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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
Microalgae represent a potential sustainable alternative for the enhancement and protection of agricultural crops. The dry biomass and cellular extracts of Scenedesmus dimorphus were applied as a biofertilizer, a foliar spray, and a seed primer to evaluate seed germination, plant growth, and crop yield of Roma tomato plants. The dry

Microalgae represent a potential sustainable alternative for the enhancement and protection of agricultural crops. The dry biomass and cellular extracts of Scenedesmus dimorphus were applied as a biofertilizer, a foliar spray, and a seed primer to evaluate seed germination, plant growth, and crop yield of Roma tomato plants. The dry biomass was applied as a biofertilizer at 50 g and 100 g per plant, to evaluate its effects on plant development and crop yield. Biofertilizer treatments enhanced plant growth and led to greater crop (fruit) production. Timing of biofertilizer application proved to be of importance - earlier 50 g biofertilizer application resulted in greater plant growth. Scenedesmus dimorphus culture, growth medium, and different concentrations (1%, 5%, 10%, 25%, 50%, 75%, 100%) of aqueous cell extracts were used as seed primers to determine effects on germination. Seeds treated with Scenedesmus dimorphus culture and with extract concentrations higher than 50 % (0.75 g ml-1) triggered faster germination - 2 days earlier than the control group. Extract foliar sprays of 50 ml and 100 ml, were obtained and applied to tomato plants at various extract concentrations (10%, 25%, 50%, 75% and 100%). Plant height, flower development and number of branches were significantly enhanced with 50 % (7.5 g ml-1) extracts. Higher concentration sprays led to a decrease in growth. The extracts were further screened to assess potential antimicrobial activity against the bacterium Escherichia coli ATCC 25922, the fungi Candida albicans ATCC 90028 and Aspergillus brasiliensis ATCC 16404. No antimicrobial activity was observed from the microalga extracts on the selected microorganisms.
ContributorsGarcia-Gonzalez, Jesus (Author) / Sommerfeld, Milton (Thesis advisor) / Steele, Kelly (Committee member) / Henderson, Mark (Committee member) / Arizona State University (Publisher)
Created2014
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Description
The repression of reproductive competition and the enforcement of altruism are key components to the success of animal societies. Eusocial insects are defined by having a reproductive division of labor, in which reproduction is relegated to one or few individuals while the rest of the group members maintain the colony

The repression of reproductive competition and the enforcement of altruism are key components to the success of animal societies. Eusocial insects are defined by having a reproductive division of labor, in which reproduction is relegated to one or few individuals while the rest of the group members maintain the colony and help raise offspring. However, workers have retained the ability to reproduce in most insect societies. In the social Hymenoptera, due to haplodiploidy, workers can lay unfertilized male destined eggs without mating. Potential conflict between workers and queens can arise over male production, and policing behaviors performed by nestmate workers and queens are a means of repressing worker reproduction. This work describes the means and results of the regulation of worker reproduction in the ant species Aphaenogaster cockerelli. Through manipulative laboratory studies on mature colonies, the lack of egg policing and the presence of physical policing by both workers and queens of this species are described. Through chemical analysis and artificial chemical treatments, the role of cuticular hydrocarbons as indicators of fertility status and the informational basis of policing in this species is demonstrated. An additional queen-specific chemical signal in the Dufour's gland is discovered to be used to direct nestmate aggression towards reproductive competitors. Finally, the level of actual worker-derived males in field colonies is measured. Together, these studies demonstrate the effectiveness of policing behaviors on the suppression of worker reproduction in a social insect species, and provide an example of how punishment and the threat of punishment is a powerful force in maintaining cooperative societies.
ContributorsSmith, Adrian A. (Author) / Liebig, Juergen (Thesis advisor) / Hoelldobler, Bert (Thesis advisor) / Gadau, Juergen (Committee member) / Johnson, Robert A. (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2011
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Description
Gene-centric theories of evolution by natural selection have been popularized and remain generally accepted in both scientific and public paradigms. While gene-centrism is certainly parsimonious, its explanations fall short of describing two patterns of evolutionary and social phenomena: the evolution of sex and the evolution of social altruism. I review

