This collection includes most of the ASU Theses and Dissertations from 2011 to present. ASU Theses and Dissertations are available in downloadable PDF format; however, a small percentage of items are under embargo. Information about the dissertations/theses includes degree information, committee members, an abstract, supporting data or media.

In addition to the electronic theses found in the ASU Digital Repository, ASU Theses and Dissertations can be found in the ASU Library Catalog.

Dissertations and Theses granted by Arizona State University are archived and made available through a joint effort of the ASU Graduate College and the ASU Libraries. For more information or questions about this collection contact or visit the Digital Repository ETD Library Guide or contact the ASU Graduate College at gradformat@asu.edu.

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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
There is increasing evidence that ovarian status influcences behavioral phenotype in workers of the honey bee Apis mellifera. Honey bee workers demonstrate a complex division of labor. Young workers perform in-hive tasks (e.g. brood care), while older bees perform outside tasks (e.g. foraging for food). This age correlated division of

There is increasing evidence that ovarian status influcences behavioral phenotype in workers of the honey bee Apis mellifera. Honey bee workers demonstrate a complex division of labor. Young workers perform in-hive tasks (e.g. brood care), while older bees perform outside tasks (e.g. foraging for food). This age correlated division of labor is known as temporal polyethism. Foragers demonstrate further division of labor with some bees biasing collection towards protein (pollen) and others towards carbohydrates (nectar). The Reproductive Ground-plan Hypothesis proposes that the ovary plays a regulatory role in foraging division of labor. European honey bee workers that have been selectively bred to store larger amounts of pollen (High strain) also have a higher number of ovarioles per ovary than workers from strains bred to store less pollen (Low strain). High strain bees also initiate foraging earlier than Low strain bees. The relationship between ovariole number and foraging behavior is also observed in wild-type Apis mellifera and Apis cerana: pollen-biased foragers have more ovarioles than nectar-biased foragers. In my first study, I investigated the pre-foraging behavioral patterns of the High and Low strain bees. I found that High strain bees progress through the temporal polyethism at a faster rate than Low strain bees. To ensure that the observed relationship between the ovary and foraging bias is not due to associated separate genes for ovary size and foraging behavior, I investigated foraging behavior of African-European backcross bees. The backcross breeding program was designed to break potential gene associations. The results from this study demonstrated the relationship between the ovary and foraging behavior, supporting the proposed causal linkage between reproductive development and behavioral phenotype. The final study was designed to elucidate a regulatory mechanism that links ovariole number with sucrose sensitivity, and loading decisions. I measured ovariole number, sucrose sensitivity and sucrose solution load size using a rate-controlled sucrose delivery system. I found an interaction effect between ovariole number and sucrose sensitivity for sucrose solution load size. This suggests that the ovary impacts carbohydrate collection through modulation of sucrose sensitivity. Because nectar and pollen collection are not independent, this would also impact protein collection.
ContributorsSiegel, Adam J (Author) / Page, Jr., Robert E (Thesis advisor) / Hamilton, Andrew L. (Committee member) / Brent, Colin S (Committee member) / Amdam, Gro V (Committee member) / McGraw, Kevin J. (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
A notable feature of advanced eusocial insect groups is a division of labor within the sterile worker caste. However, the physiological aspects underlying the differentiation of behavioral phenotypes are poorly understood in one of the most successful social taxa, the ants. By starting to understand the foundations on which social

A notable feature of advanced eusocial insect groups is a division of labor within the sterile worker caste. However, the physiological aspects underlying the differentiation of behavioral phenotypes are poorly understood in one of the most successful social taxa, the ants. By starting to understand the foundations on which social behaviors are built, it also becomes possible to better evaluate hypothetical explanations regarding the mechanisms behind the evolution of insect eusociality, such as the argument that the reproductive regulatory infrastructure of solitary ancestors was co-opted and modified to produce distinct castes. This dissertation provides new information regarding the internal factors that could underlie the division of labor observed in both founding queens and workers of Pogonomyrmex californicus ants, and shows that changes in task performance are correlated with differences in reproductive physiology in both castes. In queens and workers, foraging behavior is linked to elevated levels of the reproductively-associated juvenile hormone (JH), and, in workers, this behavioral change is accompanied by depressed levels of ecdysteroid hormones. In both castes, the transition to foraging is also associated with reduced ovarian activity. Further investigation shows that queens remain behaviorally plastic, even after worker emergence, but the association between JH and behavioral bias remains the same, suggesting that this hormone is an important component of behavioral development in these ants. In addition to these reproductive factors, treatment with an inhibitor of the nutrient-sensing pathway Target of Rapamycin (TOR) also causes queens to become biased towards foraging, suggesting an additional sensory component that could play an important role in division of labor. Overall, this work provides novel identification of the possible regulators behind ant division of labor, and suggests how reproductive physiology could play an important role in the evolution and regulation of non-reproductive social behaviors.
ContributorsDolezal, Adam G (Author) / Amdam, Gro V (Thesis advisor) / Brent, Colin S. (Committee member) / Gadau, Juergen (Committee member) / Hoelldobler, Bert (Committee member) / Liebig, Juergen (Committee member) / Arizona State University (Publisher)
Created2012
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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
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
Ant colonies provide numerous opportunities to study communication systems that maintain the cohesion of eusocial groups. In many ant species, workers have retained their ovaries and the ability to produce male offspring; however, they generally refrain from producing their own sons when a fertile queen is present in the colony.

Ant colonies provide numerous opportunities to study communication systems that maintain the cohesion of eusocial groups. In many ant species, workers have retained their ovaries and the ability to produce male offspring; however, they generally refrain from producing their own sons when a fertile queen is present in the colony. Although mechanisms that facilitate the communication of the presence of a fertile queen to all members of the colony have been highly studied, those studies have often overlooked the added challenge faced by polydomous species, which divide their nests across as many as one hundred satellite nests resulting in workers potentially having infrequent contact with the queen. In these polydomous contexts, regulatory phenotypes must extend beyond the immediate spatial influence of the queen.

This work investigates mechanisms that can extend the spatial reach of fertility signaling and reproductive regulation in three polydomous ant species. In Novomessor cockerelli, the presence of larvae but not eggs is shown to inhibit worker reproduction. Then, in Camponotus floridanus, 3-methylheptacosane found on the queen cuticle and queen-laid eggs is verified as a releaser pheromone sufficient to disrupt normally occurring aggressive behavior toward foreign workers. Finally, the volatile and cuticular hydrocarbon pheromones present on the cuticle of Oecophylla smaragdina queens are shown to release strong attraction response by workers; when coupled with previous work, this result suggests that these chemicals may underly both the formation of a worker retinue around the queen as well as egg-located mechanisms of reproductive regulation in distant satellite nests. Whereas most previous studies have focused on the short-range role of hydrocarbons on the cuticle of the queen, these studies demonstrate that eusocial insects may employ longer range regulatory mechanisms. Both queen volatiles and distributed brood can extend the range of queen fertility signaling, and the use of larvae for fertility signaling suggest that feeding itself may be a non-chemical mechanism for reproductive regulation. Although trail laying in mass-recruiting ants is often used as an example of complex communication, reproductive regulation in ants may be a similarly complex example of insect communication, especially in the case of large, polydomous ant colonies.
ContributorsEbie, Jessica (Author) / Liebig, Jürgen (Thesis advisor) / Hölldobler, Bert (Thesis advisor) / Pratt, Stephen (Committee member) / Smith, Brian (Committee member) / Rutowski, Ronald (Committee member) / Arizona State University (Publisher)
Created2020
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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