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Speciation is the fundamental process that has generated the vast diversity of life on earth. The hallmark of speciation is the evolution of barriers to gene flow. These barriers may reduce gene flow either by keeping incipient species from hybridizing at all (pre-zygotic), or by reducing the fitness of hybrids

Speciation is the fundamental process that has generated the vast diversity of life on earth. The hallmark of speciation is the evolution of barriers to gene flow. These barriers may reduce gene flow either by keeping incipient species from hybridizing at all (pre-zygotic), or by reducing the fitness of hybrids (post-zygotic). To understand the genetic architecture of these barriers and how they evolve, I studied a genus of wasps that exhibits barriers to gene flow that act both pre- and post-zygotically. Nasonia is a genus of four species of parasitoid wasps that can be hybridized in the laboratory. When two of these species, N. vitripennis and N. giraulti are mated, their offspring suffer, depending on the generation and cross examined, up to 80% mortality during larval development due to incompatible genic interactions between their nuclear and mitochondrial genomes. These species also exhibit pre-zygotic isolation, meaning they are more likely to mate with their own species when given the choice. I examined these two species and their hybrids to determine the genetic and physiological bases of both speciation mechanisms and to understand the evolutionary forces leading to them. I present results that indicate that the oxidative phosphorylation (OXPHOS) pathway, an essential pathway that is responsible for mitochondrial energy generation, is impaired in hybrids of these two species. These results indicate that this impairment is due to the unique evolutionary dynamics of the combined nuclear and mitochondrial origin of this pathway. I also present results showing that, as larvae, these hybrids experience retarded growth linked to the previously observed mortality and I explore possible physiological mechanisms for this. Finally, I show that the pre-mating isolation is due to a change in a single pheromone component in N. vitripennis males, that this change is under simple genetic control, and that it evolved neutrally before being co-opted as a species recognition signal. These results are an important addition to our overall understanding of the mechanisms of speciation and showcase Nasonia as an emerging model for the study of the genetics of speciation.
ContributorsGibson, Joshua D (Author) / Gadau, Jürgen (Thesis advisor) / Harrison, Jon (Committee member) / Pratt, Stephen (Committee member) / Verrelli, Brian (Committee member) / Willis, Wayne (Committee member) / Arizona State University (Publisher)
Created2013
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Description
Foraging has complex effects on whole-organism homeostasis, and there is considerable evidence that foraging behavior is influenced by both environmental factors (e.g., food availability, predation risk) and the physiological condition of an organism. The optimization of foraging behavior to balance costs and benefits is termed state-dependent foraging (SDF) while behavior

Foraging has complex effects on whole-organism homeostasis, and there is considerable evidence that foraging behavior is influenced by both environmental factors (e.g., food availability, predation risk) and the physiological condition of an organism. The optimization of foraging behavior to balance costs and benefits is termed state-dependent foraging (SDF) while behavior that seeks to protect assets of fitness is termed the asset protection principle (APP). A majority of studies examining SDF have focused on the role that energy balance has on the foraging of organisms with high metabolism and high energy demands ("high-energy systems" such as endotherms). In contrast, limited work has examined whether species with low energy use ("low-energy systems" such as vertebrate ectotherms) use an SDF strategy. Additionally, there is a paucity of evidence demonstrating how physiological and environmental factors other than energy balance influence foraging behavior (e.g. hydration state and free-standing water availability). Given these gaps in our understanding of SDF behavior and the APP, I examined the state-dependency and consequences of foraging in a low-energy system occupying a resource-limited environment - the Gila monster (Heloderma suspectum, Cope 1869). In contrast to what has been observed in a wide variety of taxa, I found that Gila monsters do not use a SDF strategy to manage their energy reserves and that Gila monsters do not defend their energetic assets. However, hydration state and free-standing water availability do affect foraging behavior of Gila monsters. Additionally, as Gila monsters become increasingly dehydrated, they reduce activity to defend hydration state. The SDF behavior of Gila monsters appears to be largely driven by the fact that Gila monsters must separately satisfy energy and water demands with food and free-standing water, respectively, in conjunction with the timescale within which Gila monsters balance their energy and water budgets (supra-annually versus annually, respectively). Given these findings, the impact of anticipated changes in temperature and rainfall patterns in the Sonoran Desert are most likely going to pose their greatest risks to Gila monsters through the direct and indirect effects on water balance.
ContributorsWright, Christian (Author) / Denardo, Dale F. (Thesis advisor) / Harrison, Jon (Committee member) / McGraw, Kevin (Committee member) / Sullivan, Brian (Committee member) / Wolf, Blair (Committee member) / Arizona State University (Publisher)
Created2014
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Description
While exercising mammalian muscle increasingly relies on carbohydrates for fuel as aerobic exercise intensity rises above the moderate range, flying birds are extraordinary endurance athletes and fuel flight, a moderate-high intensity exercise, almost exclusively with lipid. In addition, Aves have long lifespans compared to weight-matched mammals. As skeletal muscle mitochondria

