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Division of labor, whereby different group members perform different functions, is a fundamental attribute of sociality. It appears across social systems, from simple cooperative groups to complex eusocial colonies. A core challenge in sociobiology is to explain how patterns of collective organization are generated. Theoretical models propose that division of

Division of labor, whereby different group members perform different functions, is a fundamental attribute of sociality. It appears across social systems, from simple cooperative groups to complex eusocial colonies. A core challenge in sociobiology is to explain how patterns of collective organization are generated. Theoretical models propose that division of labor self-organizes, or emerges, from interactions among group members and the environment; division of labor is also predicted to scale positively with group size. I empirically investigated the emergence and scaling of division of labor in evolutionarily incipient groups of sweat bees and in eusocial colonies of harvester ants. To test whether division of labor is an emergent property of group living during early social evolution, I created de novo communal groups of the normally solitary sweat bee Lasioglossum (Ctenonomia) NDA-1. A division of labor repeatedly arose between nest excavation and guarding tasks; results were consistent with hypothesized effects of spatial organization and intrinsic behavioral variability. Moreover, an experimental increase in group size spontaneously promoted higher task specialization and division of labor. Next, I examined the influence of colony size on division of labor in larger, more integrated colonies of the harvester ant Pogonomyrmex californicus. Division of labor scaled positively with colony size in two contexts: during early colony ontogeny, as colonies grew from tens to hundreds of workers, and among same-aged colonies that varied naturally in size. However, manipulation of colony size did not elicit a short-term response, suggesting that the scaling of division of labor in P. californicus colonies is a product of functional integration and underlying developmental processes, rather than a purely emergent epiphenomenon. This research provides novel insights into the organization of work in insect societies, and raises broader questions about the role of size in sociobiology.
ContributorsHolbrook, Carter Tate (Author) / Fewell, Jennifer H (Thesis advisor) / Gadau, Jürgen (Committee member) / Harrison, Jon F. (Committee member) / Hölldobler, Berthold (Committee member) / Johnson, Robert A. (Committee member) / Arizona State University (Publisher)
Created2011
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Description
Social insect colonies exhibit striking diversity in social organization. Included in this overwhelming variation in structure are differences in colony queen number. The number of queens per colony varies both intra- and interspecifically and has major impacts on the social dynamics of a colony and the fitness of its members.

Social insect colonies exhibit striking diversity in social organization. Included in this overwhelming variation in structure are differences in colony queen number. The number of queens per colony varies both intra- and interspecifically and has major impacts on the social dynamics of a colony and the fitness of its members. To understand the evolutionary transition from single to multi-queen colonies, I examined a species which exhibits variation both in mode of colony founding and in the queen number of mature colonies. The California harvester ant Pogonomyrmex californicus exhibits both variation in the number of queens that begin a colony (metrosis) and in the number of queens in adult colonies (gyny). Throughout most of its range, colonies begin with one queen (haplometrosis) but in some populations multiple queens cooperate to initiate colonies (pleometrosis). I present results that confirm co-foundresses are unrelated. I also map the geographic occurrence of pleometrotic populations and show that the phenomenon appears to be localized in southern California and Northern Baja California. Additionally, I provide genetic evidence that pleometrosis leads to primary polygyny (polygyny developing from pleometrosis) a phenomenon which has received little attention and is poorly understood. Phylogenetic and haplotype analyses utilizing mitochondrial markers reveal that populations of both behavioral types in California are closely related and have low mitochondrial diversity. Nuclear markers however, indicate strong barriers to gene flow between focal populations. I also show that intrinsic differences in queen behavior lead to the two types of populations observed. Even though populations exhibit strong tendencies on average toward haplo- or pleometrosis, within population variation exists among queens for behaviors relevant to metrosis and gyny. These results are important in understanding the dynamics and evolutionary history of a distinct form of cooperation among unrelated social insects. They also help to understand the dynamics of intraspecific variation and the conflicting forces of local adaptation and gene flow.
ContributorsOverson, Rick P (Author) / Gadau, Jürgen (Thesis advisor) / Fewell, Jennifer H (Committee member) / Hölldobler, Bert (Committee member) / Johnson, Robert A. (Committee member) / Liebig, Jürgen (Committee member) / Arizona State University (Publisher)
Created2011
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Description

