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Of all the signals and cues that orchestrate the activities of a social insect colony, the reproductives' fertility pheromones are perhaps the most fundamental. These pheromones regulate reproductive division of labor, a defining characteristic of eusociality. Despite their critical role, reproductive fertility pheromones are not evenly expressed across the development

Of all the signals and cues that orchestrate the activities of a social insect colony, the reproductives' fertility pheromones are perhaps the most fundamental. These pheromones regulate reproductive division of labor, a defining characteristic of eusociality. Despite their critical role, reproductive fertility pheromones are not evenly expressed across the development of a social insect colony and may even be absent in the earliest colony stages. In the ant Camponotus floridanus, queens of incipient colonies do not produce the cuticular hydrocarbons that serve as fertility and egg-marking signals in this species. My dissertation investigates the consequences of the dramatic change in the quantity of these pheromones that occurs as the colony grows. C. floridanus workers from large, established colonies use egg surface hydrocarbons to discriminate among eggs. Eggs with surface hydrocarbons typical of eggs laid by established queens are nurtured, whereas eggs lacking these signals (i.e., eggs laid by workers and incipient queens) are destroyed. I characterized how workers from incipient colonies responded to eggs lacking queen fertility hydrocarbons. I found that established-queen-laid eggs, incipient-queen-laid eggs, and worker-laid eggs were not destroyed by workers at this colony stage. Destruction of worker-laid eggs is a form of policing, and theoretical models predict that policing should be strongest in incipient colonies. Since there was no evidence of policing by egg-eating in incipient C. floridanus colonies, I searched for evidence of another policing mechanism at this colony stage. Finding none, I discuss reasons why policing behavior may not be expressed in incipient colonies. I then considered the mechanism that accounts for the change in workers' response to eggs. By manipulating ants' egg experience and testing their egg-policing decisions, I found that ants use a combination of learned and innate criteria to discriminate between targets of care and destruction. Finally, I investigated how the increasing strength of queen-fertility hydrocarbons affects nestmate recognition, which also relies on cuticular hydrocarbons. I found that queens with strong fertility hydrocarbons can be transferred between established colonies without aggression, but they cannot be introduced into incipient colonies. Queens from incipient colonies cannot be transferred into incipient or established colonies.
ContributorsMoore, Dani (Author) / Liebig, Juergen (Thesis advisor) / Gadau, Juergen (Committee member) / Pratt, Stephen (Committee member) / Smith, Brian (Committee member) / Rutowski, Ronald (Committee member) / Arizona State University (Publisher)
Created2012
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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

When examining the medical doctrines of previous empires, they reveal the influence of religion, societal attitudes, and the historical context that influenced the scholars that penned them. The advancements during the Islamic Golden age can be seen in the field of medicine, which had the Greco-Roman medical corpus as their

When examining the medical doctrines of previous empires, they reveal the influence of religion, societal attitudes, and the historical context that influenced the scholars that penned them. The advancements during the Islamic Golden age can be seen in the field of medicine, which had the Greco-Roman medical corpus as their foundation and the source of the theory of the four humors and anatomical beliefs. This paper will analyze the effect of cultural, societal, and historical influences on the medical doctrines of Muslim medieval physicians in the Golden Age and the works of the Roman physician Galen, and demonstrate how these effects result in similarities and differences in medical practice and the understanding of disease and anatomy. Due to translation efforts that were supported by religious views on the accumulation of knowledge and the efforts of the Abbasid empire, resultant acceptance of the theory of the four humors and anatomical doctrines is observed in the treatment and perception of disease, which would consist of this paper's focus on surgery, diet therapy and associations with nature. However, with further analysis of the extent of this acceptance and the findings in the Islamic medical doctrines, the differences in experimental methods, religious interpretations, and cultural attitudes shows a deviation from the Galenic tradition, with the second set of the paper's focus being human dissection, cause of disease, and experimentation. The purpose of this research is to demonstrate the impact of religion, societal attitudes, culture and the accepted paradigm on the practice of medicine and the study of anatomy, and what would cause a challenge against the legacy of Galen.