Gene-centric theories of evolution by natural selection have been popularized and remain generally accepted in both scientific and public paradigms. While gene-centrism is certainly parsimonious, its explanations fall short of describing two patterns of evolutionary and social phenomena: the evolution of sex and the evolution of social altruism. I review and analyze current theories on the evolution of sex. I then introduce the conflict presented to gene-centric evolution by social phenomena such as altruism and caste sterility in eusocial insects. I review gene-centric models of inclusive fitness and kin selection proposed by Hamilton and Maynard Smith. Based their assumptions, that relatedness should be equal between sterile workers and reproductives, I present several empirical examples that conflict with their models. Following that, I introduce a unique system of genetic caste determination (GCD) observed in hybrid populations of two sister-species of seed harvester ants, Pogonomyrmex rugosus and Pogonomyrmex barbatus. I review the evidence for GCD in those species, followed by a critique of the current gene-centric models used to explain it. In chapter two I present my own theoretical model that is both simple and extricable in nature to explain the origin, evolution, and maintenance of GCD in Pogonomyrmex. Furthermore, I use that model to fill in the gaps left behind by the contributing authors of the other GCD models. As both populations in my study system formed from inter-specific hybridization, I review modern discussions of heterosis (also called hybrid vigor) and use those to help explain the ecological competitiveness of GCD. I empirically address the inbreeding depression the lineages of GCD must overcome in order to remain ecologically stable, demonstrating that as a result of their unique system of caste determination, GCD lineages have elevated recombination frequencies. I summarize and conclude with an argument for why GCD evolved under selective mechanisms which cannot be considered gene-centric, providing evidence that natural selection can effectively operate on non-heritable genotypes appearing in groups and other social contexts.
ContributorsJacobson, Neal (Author) / Gadau, Juergen (Thesis advisor) / Laubichler, Manfred (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2012
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Description
For the past 30 years wildlife biologists have debated the need of pronghorn antelope (Antilocapra americana) to drink freestanding water (free water). Some have suggested that pronghorn may feed at night to increase preformed water (plant moisture) intake, thus decreasing their dependence on free water. Pronghorn diet composition and nutrient

For the past 30 years wildlife biologists have debated the need of pronghorn antelope (Antilocapra americana) to drink freestanding water (free water). Some have suggested that pronghorn may feed at night to increase preformed water (plant moisture) intake, thus decreasing their dependence on free water. Pronghorn diet composition and nutrient intake is integral to understanding water available to pronghorn through preformed and metabolic sources. The dual purpose of this study was to determine plant composition of pronghorn diets, and to examine whether night feeding provides a water allocation advantage by testing for differences between day and night and modeling free water requirements during biologically critical seasons and years of different precipitation. I determined species composition, selected nutrients, and moisture content of American pronghorn diets on Perry Mesa, Arizona in March, May, June and August of 2008 and 2009. I used microhistological analysis of fecal samples to determine percent plant composition of pronghorn diets. I used forage samples to evaluate the nutrient composition of those diets for moisture, crude protein and structural carbohydrates, and to calculate metabolic water. I used calculations proposed by Fox et al. (2000) to model free water requirements and modified the equations to reflect increased requirements for lactation. Diet analysis revealed that pronghorn used between 67% and 99% forbs and suggested fair range conditions. Preformed water was not significantly different between night and day. Night feeding appeared to be of marginal advantage, providing an average potential 9% preformed water increase in 2008, and 3% in 2009. The model indicated that neither male nor female pronghorn could meet their water requirements from preformed and metabolic water during any time period, season or year. The average free water requirements for females ranged from 0.67 L/animal/day (SE 0.06) in March, 2008 to 3.12 L/animal/day (SE 0.02) in June, 2009. The model showed that American pronghorn on Perry Mesa require access to free water during biological stress periods.
ContributorsTluczek, Melanie (Author) / Miller, William H. (Thesis advisor) / Brown, David E. (Committee member) / Steele, Kelly (Committee member) / Arizona State University (Publisher)
Created2012
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Description
Persistent cooperation between unrelated conspecifics rarely occurs in mature eusocial insect societies. In this dissertation, I present evidence of non-kin cooperation in the Nearctic honey ant Myrmecocystus mendax. Using microsatellite markers, I show that mature colonies in the Sierra Ancha Mountain of central Arizona contain multiple unrelated matrilines, an observation