While exercising mammalian muscle increasingly relies on carbohydrates for fuel as aerobic exercise intensity rises above the moderate range, flying birds are extraordinary endurance athletes and fuel flight, a moderate-high intensity exercise, almost exclusively with lipid. In addition, Aves have long lifespans compared to weight-matched mammals. As skeletal muscle mitochondria account for the majority of oxygen consumption during aerobic exercise, the primary goal was to investigate differences in isolated muscle mitochondria between these species and to examine to what extent factors intrinsic to mitochondria may account for the behavior observed in the intact tissue and whole organism. First, maximal enzyme activities were assessed in sparrow and rat mitochondria. Citrate synthase and aspartate aminotransferase activity were higher in sparrow compared to rat mitochondria, while glutamate dehydrogenase activity was lower. Sparrow mitochondrial NAD-linked isocitrate dehydrogenase activity was dependent on phosphate, unlike the mammalian enzyme. Next, the rate of oxygen consumption (JO), electron transport chain (ETC) activity, and reactive oxygen species (ROS) production were assessed in intact mitochondria. Maximal rates of fat oxidation were lower than for carbohydrate in rat but not sparrow mitochondria. ETC activity was higher in sparrows, but no differences were found in ROS production between species. Finally, fuel selection and control of respiration at three rates between rest and maximum were assessed. Mitochondrial fuel oxidation and selection mirrored that of the whole body; in rat mitochondria the reliance on carbohydrate increased as the rate of oxygen consumption increased, whereas fat dominated under all conditions in the sparrow. These data indicate fuel selection, at least in part, can be modulated at the level of the mitochondrial matrix when multiple substrates are present at saturating levels. As an increase in matrix oxidation-reduction potential has been linked to a suppression of fat oxidation and high ROS production, the high ETC activity relative to dehydrogenase activity in avian compared to mammalian mitochondria may result in lower matrix oxidation-reduction potential, allowing fatty acid oxidation to proceed while also resulting in low ROS production in vivo.
ContributorsKuzmiak, Sarah (Author) / Willis, Wayne T (Thesis advisor) / Mandarino, Lawrence (Committee member) / Sweazea, Karen (Committee member) / Harrison, Jon (Committee member) / Gadau, Juergen (Committee member) / Arizona State University (Publisher)
Created2012
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Description
Land management practices such as domestic animal grazing can alter plant communities via changes in soil structure and chemistry, species composition, and plant nutrient content. These changes can affect the abundance and quality of plants consumed by insect herbivores with consequent changes in population dynamics. These population changes can translate

Land management practices such as domestic animal grazing can alter plant communities via changes in soil structure and chemistry, species composition, and plant nutrient content. These changes can affect the abundance and quality of plants consumed by insect herbivores with consequent changes in population dynamics. These population changes can translate to massive crop damage and pest control costs. My dissertation focused on Oedaleus asiaticus, a dominant Asian locust, and had three main objectives. First, I identified morphological, physiological, and behavioral characteristics of the migratory ("brown") and non-migratory ("green") phenotypes. I found that brown morphs had longer wings, larger thoraxes and higher metabolic rates compared to green morphs, suggesting that developmental plasticity allows greater migratory capacity in the brown morph of this locust. Second, I tested the hypothesis of a causal link between livestock overgrazing and an increase in migratory swarms of O. asiaticus. Current paradigms generally assume that increased plant nitrogen (N) should enhance herbivore performance by relieving protein-limitation, increasing herbivorous insect populations. I showed, in contrast to this scenario, that host plant N-enrichment and high protein artificial diets decreased the size and viability of O. asiaticus. Plant N content was lowest and locust abundance highest in heavily livestock-grazed fields where soils were N-depleted, likely due to enhanced erosion and leaching. These results suggest that heavy livestock grazing promotes outbreaks of this locust by reducing plant protein content. Third, I tested for the influence of dietary imbalance, in conjunction with high population density, on migratory plasticity. While high population density has clearly been shown to induce the migratory morph in several locusts, the effect of diet has been unclear. I found that locusts reared at high population density and fed unfertilized plants (i.e. high quality plants for O. asiaticus) had the greatest migratory capacity, and maintained a high percent of brown locusts. These results did not support the hypothesis that poor-quality resources increased expression of migratory phenotypes. This highlights a need to develop new theoretical frameworks for predicting how environmental factors will regulate migratory plasticity in locusts and perhaps other insects.
ContributorsCease, Arianne (Author) / Harrison, Jon (Thesis advisor) / Elser, James (Thesis advisor) / DeNardo, Dale (Committee member) / Quinlan, Michael (Committee member) / Sabo, John (Committee member) / Arizona State University (Publisher)
Created2012
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Description
Phosphorus (P), an essential nutrient for growth of all organisms, is often in limited biological supply for herbivore consumers compared to other elements, such as carbon (C). Ecological stoichiometry studies have assessed responses of filter-feeding zooplankton from the genus Daphnia to single and multi-species food resources that are P-limited,