Created2021-05
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ABSTRACT Communication is vital in the context of everyday life for all organisms, but particularly so in social insects, such as Z. nevadensis. The overall lifestyle and need for altruistic acts of individuals within a colony depends primarily on intracolony chemical communication, with a focus on odorants. The perception of

ABSTRACT Communication is vital in the context of everyday life for all organisms, but particularly so in social insects, such as Z. nevadensis. The overall lifestyle and need for altruistic acts of individuals within a colony depends primarily on intracolony chemical communication, with a focus on odorants. The perception of these odorants is made possible by the chemoreceptive functions of sensilla basiconica and sensilla trichoid which exist on the antennal structure. The present study consists of both a morphological analysis and electrophysiological experiment concerning sensilla basiconica. It attempts to characterize the function of neurons present in sensilla basiconica through single sensillum recordings and contributes to existing literature by determining if a social insect, such as the dampwood termite, is able to perceive a wide spectrum of odorants despite having significantly fewer olfactory receptors than most other social insect species. Results indicated that sensilla basiconica presence significantly out-paced that of sensilla trichoid and sensilla chaetica combined, on all flagellomeres. Analysis demonstrated significant responses to all general odorants and several cuticular hydrocarbons. Combined with the knowledge of fewer olfactory receptors present in this species and their lifestyle, results may indicate a positive association between the the social complexity of an insect's lifestyle and the number of ORs the individuals within that colony possess.
ContributorsMcGlone, Taylor (Author) / Liebig, Juergen (Thesis director) / Ghaninia, Majid (Committee member) / Barrett, The Honors College (Contributor)
Created2015-05
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Self-maintenance behaviors, like preening in birds, can have important effects on fitness in many animals. Birds produce preen oil, which is a mixture of volatile and non-volatile compounds, that is spread through their feathers during grooming and influences feather integrity, waterproofing, and coloration. As urban areas grow and present conditions

Self-maintenance behaviors, like preening in birds, can have important effects on fitness in many animals. Birds produce preen oil, which is a mixture of volatile and non-volatile compounds, that is spread through their feathers during grooming and influences feather integrity, waterproofing, and coloration. As urban areas grow and present conditions that may demand increased feather self-maintenance (e.g. due to soiling, pollution, elevated UV exposure due to natural habitat alterations), it is important to examine how preening and preen oil may be affected by this process. I assessed variation in preen oil composition in house finches (Haemorhous mexicanus) as a function of sex, urbanization, and plumage hue, a sexually selected indicator of male quality. Preen oil samples from birds captured at urban and rural sites were analyzed using gas chromatography-mass spectrometry. We detected 18 major peaks, which we tentatively identified as esters, and found that, although there were no sex or urban-rural differences in preen oil constituents, there was a significant interactive effect of sex and urbanization, with rural females and urban males having higher amounts of some components. This suggests that factors that vary with sex or urbanization, such as the timing of seasonal cycles, are affecting preen oil composition. There were no significant relationships between coloration and preen oil composition, suggesting that preen oil composition does not vary with male quality.
ContributorsBrooks, Ellen Elizabeth (Author) / McGraw, Kevin (Thesis director) / Liebig, Juergen (Committee member) / Weaver, Melinda (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor)
Created2015-05
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Honey bee (Apis mellifera) colonies have experienced substantial losses due to colony collapse disorder (CCD) since the first officially reported cases in 2006. Many factors have been implicated in CCD, including pests, pathogens, malnutrition, and pesticide use, but no correlation has been found between a single factor and the occurrence

Honey bee (Apis mellifera) colonies have experienced substantial losses due to colony collapse disorder (CCD) since the first officially reported cases in 2006. Many factors have been implicated in CCD, including pests, pathogens, malnutrition, and pesticide use, but no correlation has been found between a single factor and the occurrence of CCD. Fungicides have received less research attention compared to insecticides, despite the fact that fungicide application coincides with bloom and the presence of bees. Pristine fungicide is widely used in agriculture and is commonly found as a residue in hives. Several studies have concluded that Pristine can be used without harming bees, but reports of brood loss following Pristine application continue to surface across the country. The primary objectives of this study were to determine whether Pristine causes an aversive gustatory response in bees and whether consumption of an acute dose affects responsiveness to sucrose. An awareness of how foragers interact with contaminated food is useful to understand the likelihood that Pristine is ingested and how that may affect bees' ability to evaluate floral resources. Our results indicated that Pristine has no significant effect on gustatory response or sucrose responsiveness. There was no significant difference between bee responses to Pristine contaminated sucrose and sucrose alone, and no significant effect of Pristine on sucrose responsiveness. These results indicate that honey bees do not have a gustatory aversion to Pristine. A lack of aversion means that honey bees will continue collecting contaminated resources and dispersing them throughout the colony where it can affect brood and clean food stores.
ContributorsMcHugh, Cora Elizabeth (Co-author) / Jernigan, Christopher (Co-author, Committee member) / Burden, Christina (Co-author) / DeGrandi-Hoffman, Gloria (Co-author) / Smith, Brian (Thesis director) / Fewell, Jennifer (Committee member) / Barrett, The Honors College (Contributor) / School of Geographical Sciences and Urban Planning (Contributor) / School of Life Sciences (Contributor) / School of Art (Contributor)
Created2015-05
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This study illustrates the abilities of the honeybee, Apis mellifera, to learn and differentiate between patterns solely off their spatial frequencies. Patterns were chosen based off of calculations derived from the measurements of the physical construction of the apposition compound eye, which led to predictions of what the bees could