Persistent cooperation between unrelated conspecifics rarely occurs in mature eusocial insect societies. In this dissertation, I present evidence of non-kin cooperation in the Nearctic honey ant Myrmecocystus mendax. Using microsatellite markers, I show that mature colonies in the Sierra Ancha Mountain of central Arizona contain multiple unrelated matrilines, an observation that is consistent with primary polygyny. In contrast, similar analyses suggest that colonies in the Chiricahua Mountains of southeastern Arizona are primarily monogynous. These interpretations are consistent with field and laboratory observations. Whereas cooperative colony founding was observed frequently among groups of Sierra Ancha foundresses, founding in the Chiricahua population was restricted to individual foundresses. Furthermore, Sierra Ancha foundresses successfully established incipient laboratory colonies without undergoing queen culling following emergence of the first workers. Multi-queen laboratory Sierra Ancha colonies also produced more workers and repletes than haplometrotic colonies, and when brood raiding was induced between colonies, queens of those with more workers had a higher survival probability.

Microsatellite analyses of additional locations within the M. mendax range suggest that polygyny is also present in some other populations, especially in central-northern Arizona, albeit at lower frequencies than that in the Sierra Anchas. In addition, analyses of multiple types of genetic data, including microsatellites, the mitochondrial barcoding region, and over 2000 nuclear ultra-conserved elements indicate that M. mendax populations within the southwestern U.S. and northwestern Mexico are geographically structured, with strong support for the existence of two or more divergent clades as well as isolation-by-distance within clades. This structure is further shown to correlate with variation in queen number and hair length, a diagnostic taxonomic feature used to distinguish honey ant species.

Together, these findings suggest that regional ecological pressures (e.g. colony density , climate) may have acted on colony founding and social strategy to select for increasing workforce size and, along with genetic drift, have driven geographically isolated M. mendax populations to differentiate genetically and morphologically. The presence of colony fusion in the laboratory and life history traits in honey ant that are influenced by colony size, including repletism, brood raiding, and tournament, support this evolutionary scenario.
ContributorsEriksson, Ti (Author) / Gadau, Jürgen (Thesis advisor) / Taylor, Jay (Thesis advisor) / Fewell, Jennifer (Committee member) / Hӧlldobler, Bert (Committee member) / Johnson, Robert (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Human-inhabited or -disturbed areas pose many unique challenges for wildlife, including increased human exposure, novel challenges, such as finding food or nesting sites in novel structures, anthropogenic noises, and novel predators. Animals inhabiting these environments must adapt to such changes by learning to exploit new resources and avoid danger. To

