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Description
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
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
At the heart of every eusocial insect colony is a reproductive division of labor. This division can emerge through dominance interactions at the adult stage or through the production of distinct queen and worker castes at the larval stage. In both cases, this division depends on plasticity within an individual

At the heart of every eusocial insect colony is a reproductive division of labor. This division can emerge through dominance interactions at the adult stage or through the production of distinct queen and worker castes at the larval stage. In both cases, this division depends on plasticity within an individual to develop reproductive characteristics or serve as a worker. In order to gain insight into the evolution of reproductive plasticity in the social insects, I investigated caste determination and dominance in the ant Harpegnathos saltator, a species that retains a number of ancestral characteristics. Treatment of worker larvae with a juvenile hormone (JH) analog induced late-instar larvae to develop as queens. At the colony level, workers must have a mechanism to regulate larval development to prevent queens from developing out of season. I identified a new behavior in H. saltator where workers bite larvae to inhibit queen determination. Workers could identify larval caste based on a chemical signal specific to queen-destined larvae, and the production of this signal was directly linked to increased JH levels. This association provides a connection between the physiological factors that induce queen development and the production of a caste-specific larval signal. In addition to caste determination at the larval stage, adult workers of H. saltator compete to establish a reproductive hierarchy. Unlike other social insects, dominance in H. saltator was not related to differences in JH or ecdysteroid levels. Instead, changes in brain levels of biogenic amines, particularly dopamine, were correlated with dominance and reproductive status. Receptor genes for dopamine were expressed in both the brain and ovaries of H. saltator, and this suggests that dopamine may coordinate changes in behavior at the neurological level with ovarian status. Together, these studies build on our understanding of reproductive plasticity in social insects and provide insight into the evolution of a reproductive division of labor.
ContributorsPenick, Clint A (Author) / Liebig, Jürgen (Thesis advisor) / Brent, Colin (Committee member) / Gadau, Jürgen (Committee member) / Hölldobler, Bert (Committee member) / Rutowski, Ron (Committee member) / Arizona State University (Publisher)
Created2012
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Description
Introduction: Human papillomavirus (HPV) infection is seen in up to 90% of cases of cervical cancer, the third leading cancer cause of death in women. Current HPV screening focuses on only two HPV types and covers roughly 75% of HPV-associated cervical cancers. A protein based assay to test for antibody

Introduction: Human papillomavirus (HPV) infection is seen in up to 90% of cases of cervical cancer, the third leading cancer cause of death in women. Current HPV screening focuses on only two HPV types and covers roughly 75% of HPV-associated cervical cancers. A protein based assay to test for antibody biomarkers against 98 HPV antigens from both high and low risk types could provide an inexpensive and reliable method to screen for patients at risk of developing invasive cervical cancer. Methods: 98 codon optimized, commercially produced HPV genes were cloned into the pANT7_cGST vector, amplified in a bacterial host, and purified for mammalian expression using in vitro transcription/translation (IVTT) in a luminescence-based RAPID ELISA (RELISA) assay. Monoclonal antibodies were used to determine immune cross-reactivity between phylogenetically similar antigens. Lastly, several protein characteristics were examined to determine if they correlated with protein expression. Results: All genes were successfully moved into the destination vector and 86 of the 98 genes (88%) expressed protein at an adequate level. A difference was noted in expression by gene across HPV types but no correlation was found between protein size, pI, or aliphatic index and expression. Discussion: Further testing is needed to express the remaining 12 HPV genes. Once all genes have been successfully expressed and purified at high concentrations, DNA will be printed on microscope slides to create a protein microarray. This microarray will be used to screen HPV-positive patient sera for antibody biomarkers that may be indicative of cervical cancer and precancerous cervical neoplasias.
ContributorsMeshay, Ian Matthew (Author) / Anderson, Karen (Thesis director) / Magee, Mitch (Committee member) / Katchman, Benjamin (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor)
Created2015-05
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Description
Adaptive therapy utilizes competitive interactions between resistant and sensitive cells by keeping some sensitive cells to control tumor burden with the aim of increasing overall survival and time to progression. The use of adaptive therapy to treat breast cancer, ovarian cancer, and pancreatic cancer in preclinical models has shown significant