Phosphorus (P), an essential nutrient for growth of all organisms, is often in limited biological supply for herbivore consumers compared to other elements, such as carbon (C). Ecological stoichiometry studies have assessed responses of filter-feeding zooplankton from the genus Daphnia to single and multi-species food resources that are P-limited, finding decreased growth as a result to changes in metabolic processes and feeding behavior. Conversely, recent laboratory studies have shown that P-rich algal food resources also result in decreased growth rates for Daphnia, though the possible mechanisms behind this maladaptive response is understudied. Moreover, no published study tests the existence of the “stoichiometric knife edge” hypothesis for low C:P under field conditions. To address this lack of information, I measured growth rate as well as respiration and ingestion rates for D. magna, D. pulicaria, and D. pulex that were fed natural lake seston experimentally enriched with different levels of PO43-. I found heterogeneous effects of high dietary P across Daphnia species. Growth rate responses for D. magna were strong and indicated a negative effect of high-P, most likely as a result to decreased ingestion rates that were observed. The seston treatments did not elicit significant growth rate responses for D. pulex and D. pulicaria, but significant responses to respiration rates were observed for all species. Consumer body stoichiometry, differences in seston C:P for each experiment, or differential assimilation by producer types may be driving these results. My study suggests that the stoichiometric knife edge documented in laboratory studies under low C:P conditions may not operate to the same degree when natural seston is the food source; diet diversity may be driving complex nuances for consumer performance that were previously overlooked.
ContributorsCurrier, Courtney M (Author) / Currier, James (Thesis advisor) / Harrison, Jon (Committee member) / Neuer, Susanne (Committee member) / Arizona State University (Publisher)
Created2015
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Description
Honey bees (Apis mellifera) are responsible for pollinating nearly 80\% of all pollinated plants, meaning humans depend on honey bees to pollinate many staple crops. The success or failure of a colony is vital to global food production. There are various complex factors that can contribute to a colony's failure,

Honey bees (Apis mellifera) are responsible for pollinating nearly 80\% of all pollinated plants, meaning humans depend on honey bees to pollinate many staple crops. The success or failure of a colony is vital to global food production. There are various complex factors that can contribute to a colony's failure, including pesticides. Neonicotoids are a popular pesticide that have been used in recent times. In this study we concern ourselves with pesticides and its impact on honey bee colonies. Previous investigations that we draw significant inspiration from include Khoury et Al's \emph{A Quantitative Model of Honey Bee Colony Population Dynamics}, Henry et Al's \emph{A Common Pesticide Decreases Foraging Success and Survival in Honey Bees}, and Brown's \emph{ Mathematical Models of Honey Bee Populations: Rapid Population Decline}. In this project we extend a mathematical model to investigate the impact of pesticides on a honey bee colony, with birth rates and death rates being dependent on pesticides, and we see how these death rates influence the growth of a colony. Our studies have found an equilibrium point that depends on pesticides. Trace amounts of pesticide are detrimental as they not only affect death rates, but birth rates as well.
ContributorsSalinas, Armando (Author) / Vaz, Paul (Thesis director) / Jones, Donald (Committee member) / School of Mathematical and Statistical Sciences (Contributor) / School of International Letters and Cultures (Contributor) / Barrett, The Honors College (Contributor)
Created2016-05
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Description
We examined the evolutionary morphological responses of Drosophila melanogaster that had evolved at constant cold (16°), constant hot (25°C), and fluctuating (16° and 25°C). Flies that were exposed to the constant low mean temperature developed larger thorax, wing, and cell sizes than those exposed to constant high mean temperatures. Males