Human-inhabited or -disturbed areas pose many unique challenges for wildlife, including increased human exposure, novel challenges, such as finding food or nesting sites in novel structures, anthropogenic noises, and novel predators. Animals inhabiting these environments must adapt to such changes by learning to exploit new resources and avoid danger. To my knowledge no study has comprehensively assessed behavioral reactions of urban and rural populations to numerous novel environmental stimuli. I tested behavioral responses of urban, suburban, and rural house finches (Haemorhous mexicanus) to novel stimuli (e.g. objects, noises, food), to presentation of a native predator model (Accipiter striatus) and a human, and to two problem-solving challenges (escaping confinement and food-finding). Although I found few population-level differences in behavioral responses to novel objects, environment, and food, I found compelling differences in how finches from different sites responded to novel noise. When played a novel sound (whale call or ship horn), urban and suburban house finches approached their food source more quickly and spent more time on it than rural birds, and urban and suburban birds were more active during the whale-noise presentation. In addition, while there were no differences in response to the native predator, rural birds showed higher levels of stress behaviors when presented with a human. When I replicated this study in juveniles, I found that exposure to humans during development more accurately predicted behavioral differences than capture site. Finally, I found that urban birds were better at solving an escape problem, whereas rural birds were better at solving a food-finding challenge. These results indicate that not all anthropogenic changes affect animal populations equally and that determining the aversive natural-history conditions and challenges of taxa may help urban ecologists better understand the direction and degree to which animals respond to human-induced rapid environmental alterations.
ContributorsWeaver, Melinda (Author) / McGraw, Kevin J. (Thesis advisor) / Rutowski, Ronald (Committee member) / Pratt, Stephen (Committee member) / Bateman, Heather (Committee member) / Deviche, Pierre (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Is it possible to treat the mouth as a natural environment, and determine new methods to keep the microbiome in check? The need for biodiversity in health may suggest that every species carries out a specific function that is required to maintain equilibrium and homeostasis within the oral cavity. Furthermore,

Is it possible to treat the mouth as a natural environment, and determine new methods to keep the microbiome in check? The need for biodiversity in health may suggest that every species carries out a specific function that is required to maintain equilibrium and homeostasis within the oral cavity. Furthermore, the relationship between the microbiome and its host is mutually beneficial because the host is providing microbes with an environment in which they can flourish and, in turn, keep their host healthy. Reviewing examples of larger scale environmental shifts could provide a window by which scientists can make hypotheses. Certain medications and healthcare treatments have been proven to cause xerostomia. This disorder is characterized by a dry mouth, and known to be associated with a change in the composition, and reduction, of saliva. Two case studies performed by Bardow et al, and Leal et al, tested and studied the relationships of certain medications and confirmed their side effects on the salivary glands [2,3]. Their results confirmed a relationship between specific medicines, and the correlating complaints of xerostomia. In addition, Vissink et al conducted case studies that helped to further identify how radiotherapy causes hyposalivation of the salivary glands [4]. Specifically patients that have been diagnosed with oral cancer, and are treated by radiotherapy, have been diagnosed with xerostomia. As stated prior, studies have shown that patients having an ecologically balanced and diverse microbiome tend to have healthier mouths. The oral cavity is like any biome, consisting of commensalism within itself and mutualism with its host. Due to the decreased salivary output, caused by xerostomia, increased parasitic bacteria build up within the oral cavity thus causing dental disease. Every human body contains a personalized microbiome that is essential to maintaining health but capable of eliciting disease. The Human Oral Microbiomics Database (HOMD) is a set of reference 16S rRNA gene sequences. These are then used to define individual human oral taxa. By conducting metagenomic experiments at the molecular and cellular level, scientists can identify and label micro species that inhabit the mouth during parasitic outbreaks or a shifting of the microbiome. Because the HOMD is incomplete, so is our ability to cure, or prevent, oral disease. The purpose of the thesis is to research what is known about xerostomia and its effects on the complex microbiome of the oral cavity. It is important that researchers determine whether this particular perspective is worth considering. In addition, the goal is to create novel experiments for treatment and prevention of dental diseases.
ContributorsHalcomb, Michael Jordan (Author) / Chen, Qiang (Thesis director) / Steele, Kelly (Committee member) / Barrett, The Honors College (Contributor) / College of Letters and Sciences (Contributor)
Created2015-05
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Description
Abstract
Purpose—Use a framework of genetic knowledge to investigate the association between the genotypes of various genes with phenotypes, specifically the traits of elite athletes, in order to establish a personal opinion on their relevance to athletic performance.
Methods—Assemble and analyze selected published scientific studies on genotype and athletic performance