Adaptive therapy utilizes competitive interactions between resistant and sensitive cells by keeping some sensitive cells to control tumor burden with the aim of increasing overall survival and time to progression. The use of adaptive therapy to treat breast cancer, ovarian cancer, and pancreatic cancer in preclinical models has shown significant results in controlling tumor growth. The adaptive therapy model comes from the integrated pest management agricultural strategy, predator prey model, and the unique intra- and inter-tumor heterogeneity of tumors. The purpose of this thesis is to analyze and compare gemcitabine dose response on hormone refractory breast cancer cells retrieved from mice using an adaptive therapy strategy with standard therapy treatment. In this study, we compared intermittent (drug holiday) adaptive therapy with maximum tolerated dose therapy. The MCF7 resistant cell lines to both fulvestrant and palbociclib were injected into the mammary fat pads of 8 weeks old NOD/SCID gamma (NSG) mice which were then treated with gemcitabine. Tumor burden graphs were made to track tumor growth/decline during different treatments while Drug Dose Response (DDR) curves were made to test the sensitivity of the cell lines to the drug gemcitabine. The tumor burden graphs showed success in controlling the tumor burden with intermittent treatment. The DDR curves showed a positive result in using the adaptive therapy treatment method to treat mice with gemcitabine. Due to some fluctuating DDR results, the sensitivity of the cell lines to gemcitabine needs to be further studied by repeating the DDR experiment on the other mice cell lines for stronger results.
ContributorsConti, Aviona Christina (Author) / Maley, Carlo (Thesis advisor) / Blattman, Joseph (Committee member) / Anderson, Karen (Committee member) / Arizona State University (Publisher)
Created2022
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Description
Dominance behavior can regulate a division of labor in a group, such as that between reproductive and non-reproductive individuals. Manipulations of insect societies in a controlled environment can reveal how dominance behavior is regulated. Here, I examined how morphological caste, fecundity, group size, and age influence the expression of

Dominance behavior can regulate a division of labor in a group, such as that between reproductive and non-reproductive individuals. Manipulations of insect societies in a controlled environment can reveal how dominance behavior is regulated. Here, I examined how morphological caste, fecundity, group size, and age influence the expression of dominance behavior using the ponerine ant Harpegnathos saltator. All H. saltator females have the ability to reproduce. Only those with a queen morphology that enables dispersal, however, show putative sex pheromones. In contrast, those with a worker morphology normally express dominance behavior. To evaluate how worker-like dominance behavior and associated traits could be expressed in queens, I removed the wings from alate gynes, those with a queen morphology who had not yet mated or left the nest, making them dealate. Compared to gynes with attached wings, dealates frequently performed dominance behavior. In addition, only the dealates demonstrated worker-like ovarian activity in the presence of reproductive individuals, whereas gynes with wings produced sex pheromones exclusively. Therefore, the attachment of wings determines a gyne’s expression of worker-like dominance behavior and physiology. When the queen dies, workers establish a reproductive hierarchy among themselves by performing a combination of dominance behaviors. To understand how reproductive status depends on these interactions as well as a worker’s age, I measured the frequency of dominance behaviors in groups of different size composed of young and old workers. The number of workers who expressed dominance scaled with the size of the group, but younger ones were more likely to express dominance behavior and eventually become reproductive. Therefore, the predisposition of age integrates with a self-organized process to form this reproductive hierarchy. A social insect’s fecundity and fertility signal depends on social context because fecundity increases with colony size. To evaluate how a socially dependent signal regulates dominance behavior, I manipulated a reproductive worker’s social context. Reproductive workers with reduced fecundity and a less prominent fertility signal expressed more dominance behavior than those with a stronger fertility signal and higher fecundity. Therefore, dominance behavior reinforces rank to compensate for a weak signal, indicating how social context can feed back to influence the maintenance of dominance. Mechanisms that regulate H. saltator’s reproductive hierarchy can inform how the reproductive division of labor is regulated in other groups of animals.
ContributorsPyenson, Benjamin (Author) / Liebig, Jürgen (Thesis advisor) / Hölldobler, Bert (Committee member) / Fewell, Jennifer (Committee member) / Pratt, Stephen (Committee member) / Kang, Yun (Committee member) / Arizona State University (Publisher)
Created2022
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Description
The purpose of this paper is to create awareness around breast cancer risk factorsand screening methods. Five overarching intrinsic risk factors, including: the patient’s age at the time of diagnosis, race, familial susceptibility, and the role of natural hormone changes, and one extrinsic risk factor, dietary habits, were selected for consideration. Along with

The purpose of this paper is to create awareness around breast cancer risk factorsand screening methods. Five overarching intrinsic risk factors, including: the patient’s age at the time of diagnosis, race, familial susceptibility, and the role of natural hormone changes, and one extrinsic risk factor, dietary habits, were selected for consideration. Along with risk factors, four screening methods were taken into consideration. These included self-breast exams, mammograms, magnetic resonance imaging (MRI), and ultrasound. The recommendation of screening methods was then determined in relation to a women’s risk for breast cancer. Two categories of risk (average and high risk) were defined and the recommended screening methods were determined based on the risk. Overall, mammography was found to be a useful tool in both average and high risk women. For high risk women, mammography with MRI had a greater sensitivity and was able to detect more breast cancers. More research needs to be conducted on the efficacy of Breast MRI, Ultrasound, and breast self-exams as supplemental tools to mammography in both average and high-risk women
ContributorsTodd, Julia M (Author) / Compton, Carolyn (Thesis advisor) / Pepin, Susan (Committee member) / Anderson, Karen (Committee member) / Arizona State University (Publisher)
Created2023
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Description
The most abundantly studied societies, with the exception of humans, are those of the eusocial insects, which include all ants. Eusocial insect societies are typically composed of many dozens to millions of individuals, referred to as nestmates, which require some form of communication to maintain colony cohesion and coordinate the