Lyme disease is a common tick-borne illness caused by the Gram-negative bacterium Borrelia burgdorferi. An outer membrane protein of Borrelia burgdorferi, P66, has been suggested as a possible target for Lyme disease treatments. However, a lack of structural information available for P66 has hindered attempts to design medications to target

Lyme disease is a common tick-borne illness caused by the Gram-negative bacterium Borrelia burgdorferi. An outer membrane protein of Borrelia burgdorferi, P66, has been suggested as a possible target for Lyme disease treatments. However, a lack of structural information available for P66 has hindered attempts to design medications to target the protein. Therefore, this study attempted to find methods for expressing and purifying P66 in quantities that can be used for structural studies. It was found that by using the PelB signal sequence, His-tagged P66 could be directed to the outer membrane of Escherichia coli, as confirmed by an anti-His Western blot. Further attempts to optimize P66 expression in the outer membrane were made, pending verification via Western blotting. The ability to direct P66 to the outer membrane using the PelB signal sequence is a promising first step in determining the overall structure of P66, but further work is needed before P66 is ready for large-scale purification for structural studies.

ContributorsRamirez, Christopher Nicholas (Author) / Fromme, Petra (Thesis director) / Hansen, Debra (Committee member) / Department of Physics (Contributor) / School of Molecular Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2021-05
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Description
All multicellular organisms are susceptible to developing cancer, but some organisms have varying sensitivities to the disease. One such organism is the Trichoplax adhaerens which has no documented case of cancer development. T. adhaerens cancer resistance was studied by observing physiological and morphological changes of the organism after radiation treatment.

All multicellular organisms are susceptible to developing cancer, but some organisms have varying sensitivities to the disease. One such organism is the Trichoplax adhaerens which has no documented case of cancer development. T. adhaerens cancer resistance was studied by observing physiological and morphological changes of the organism after radiation treatment. Preliminary experiments suggested that this organism is able to survive exposure to 160 gray radiation treatment almost as well as untreated organisms. The T. adhaerens have two genes, TriadG6402 and TriadG5479, similar to the human genes TP53 and MDM2 respectively. TP53 and MDM2 are the two main genes associated with apoptosis in humans: an important cell regulatory checkpoint involved in cancer prevention. PCR analysis, done after radiation treatment, showed an overexpression of the ortholog gene MDM2 in the T. adhaerens. This may suggest that T. adhaerens block apoptosis from occurring and that their ortholog gene is involved in DNA repair. It is significant to study the gene expression of TriadG6402 and TriadG54791 in T. adhaerens because these genes are well conserved in humans. Future studies of these genes in the T. adhaerens can be used to understand the evolution of the function of these genes in more complex organisms and be used for human cancer prevention.
ContributorsKulkarni, Arathi (Author) / Fortunato, Angelo (Thesis director) / Maley, Carlo (Committee member) / Department of Economics (Contributor) / School of Molecular Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2019-05
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Description
Enzyme Replacement Therapy (ERT) is a treatment often used for patients with disorders that affect the production of various enzymes within the body, such as Cystic Fibrosis and Fabry Disease. ERT involves the use of artificially-produced enzymes, which can be derived from humans, pigs, and bacteria. Generally, enzymes derived from