This study illustrates the abilities of the honeybee, Apis mellifera, to learn and differentiate between patterns solely off their spatial frequencies. Patterns were chosen based off of calculations derived from the measurements of the physical construction of the apposition compound eye, which led to predictions of what the bees could theoretically see. The hypothesis was then that bees would have a visual threshold where patterns with spatial frequencies that fall below this line should be easily distinguishable, and patterns above the threshold would have scores that mimic if the bees made choices randomly. There were 9 patterns tested, all with different spatial frequencies and in the colors of black, white, and gray. The bees were tested on their learning and pattern differentiation abilities with 10 pattern comparisons, with the lower frequency of the two being associated with an unscented sucrose solution reward. The results were surprising in that the previous studies pointing towards this visual threshold were inaccurate because of some of the patterns being learning in an intermediate ability. These intermediate scores suggest that the calculations predicting what the bees could see clearly were slightly wrong because it was more likely that the bees saw those images in more of a blur, which resulted in their intermediate score. Honeybees have served as a useful model organisms over the decades with studying learning involving visual information. This study lacked in its total numbers of trials and bees tested, which could have led to incomplete results, and this showing of an intermediate score and ability. Future studies should continue in order to advance this understanding of a perceptually and cognitively advance processing animal.
ContributorsBalsino, Brandon Bartholomew (Author) / Harrison, Jon (Thesis director) / Smith, Brian (Committee member) / Duell, Meghan (Committee member) / School of Life Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2015-12
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Selenium, a group 16 metalloid on the periodic table, is a necessary mineral for many organisms. Trace amounts of selenium are essential for normal development, antioxidant protein function, enzyme function, and hormone regulation (Burden et al., 2016). However, when selenium is found in toxic amounts in organisms, it has been

Selenium, a group 16 metalloid on the periodic table, is a necessary mineral for many organisms. Trace amounts of selenium are essential for normal development, antioxidant protein function, enzyme function, and hormone regulation (Burden et al., 2016). However, when selenium is found in toxic amounts in organisms, it has been found to substitute for sulfur in proteins, which can be toxic to these animals, and cause oxidative stress (Quinn et al., 2007). Using the previous research done with acute exposure to organic and inorganic selenium compounds, we hypothesized that the inorganic sodium selenate would significantly decrease learning and memory recall for both chronic and acute exposure. We also hypothesized that the consumption of organic methylseleno-L-cysteine by honey bees would decrease learning and memory recall for both the chronic and acute exposure. We further hypothesized that protein carbonyl content would be increased due to oxidative damage caused by selenium in both the sodium selenate and the methylseleno-L-cysteine treatment groups, but that the inorganic selenium compound would increase the carbonyl content more than the methylseleno-L-cysteine. To run the experiments, three tents outside had two colonies in each tent. One tent contained the sodium selenate group, another had the sucrose control, and one contained the methylseleno-L-cysteine group. The treatment groups were fed selenium in their sucrose feeders. The first part of the experiment was training the bees by using proboscis extension response (PER) to teach them to extend their proboscis to the rewarded odor and not to the unrewarded odor. This was done by pairing the rewarded odor with a sucrose reward and not pairing it with the unrewarded odor. Then their short-term and long-term memory recall was tested. The second part of the experiment was checking for oxidative damage by measuring the protein carbonyl content in the bees. Three boxes were set up with the same three treatment groups as used in the tents. The treatment group bees were exposed to selenium in the sucrose feeders and in the pollen patties. After one week, the living bees were removed and frozen. They were then homogenized to extract protein. The first assay run was the protein content assay to establish a standard protein concentration for samples. Then a protein carbonyl assay was run, to determine the protein carbonyl content. Overall, the experiment found that exposure to selenium negatively impacted honey bees learning and memory recall significantly. Chronic exposure to the inorganic selenate reduced the bees' long-term memory abilities to differentiate between odors. With methylseleno-L-cysteine, it had no significant effect for the chronic exposure, but for the acute exposure, it had a significant impairment on their abilities to distinguish between the rewarded and unrewarded odors during conditioning. Our results showed that from our experiment there appeared to be no significant effect of selenium exposure on the increase of carbonylation content in the different treatment groups. This is most likely due to the fact the carbonyl content was not detectable because the protein concentration was low in the samples (approximately 3.5 mg/mL).
ContributorsWinski, Alexandra (Co-author) / Winski, Brandon (Co-author) / Smith, Brian (Thesis director) / Harrison, Jon (Committee member) / Burden, Christina (Committee member) / School of Life Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2016-05
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Collective decision making in social organism societies involves a large network of communication systems. Studying the processes behind the transmission of information allows for greater understanding of the decision making capabilities of a group. For Temnothorax rugatulus colonies, information is commonly spread in the form of tandem running, a linear

