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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: 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: The evolutionary origins of memory use in navigation
Creator: Grabowski, Laura M.
Date: 2009
Collection: Electronic Theses & Dissertations

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: Teaching and learning with digital evolution : factors influencing implementation and student outcomes
Creator: Lark, Amy M.
Date: 2014
Collection: Electronic Theses & Dissertations
Description: Science literacy for all Americans has been the rallying cry of science education in the United States for decades. Regardless, Americans continue to fall short when it comes to keeping pace with other developed nations on international science education assessments. To combat this problem, recent national reforms have reinvigorated the discussion of what and how we should teach science, advocating for the integration of disciplinary core ideas, crosscutting concepts, and science practices....
Show moreScience literacy for all Americans has been the rallying cry of science education in the United States for decades. Regardless, Americans continue to fall short when it comes to keeping pace with other developed nations on international science education assessments. To combat this problem, recent national reforms have reinvigorated the discussion of what and how we should teach science, advocating for the integration of disciplinary core ideas, crosscutting concepts, and science practices. In the biological sciences, teaching the core idea of evolution in ways consistent with reforms is fraught with challenges. Not only is it difficult to observe biological evolution in action, it is nearly impossible to engage students in authentic science practices in the context of evolution. One way to overcome these challenges is through the use of evolving populations of digital organisms.Avida-ED is digital evolution software for education that allows for the integration of science practice and content related to evolution. The purpose of this study was to investigate the effects of Avida-ED on teaching and learning evolution and the nature of science. To accomplish this I conducted a nationwide, multiple-case study, documenting how instructors at various institutions were using Avida-ED in their classrooms, factors influencing implementation decisions, and effects on student outcomes. I found that all of the participating instructors held views on teaching and learning that were well aligned with reform-based pedagogy, and although instructors used Avida-ED in a variety of ways, all adopted learner-centered pedagogical strategies that focused on the use of inquiry. After implementation, all of the instructors indicated that Avida-ED had allowed them to teach evolution and the nature of science in ways consistent with their personal teaching philosophies. In terms of assessment outcomes, students in lower-division courses significantly improved both their understanding and acceptance of evolution after using Avida-ED, and learning of content was positively associated with increased acceptance. Although student learning outcomes and instructor familiarity with Avida-ED were not associated with student affective response to the program, instructor familiarity was highly influential with regard to both how Avida-ED was implemented and student affective response, particularly student interest, enjoyment, and self-efficacy. The results of this dissertation provide strong evidence suggesting that Avida-ED is a promising tool for teaching and learning about evolution in reform-based ways, and suggest that improving instructor pedagogical content knowledge with regard to research-based tools like Avida-ED may be implicated in generating student interest in STEM.
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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: 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: Species pluralism
Creator: Holmes, John Alan
Date: 2000
Collection: Electronic Theses & Dissertations