Enzyme Replacement Therapy (ERT) is a treatment often used for patients with disorders that affect the production of various enzymes within the body, such as Cystic Fibrosis and Fabry Disease. ERT involves the use of artificially-produced enzymes, which can be derived from humans, pigs, and bacteria. Generally, enzymes derived from porcine and bacterial sources are much less expensive and more accessible than those derived from a human source. This, and the ethical implications that porcine enzymes carry, make the decision of choosing treatment simple to some and complex to others. Ethically, human-derived enzymes are often considered more ethical, while not conflicting with religious beliefs and practices as porcine-derived enzymes do.
In order to further compare porcine and human-derived enzymes, a determination of the enzyme effectiveness was done via digestion simulation. The digestion for both the human and porcine-derived enzymes consisted of three steps: oral, gastric, and intestinal. After the digestion, the absorbance for each enzyme class as well as a dilution curve of the formula used was read and recorded. Using the standard dilution curve and the absorbance values for each unknown, the formula and thus enzyme concentration that was lost through the reaction was able to be calculated.
The effectiveness of both the human and porcine enzymes, determined by the percent of formula lost, was 18.2% and 19.7%, respectively, with an error of 0.6% from the spectrophotometer, and an error of about 10% from the scale used for measuring the enzymes. This error was likely due to the small mass required of the enzymes and can be prevented in the future by performing the experiment at a larger scale.
ContributorsBlevins, Brianna R (Author) / Martin, Thomas (Thesis director) / McILwraith, Heide (Committee member) / College of Integrative Sciences and Arts (Contributor) / School of Molecular Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2020-05
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Description
Bee communities form the keystone of many ecosystems through their pollination services. They are dynamic and often subject to significant changes due to several different factors such as climate, urban development, and other anthropogenic disturbances. As a result, the world has been experiencing a decline in bee diversity and abundance,

Bee communities form the keystone of many ecosystems through their pollination services. They are dynamic and often subject to significant changes due to several different factors such as climate, urban development, and other anthropogenic disturbances. As a result, the world has been experiencing a decline in bee diversity and abundance, which can have detrimental effects in the ecosystems they inhabit. One of the largest factors that impacts bees in today's world is the rapid urbanization of our planet, and it impacts the bee community in mixed ways. Not very much is understood about the bee communities that exist in urban habitats, but as urbanization is inevitably going to continue, knowledge on bee communities will need to strengthen. This study aims to determine the levels of variance in bee communities, considering multiple variables that bee communities can differ in. The following three questions are posed: do bee communities that are spatially separated differ significantly? Do bee communities that are separated by seasons differ significantly? Do bee communities that are separated temporally (by year, interannually) differ significantly? The procedure to conduct this experiment consists of netting and trapping bees at two sites at various times using the same methods. The data is then statistically analyzed for differences in abundance, richness, diversity, and species composition. After performing the various statistical analyses, it has been discovered that bee communities that are spatially separated, seasonally separated, or interannually separated do not differ significantly when it comes to abundance and richness. Spatially separated bee communities and interannually separated bee communities show a moderate level of dissimilarity in their species composition, while seasonally separated bee communities show a greater level of dissimilarity in species composition. Finally, seasonally separated bee communities demonstrate the greatest disparity of bee diversity, while interannually separated bee communities show the least disparity of bee diversity. This study was conducted over the time span of two years, and while the levels of variance of an urban area between these variables were determined, further variance studies of greater length or larger areas should be conducted to increase the currently limited knowledge of bee communities in urban areas. Additional studies on precipitation amounts and their effects on bee communities should be conducted, and studies from other regions should be taken into consideration while attempting to understand what is likely the most environmentally significant group of insects.
ContributorsPhan, James Thien (Author) / Sweat, Ken (Thesis director) / Foltz-Sweat, Jennifer (Committee member) / School of Music (Contributor) / School of Molecular Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2017-05
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Description
The college textbook is the most commonly required component of almost any college course, regardless of a student's academic discipline. Professors often expect students to have access to the textbook and to use it to complete assigned readings. Textbooks often contain features that are designed to facilitate active reading, or

