pleiotropic effects on other traits. Using a set of Near Isogenic Lines (NILs), I found that the flower color locus has an effect on survival in field soil and fecundity in benign conditions. Ongoing work is aimed at addressing the mechanisms underlying the relationship between flower color and soil adaptation.
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Title: The evolutionary origins of memory use in navigation
Creator: Grabowski, Laura M.
Date: 2009
Collection: Electronic Theses & Dissertations

Title: A differential evolution approach to feature selection in genomic prediction
Creator: Whalen, Ian
Date: 2018
Collection: Electronic Theses & Dissertations
Description: The use of genetic markers has become widespread for prediction of genetic merit in agricultural applications and is a beginning to show promise for estimating propensity to disease in human medicine. This process is known as genomic prediction and attempts to model the mapping between an organism's genotype and phenotype. In practice, this process presents a challenging problem. Sequencing and recording phenotypic traits are often expensive and time consuming. This leads to datasets often...
Show moreThe use of genetic markers has become widespread for prediction of genetic merit in agricultural applications and is a beginning to show promise for estimating propensity to disease in human medicine. This process is known as genomic prediction and attempts to model the mapping between an organism's genotype and phenotype. In practice, this process presents a challenging problem. Sequencing and recording phenotypic traits are often expensive and time consuming. This leads to datasets often having many more features than samples. Common models for genomic prediction often fall victim to overfitting due to the curse of dimensionality. In this domain, only a fraction of the markers that are present significantly affect a particular trait. Models that fit to non-informative markers are in effect fitting to statistical noise, leading to a decrease in predictive performance. Therefore, feature selection is desirable to remove markers that do not appear to have a significant effect on the trait being predicted. The method presented here uses differential evolution based search for feature selection. This study will characterize differential evolution's efficacy in feature selection for genomic prediction and present several extensions to the base search algorithm in an attempt to apply domain knowledge to guide the search toward better solutions.
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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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