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: 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: 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: Phylogeny of Astylopsis Casey (Coleoptera : Cerambycidae) species and patterns of host use
Creator: Camerato, Ellen M.
Date: 2021
Collection: Electronic Theses & Dissertations
Description: Cerambycid (longhorn) beetles are diverse in their morphology and life history traits, but all share the common trait of being larval plant-borers. The larvae bore into and develop inside various plant tissues thus they can potentially cause significant economic and ecological damage, especially when transported to non-native localities. There is little empirical data on cerambycid life history traits that are essential in understanding their ecological and economic effects. Astylopsis Casey ...
Show moreCerambycid (longhorn) beetles are diverse in their morphology and life history traits, but all share the common trait of being larval plant-borers. The larvae bore into and develop inside various plant tissues thus they can potentially cause significant economic and ecological damage, especially when transported to non-native localities. There is little empirical data on cerambycid life history traits that are essential in understanding their ecological and economic effects. Astylopsis Casey (Lamiinae: Acanthocinini) is an eastern North American genus of six species. Host preference varies greatly among the species, including both angiosperms and gymnosperms. I used morphological characters and molecular data to reconstruct phylogenies of Astylopsis to test the hypothesis that host plant use among Astylopsis species is conserved. I constructed phylogenies using partial COI and CAD DNA sequences from Astylopsis species and outgroups using parsimony methods. Astylopsis collaris, A. macula, A. sexguttata, and A. arcuata were monophyletic in both COI and combined gene phylogenies, with the genus also exhibiting monophyly in the combined gene tree. Evidence of host shift from angiosperms to gymnosperms in some species was also observed. These results confirm current taxonomic separations among the four species and their outgroups and provide important host use information. No conclusions could be drawn regarding DNA variation in association with geographic locality. These findings will inform future studies expanding the molecular dataset for Astylopsis with additional genes (arginine kinase, 28S, and EF1-α) and species (Astylopsis perplexa and A. fascipennis).
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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: 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: Microbial diversity and metabolic potential of the serpentinite subsurface environment
Creator: Twing, Katrina Irene
Date: 2015
Collection: Electronic Theses & Dissertations
Description: Serpentinization is the hydrous alteration of mafic rocks to form serpentine minerals and magnetite. The reactions of this alteration result in elevated pH of the surrounding fluids, abiotic generation of H2, CH4 (and other organic molecules), and depletion of dissolved inorganic carbon. Thus, serpentinization has implications for the origin of life on Earth and possibly Mars and other planetary bodies with water. The microbial diversity of continental serpentinite systems consistently shows...
Show moreSerpentinization is the hydrous alteration of mafic rocks to form serpentine minerals and magnetite. The reactions of this alteration result in elevated pH of the surrounding fluids, abiotic generation of H2, CH4 (and other organic molecules), and depletion of dissolved inorganic carbon. Thus, serpentinization has implications for the origin of life on Earth and possibly Mars and other planetary bodies with water. The microbial diversity of continental serpentinite systems consistently shows communities that are dominated by two major taxa – microaerophilic Betaproteobacteria and anaerobic Clostridia. Previous studies relied on few samples collected from natural springs or seeps, meaning that the flow path of fluids from the subsurface process of serpentinization was unknown. The Coast Range Ophiolite Microbial Observatory (CROMO), a set of wells drilled into the actively serpentinizing subsurface environment in northern California, was established in northern California to gain a better understanding of the habitability and microbial functions within the serpentinite subsurface environment. This dissertation represents a culmination of microbiological investigations into the serpentinite subsurface environment at CROMO to identify the microbial inhabitants of subsurface fluids, rocks, and in situ colonization experiments using molecular methods and high-throughput sequencing. The CROMO wells represent a broad range of geochemical gradients and pH and the concentrations of carbon monoxide and methane have the strongest correlation with microbial community composition. The most extremely high pH wells were inhabited exclusively by a single operational taxonomic unit (OTU) of Betaproteobacteria and a few OTUs of Clostridia, while more moderate pH wells exhibited greater diversity. Genes involved in the metabolism of hydrogen, carbon monoxide, and carbon fixation were abundant in the extreme pH fluids, while genes for metabolizing methane were exclusively in the moderate pH wells. The subsurface environment is an amalgamation of fluids and rocks, and as such, studying fluids alone only gives half the story. CROMO represents the first drill campaign into the continental serpentinite environment and the microbial diversity of serpentinite cores to a depth of 45 meters below surface suggests that specific geological features harbor different microbial communities. Archaea, previously undetected at CROMO, dominated cores containing magnetite-bearing serpentine, while bacteria were more abundant in layers containing clay particles. Additionally, organisms involved in the cycling of nitrogen and methane were found associated with core materials, indicating core-associated communities may have strong biogeochemical roles within the serpentinite subsurface environment. Given that microbial communities appear to vary with geological composition and that serpentinite fluids are a challenging habitat for life, depleted in inorganic carbon and electron acceptors, microbe-mineral interactions within the serpentinite subsurface environment through the use of in situ colonization devices to see if communities were able to utilize inorganic carbon in calcite or ferric iron as a terminal electron acceptor from magnetite. In the highest pH well, calcite led to an increased abundance of Clostridia and Deinococcus, while magnetite led to an increase in diversity, including Alphaproteobacteria, Gammaproteobacteria, and Actinobacteria, suggesting further that mineralogical composition of solids within the subsurface impact community composition. The data discussed here further our understanding of life associated with serpentinite fluids and minerals within the subsurface environment.
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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: 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: 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: Increasing student comprehension of evolution through laboratory investigations and simulations
Creator: McClintock, Steven W.
Date: 2008
Collection: Electronic Theses & Dissertations

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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