The college textbook is the most commonly required component of almost any college course, regardless of a student's academic discipline. Professors often expect students to have access to the textbook and to use it to complete assigned readings. Textbooks often contain features that are designed to facilitate active reading, or critical engagement with the information being read, to enhance learning of the material. However, students often do not read or prioritize reading the textbook. Students who do read, tend not to read the textbook as intended or use many of the features designed to promote active reading and enhanced learning of the material. Educational studies of textbooks tend to focus on aspects related to topics more relevant to publishers or professors with less research on aspects of the textbook applicable to students at the college level. The purpose of this study is to evaluate students' textbook use and their attitudes toward the textbook in an introductory biology course. Results of this study indicate students hold positive attitudes toward their textbook in an introductory biology course and majority of students do not use components meant to facilitate active learning. Although students report completing assigned readings, students may actually be reading select portions of what is assigned in using the textbook to prepare for exams. These results suggest that students may only be using their textbook to enhance their understanding of materials they expect to be tested on. The findings of this study help to understand the role of the textbook from the perspective of the student and provide insight for improving textbook design and use in science courses at the college level.
ContributorsRudolph, Alexia Marion (Author) / Vanmali, Binaben (Thesis director) / Chen, Ying-Chih (Committee member) / Yoho, Rachel (Committee member) / Department of Psychology (Contributor) / School of Molecular Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2016-05
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Description
Understanding why animals form social groups is a fundamental aim of sociobiology. To date, the field has been dominated by studies of kin groups, which have emphasized indirect fitness benefits as key drivers of grouping among relatives. Nevertheless, many animal groups are comprised of unrelated individuals. These cases provide unique

Understanding why animals form social groups is a fundamental aim of sociobiology. To date, the field has been dominated by studies of kin groups, which have emphasized indirect fitness benefits as key drivers of grouping among relatives. Nevertheless, many animal groups are comprised of unrelated individuals. These cases provide unique opportunities to illuminate drivers of social evolution beyond indirect fitness, especially ecological factors. This dissertation combines behavioral, physiological, and ecological approaches to explore the conditions that favor group formation among non-kin, using as a model the facultatively social carpenter bee, Xylocopa sonorina. Using behavioral and genetic techniques, I found that nestmates in this species are often unrelated, and that non-kin groups form following extensive inter-nest migration.Group living may arise as a strategy to mitigate constraints on available breeding space. To test the hypothesis that nest construction is prohibitively costly for carpenter bees, I measured metabolic rates of excavating bees and used imaging techniques to quantify nest volumes. From these measurements, I found that nest construction is highly energetically costly, and that bees who inherit nests through social queuing experience substantial energetic savings. These costs are exacerbated by limitations on the reuse of existing nests. Using repeated CT scans of nesting logs, I examined changes in nest architecture over time and found that repeatedly inherited tunnels become indefensible to intruders, and are subsequently abandoned. Together, these factors underlie intense competition over available breeding space. The imaging analysis of nesting logs additionally revealed strong seasonal effects on social strategy, with social nesting dominating during winter. To test the hypothesis that winter social nesting arises from intrinsic physiological advantages of grouping, I experimentally manipulated social strategy in overwintering bees. I found that social bees conserve heat and body mass better than solitary bees, suggesting fitness benefits to grouping in cold, resource-scarce conditions. Together, these results suggest that grouping in X. sonorina arises from dynamic strategies to maximize direct fitness in response to harsh and/or competitive conditions. These studies provide empirical insights into the ecological conditions that favor non-kin grouping, and emphasize the importance of ecology in shaping sociality at its evolutionary origins.
ContributorsOstwald, Madeleine (Author) / Fewell, Jennifer H (Thesis advisor) / Amdam, Gro (Committee member) / Harrison, Jon (Committee member) / Pratt, Stephen (Committee member) / Kapheim, Karen (Committee member) / Arizona State University (Publisher)
Created2022
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Description
Understanding how and why animals choose what to eat is one of the fundamental goals of nutritional and behavioral biology. This question can be scaled to animals that live in social groups, including eusocial insects. One of the factors that plays an important role in foraging decisions is the prevalence