The most abundantly studied societies, with the exception of humans, are those of the eusocial insects, which include all ants. Eusocial insect societies are typically composed of many dozens to millions of individuals, referred to as nestmates, which require some form of communication to maintain colony cohesion and coordinate the activities within them. Nestmate recognition is the process of distinguishing between nestmates and non-nestmates, and embodies the first line of defense for social insect colonies. In ants, nestmate recognition is widely thought to occur through olfactory cues found on the exterior surfaces of individuals. These cues, called cuticular hydrocarbons (CHCs), comprise the overwhelming majority of ant nestmate profiles and help maintain colony identity. In this dissertation, I investigate how nestmate recognition is influenced by evolutionary, ontogenetic, and environmental factors. First, I contributed to the sequencing and description of three ant genomes including the red harvester ant, Pogonomyrmex barbatus, presented in detail here. Next, I studied how variation in nestmate cues may be shaped through evolution by comparatively studying a family of genes involved in fatty acid and hydrocarbon biosynthesis, i.e., the acyl-CoA desaturases, across seven ant species in comparison with other social and solitary insects. Then, I tested how genetic, developmental, and social factors influence CHC profile variation in P. barbatus, through a three-part study. (1) I conducted a descriptive, correlative study of desaturase gene expression and CHC variation in P. barbatus workers and queens; (2) I explored how larger-scale genetic variation in the P. barbatus species complex influences CHC variation across two genetically isolated lineages (J1/J2 genetic caste determining lineages); and (3) I experimentally examined how CHC development is influenced by an individual’s social environment. In the final part of my work, I resolved discrepancies between previous findings of nestmate recognition behavior in P. barbatus by studying how factors of territorial experience, i.e., spatiotemporal relationships, affect aggressive behaviors among red harvester ant colonies. Through this research, I was able to identify promising methodological approaches and candidate genes, which both broadens our understanding of P. barbatus nestmate recognition systems and supports future functional genetic studies of CHCs in ants.
ContributorsCash, Elizabeth I (Author) / Gadau, Jürgen (Thesis advisor) / Liebig, Jürgen (Thesis advisor) / Fewell, Jennifer (Committee member) / Hölldobler, Berthold (Committee member) / Kusumi, Kenro (Committee member) / Arizona State University (Publisher)
Created2016
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Description
Measles is a contagious, vaccine-preventable disease that continues to be the leading

cause of death in children younger than the age of 5 years. While the introduction of the Measles, Mumps, and Rubella vaccine (MMR) has significantly decreased morbidity and mortality rates worldwide, vaccine coverage is highly variable across global regions.

Measles is a contagious, vaccine-preventable disease that continues to be the leading

cause of death in children younger than the age of 5 years. While the introduction of the Measles, Mumps, and Rubella vaccine (MMR) has significantly decreased morbidity and mortality rates worldwide, vaccine coverage is highly variable across global regions. Current diagnostic methods rely on enzyme immunoassays (EIA) to detect IgM or IgG Abs in serum. Commercially available Diamedix Immunosimplicity® Measles IgG test kit has been shown to have 91.1% sensitivity and 93.8% specificity, with a positive predictive value of 88.7% and a negative predictive value of 90.9% on the basis of a PRN titer of 120. There is an increasing need for rapid screening for measles specific immunity in outbreak settings. This study aims to develop a rapid molecular diagnostic assay to detect IgG reactive to three individual measles virus (MeV) proteins.

Measles virus (MeV) genes were subcloned into the pJFT7_nGST vector to generate N- terminal GST fusion proteins. Single MeV cistrons were expressed using in vitro transcription/translation (IVTT) with human cell lysate. Expression of GST-tagged proteins was measured with mouse anti-GST mAb and sheep anti-mouse IgG. Relative light units (RLUs) as luminescence was measured. Antibodies to MeV antigens were measured in 40 serum samples from healthy subjects.

Protein expression of three MeV genes of interest was measured in comparison with vector control and statistical significance was determined using the Student’s t-test (p<0.05). N expressed at the highest level with an average RLU value of 3.01 x 109 (p<0.001) and all proteins were expressed at least 50% greater than vector control (4.56 x 106 RLU). 36/40 serum samples had IgG to N (Ag:GST ratio>1.21), F (Ag:GST ratio>1.92), or H (Ag:GST ratio> 1.23).

These data indicate that the in vitro expression of MeV antigens, N, F, and H, were markedly improved by subcloning into pJFT7_nGST vector to generate N-terminal GST fusion proteins. The expression of single MeV genes N, F and H, are suitable antigens for serologic capture analysis of measles-specific antibodies. These preliminary data can be used to design a more intensive study to explore the possibilities of using these MeV antigens as a diagnostic marker.
ContributorsMushtaq, Zuena (Author) / Anderson, Karen (Thesis advisor) / Blattman, Joseph (Committee member) / Lake, Douglas (Committee member) / Arizona State University (Publisher)
Created2016