Collective decision making in social organism societies involves a large network of communication systems. Studying the processes behind the transmission of information allows for greater understanding of the decision making capabilities of a group. For Temnothorax rugatulus colonies, information is commonly spread in the form of tandem running, a linear recruitment pattern where a leading ant uses a short-ranged pheromone to direct a following ant to a target location (in tandem).The observed phenomenon of reverse tandem running (RTR), where a follower is lead from a target back to the home nest, has not been as extensively studied as forward tandem running and transportation recruitment activities. This study seeks to explain a potential reason for the presence of the RTR behavior; more specifically, the study explores the idea that reverse tandem run followers are being shown a specific route to the home nest by a highly experienced and efficient leading ant. Ten colonies had migrations induced experimentally in order to generate some reverse tandem running activity. Once an RTR has been observed, the follower and leader were studied for behavior and their pathways were analyzed. It was seen that while RTR paths were quite efficient (1.4x a straight line distance), followers did not experience a statistically significant improvement in their pathways between the home and target nests (based on total distance traveled) when compared to similar non-RTR ants. Further, RTR leading ants were no more efficient than other non-RTR ants. It was observed that some followers began recruiting after completion of an RTR, but the number than changed their behavior was not significant. Thus, the results of this experiment cannot conclusively show that RTR followers are utilizing reverse tandem runs to improve their routes between the home and target nests.
ContributorsColling, Blake David (Author) / Pratt, Stephen (Thesis director) / Liebig, Juergen (Committee member) / Sasaki, Takao (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor)
Created2014-12
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Communication amongst eusocial insect is key to their success. Ants rely on signaling to mediate many different functions within a colony such as policing and nest mate recognition. Camponotus floridanus uses chemosensory signaling in the form of cuticular hydrocarbons to regulate these functions. Each cuticular hydrocarbon profile contains numerous hydrocarbons,

Communication amongst eusocial insect is key to their success. Ants rely on signaling to mediate many different functions within a colony such as policing and nest mate recognition. Camponotus floridanus uses chemosensory signaling in the form of cuticular hydrocarbons to regulate these functions. Each cuticular hydrocarbon profile contains numerous hydrocarbons, however it is yet to be seen if Camponotus floridanus can discriminate between linear hydrocarbons of similar length. Individual specimens were conditioned in three different ways: 5 conditioning with high concentration of sugar water (1;1 ratio), 1 conditioning with high concentration of sugar water, and 5 conditioning with low concentration of sugar water (1;4). Two linear hydrocarbons were use, C23 and C24, with C23 always being the conditioned stimulus. Specimens who were conditioned 5 times with high concentration of sugar water were the only group to show a significant response to the conditioned stimulus with a p-value of .008 and exhibited discrimination behavior 46% of the time. When compared 5 conditioning with high concentration to the other two testing conditioning groups, 1 conditioning with high concentration produced an insignificant p-value of .13 was obtained whereas when comparing it with 5 conditioning low concentration of sugar a significant p-value of .0132 was obtained. This indiciates that Camponotus floridanus are capable of discrimination however must be conditioned with high concentration of sugar water, while number of conditioning is insignificant.
ContributorsDamari, Ben Aviv (Author) / Liebig, Juergen (Thesis director) / Ghaninia, Majid (Committee member) / Pratt, Stephen (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor)
Created2014-05