Abstract
Purpose—Use a framework of genetic knowledge to investigate the association between the genotypes of various genes with phenotypes, specifically the traits of elite athletes, in order to establish a personal opinion on their relevance to athletic performance.
Methods—Assemble and analyze selected published scientific studies on genotype and athletic performance and lastly to formulate a personal opinion on the value of genetic testing of athletes. ACTN3, ACE, MSTN, and apoE were the genes selected for analyses.
Results—Two genes, ACTN3 and ACE, showed a significant relationship of genotype to phenotypic traits related to athletic performance. ApoE did not demonstrate a phenotypic association with athletic performance, however it showed a correlation with injury susceptibility leading to traumatic brain injury (TBI). MSTN did not show a phenotypic association with athletic performance.
Conclusion—When considering the multifactorial nature of athletics, each sport must be investigated individually due to the different individual requirements. ACTN3 and ACE are the most widely studied genes, therefore, considerable data on their relevance to athletic performance was easily obtained and supported a relationship between genotype and athletic performance.
ContributorsMinto, Jordan Taylor- Lloyd (Author) / Steele, Kelly (Thesis director) / Penton, C. Ryan (Committee member) / College of Integrative Sciences and Arts (Contributor) / Barrett, The Honors College (Contributor)
Created2017-05
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Description
Although mimetic animal coloration has been studied since Darwin's time, many questions on the efficacy, evolution, and function of mimicry remain unanswered. Müller (1879) hypothesized that unpalatable individuals converge on the same conspicuous coloration to reduce predation. However, there are many cases where closely related, unpalatable species have diverged from

Although mimetic animal coloration has been studied since Darwin's time, many questions on the efficacy, evolution, and function of mimicry remain unanswered. Müller (1879) hypothesized that unpalatable individuals converge on the same conspicuous coloration to reduce predation. However, there are many cases where closely related, unpalatable species have diverged from a shared conspicuous pattern. What selection pressures have led to divergence in warning colors? Environmental factors such as ambient light have been hypothesized to affect signal transmission and efficacy in animals. Using two mimetic pairs of Heliconius butterflies, Postman and Blue-white, I tested the hypothesis that animals with divergent mimetic colors segregate by light environment to maximize conspicuousness of the aposematic warning signal under their particular environmental conditions. Each mimetic pair was found in a light environment that differed in brightness and spectral composition, which affected visual conspicuousness differently depending on mimetic color patch. I then used plasticine models in the field to test the hypothesis that mimics had higher survival in the habitat where they occurred. Although predation rates differed between the two habitats, there was no interactive effect of species by habitat type. Through choice experiments, I demonstrated that mimetic individuals preferred to spend time in the light environment where they were most often found and that their absolute visual sensitivity corresponds to the ambient lighting of their respective environment. Eye morphology was then studied to determine if differences in total corneal surface area and/or facet diameters explained the differences in visual sensitivities, but the differences found in Heliconius eye morphology did not match predictions based upon visual sensitivity. To further understand how eye morphology varies with light environments, I studied many tropical butterflies from open and closed habitats to reveal that forest understory butterflies have larger facets compared to butterflies occupying open habitats. Lastly, I tested avian perception of mimicry in a putative Heliconius mimetic assemblage and show that the perceived mimetic resemblance depends upon visual system. This dissertation reveals the importance of light environments on mimicry, coloration, behavior and visual systems of tropical butterflies.
ContributorsSeymoure, Brett M (Author) / Rutowski, Ronald L (Thesis advisor) / McGraw, Kevin J. (Thesis advisor) / McMillan, W. Owen (Committee member) / Pratt, Stephen (Committee member) / Gadau, Jürgen (Committee member) / Arizona State University (Publisher)
Created2016