We examined the evolutionary morphological responses of Drosophila melanogaster that had evolved at constant cold (16°), constant hot (25°C), and fluctuating (16° and 25°C). Flies that were exposed to the constant low mean temperature developed larger thorax, wing, and cell sizes than those exposed to constant high mean temperatures. Males and females both responded similarly to thermal treatments in average wing and cell size. The resulting cell area for a given wing size in thermal fluctuating populations remains unclear and remains a subject for future research.
ContributorsAdrian, Gregory John (Author) / Angilletta, Michael (Thesis director) / Harrison, Jon (Committee member) / Rusch, Travis (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor)
Created2015-05
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Description
Currently in synthetic biology only the Las, Lux, and Rhl quorum sensing pathways have been adapted for broad engineering use. Quorum sensing allows a means of cell to cell communication in which a designated sender cell produces quorum sensing molecules that modify gene expression of a designated receiver cell. While

Currently in synthetic biology only the Las, Lux, and Rhl quorum sensing pathways have been adapted for broad engineering use. Quorum sensing allows a means of cell to cell communication in which a designated sender cell produces quorum sensing molecules that modify gene expression of a designated receiver cell. While useful, these three quorum sensing pathways exhibit a nontrivial level of crosstalk, hindering robust engineering and leading to unexpected effects in a given design. To address the lack of orthogonality among these three quorum sensing pathways, previous scientists have attempted to perform directed evolution on components of the quorum sensing pathway. While a powerful tool, directed evolution is limited by the subspace that is defined by the protein. For this reason, we take an evolutionary biology approach to identify new orthogonal quorum sensing networks and test these networks for cross-talk with currently-used networks. By charting characteristics of acyl homoserine lactone (AHL) molecules used across quorum sensing pathways in nature, we have identified favorable candidate pathways likely to display orthogonality. These include Aub, Bja, Bra, Cer, Esa, Las, Lux, Rhl, Rpa, and Sin, which we have begun constructing and testing. Our synthetic circuits express GFP in response to a quorum sensing molecule, allowing quantitative measurement of orthogonality between pairs. By determining orthogonal quorum sensing pairs, we hope to identify and adapt novel quorum sensing pathways for robust use in higher-order genetic circuits.
ContributorsMuller, Ryan (Author) / Haynes, Karmella (Thesis director) / Wang, Xiao (Committee member) / Barrett, The Honors College (Contributor) / School of Mathematical and Statistical Sciences (Contributor) / Department of Chemistry and Biochemistry (Contributor) / School of Life Sciences (Contributor)
Created2015-05
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Description
Three populations of experimentally evolved Drosophila melanogaster populations made up of high temperature (H, constant 25 ᵒC), low temperature (C, constant 16 ᵒC) and temporal homogeneity (T, environment changes between 16 ᵒC and 25 ᵒC) were prepared and assayed to determine difference in citrate synthase activity. Between the three groups,

Three populations of experimentally evolved Drosophila melanogaster populations made up of high temperature (H, constant 25 ᵒC), low temperature (C, constant 16 ᵒC) and temporal homogeneity (T, environment changes between 16 ᵒC and 25 ᵒC) were prepared and assayed to determine difference in citrate synthase activity. Between the three groups, the results were inconclusive: the resulting reaction rates in units of nmol min-1mgfly-1 were 81.8 + 20.6, 101 + 15.6, and 96.9 + 25.2 for the hot (H), cold (C), and temporally homogeneous (T) groups, respectively. We conclude that the high associated variability was due to a lack of control regarding the collection time of the experimentally evolved Drosophila.
ContributorsBelohlavek, David (Author) / Angilletta, Michael (Thesis director) / Francisco, Wilson (Committee member) / Barrett, The Honors College (Contributor) / Department of Chemistry and Biochemistry (Contributor)
Created2015-05
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Description
For colonies of ponerine ant species, sterility regulation after a founding queen's death is not totally achieved in the worker caste, and the possibility of sexual reproduction is opened to workers. The persisting survival of these colonies is dependent on capturing the optimal reproductive ratio; yet, an informational gap bounds

For colonies of ponerine ant species, sterility regulation after a founding queen's death is not totally achieved in the worker caste, and the possibility of sexual reproduction is opened to workers. The persisting survival of these colonies is dependent on capturing the optimal reproductive ratio; yet, an informational gap bounds the mechanisms detailing the selection of new reproductives and the suppression of ovarian development in rejected reproductives. We investigated the mechanisms of worker policing, one of the primary methods of ovarian suppression, through continuous video observation for a period of five days at the start of colony instability. Observations suggest policing in H. saltator is performed by a majority of a colony, including potential reproductives, and requires multiple events to fully discourage ovarian growth.
ContributorsChien, Jeffrey (Co-author) / Barat Ali, Fatima (Co-author) / Kang, Yun (Thesis director) / Liebig, Juergen (Committee member) / School of Mathematical and Statistical Sciences (Contributor) / Mechanical and Aerospace Engineering Program (Contributor) / Barrett, The Honors College (Contributor)
Created2018-12