Understanding how and why animals choose what to eat is one of the fundamental goals of nutritional and behavioral biology. This question can be scaled to animals that live in social groups, including eusocial insects. One of the factors that plays an important role in foraging decisions is the prevalence of specific nutrients and their relative balance. This dissertation explores the role of relative nutrient content in the food selection decisions of a species that is eusocial and also agricultural, the desert leafcutter ant Acromyrmex versicolor. A dietary choice assay, in which the relative amount of protein and carbohydrates in the available diets was varied, demonstrated that A. versicolor colonies regulate relative collection of protein and carbohydrates. Tracking the foraging behavior of individual workers revelaed that foragers vary in their relative collection of experimental diets and in their foraging frequency, but that there is no relationship between these key factors of foraging behavior. The high proportion of carbohydrates preferred by lab colonies suggests that they forage to nutritionally support the fungus rather than brood and workers. To test this, the relative amounts of 1) fungus, and 2) brood (larvae) was manipulated and foraging response was measured. Changing the amount of brood had no effect on foraging. Although decreasing the size of fungus gardens did not change relative P:C collection, it produced significant increases in caloric intake, supporting the assertion that the fungus is the main driver of colony nutrient regulation. The nutritional content of naturally harvested forage material collected from field colonies was measured, as was recruitment to experimental diets with varying relative macronutrient content. Field results confirmed a strong colony preference for high carbohydrate diets. They also indicated that this species may, at times, be limited in its ability to collect sufficiently high levels of carbohydrates to meet optimal intake. This dissertation provides important insights about fundamental aspects of leafcutter ant biology and extends our understanding of the role of relative nutrient content in foraging decisions to systems that span multiple trophic levels.
ContributorsSmith, Nathan Edward (Author) / Fewell, Jennifer H (Thesis advisor) / Harrison, Jon F (Committee member) / Pavlic, Ted (Committee member) / Cease, Arianne (Committee member) / Hoelldobler, Bert (Committee member) / Arizona State University (Publisher)
Created2023
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Description
In many social groups, reproduction is shared between group members, whocompete for position in the social hierarchy for reproductive dominance. This reproductive conflict can lead to different means of enforcing reproductive differences, such as dominance displays or limited control of social hierarchy through antagonistic encounters. In eusocial insects, archetypal colonies contain a single,

In many social groups, reproduction is shared between group members, whocompete for position in the social hierarchy for reproductive dominance. This reproductive conflict can lead to different means of enforcing reproductive differences, such as dominance displays or limited control of social hierarchy through antagonistic encounters. In eusocial insects, archetypal colonies contain a single, singly-mated fertile queen, such that no reproductive conflict exists within a colony. However, many eusocial insects deviate from this archetype and have multiply-mated queens (polyandry), multiple queens in a single colony (polygyny), or both. In these cases, reproductive conflict exists between the matrilines and patrilines represented in a colony, specifically over the production of sexual offspring. A possible outcome of reproductive conflict may be the emergence of cheating lineages, which favor the production of sexual offspring, taking advantage of the worker force produced by nestmate queens and/or patrilines. In extreme examples, inquiline social parasites may be an evolutionary consequence of reproductive conflict between nestmate queens. Inquiline social parasitism is a type of social parasitism that is usually defined by a partial or total loss of the worker caste, and the “infiltration” of host colonies to take advantage of the host worker force for reproduction. It has been hypothesized that these inquiline social parasites evolve through the speciation of cheating queen lineages from within their incipient host species. This “intra- specific” origin model involves a foundational hypothesis that the common ancestor of host and parasite (and thus, putatively, the host at the time of speciation) should be functionally polygynous, and that parasitism evolves as a “resolution” of reproductive conflict in colonies. In this dissertation, I investigate the hypothesized role of polygyny in the evolution of inquiline social parasites. I use molecular ecology and statistical approaches to validate the role of polygyny in the evolution of some inquiline social parasites. I further discuss potential mechanisms for the evolution and speciation of social parasites, and discuss future directions to elucidate these mechanisms.
ContributorsDahan, Romain Arvid (Author) / Rabeling, Christian (Thesis advisor) / Amdam, Gro V (Committee member) / Fewell, Jennifer H (Committee member) / Pratt, Stephen C (Committee member) / Rüppell, Olav (Committee member) / Arizona State University (Publisher)
Created2021