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How a colony regulates the division of labor to forage for nutritional resources while accommodating for changes in colony demography is a fundamental question in the sociobiology of social insects. In honey bee, Apis mellifera, brood composition impacts the division of labor, but it is unknown if colonies adjust the

How a colony regulates the division of labor to forage for nutritional resources while accommodating for changes in colony demography is a fundamental question in the sociobiology of social insects. In honey bee, Apis mellifera, brood composition impacts the division of labor, but it is unknown if colonies adjust the allocation of foragers to carbohydrate and protein resources based on changes in the age demography of larvae and the pheromones they produce. Young and old larvae produce pheromones that differ in composition and volatility. In turn, nurses differentially provision larvae, feeding developing young worker larvae a surplus diet that is more queen-like in protein composition and food availability, while old larvae receive a diet that mimics the sugar composition of the queen larval diet but is restrictively fed instead of provided ad lib. This research investigated how larval age and the larval pheromone e-β ocimene (eβ) impact foraging activity and foraging load. Additional cage studies were conducted to determine if eβ interacts synergistically with queen mandibular pheromone (QMP) to suppress ovary activation and prime worker physiology for nursing behavior. Lastly, the priming effects of larval age and eβ on worker physiology and the transition from in-hive nursing tasks to outside foraging were examined. Results indicate that workers differentially respond to larvae of different ages, likely by detecting changes in the composition of the pheromones they emit. This resulted in adjustments to the foraging division of labor (pollen vs. nectar) to ensure that the nutritional needs of the colony's brood were met. For younger larvae and eβ, this resulted in a bias favoring pollen collection. The cage studies reveal that both eβ and QMP suppressed ovary activation, but the larval pheromone was more effective. Maturing in an environment of young or old larvae primed bees for nursing and impacted important endocrine titers involved in the transition to foraging, so bees maturing in the presence of larvae foraged earlier than control bees reared with no brood.
ContributorsTraynor, Kirsten S. (Author) / Page, Robert E. (Thesis advisor) / Hölldobler, Berthold (Committee member) / Pratt, Stephen (Committee member) / Liebig, Jürgen (Committee member) / Brent, Colin (Committee member) / Baluch, Page (Committee member) / Arizona State University (Publisher)
Created2014
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Description
Food is an essential driver of animal behavior. For social organisms, the acquisition of food guides interactions with the environment and with group-mates. Studies have focused on how social individuals find and choose food sources, and share both food and information with group-mates. However, it is often not clear how

Food is an essential driver of animal behavior. For social organisms, the acquisition of food guides interactions with the environment and with group-mates. Studies have focused on how social individuals find and choose food sources, and share both food and information with group-mates. However, it is often not clear how experiences throughout an individual's life influence such interactions. The core question of this thesis is how individuals’ experience contributes to within-caste behavioral variation in a social group. I investigate the effects of individual history, including physical injury and food-related experience, on individuals' social food sharing behavior, responses to food-related stimuli, and the associated neural biogenic amine signaling pathways. I use the eusocial honey bee (Apis mellifera) system, one in which individuals exhibit a high degree of plasticity in responses to environmental stimuli and there is a richness of communicatory pathways for food-related information. Foraging exposes honey bees to aversive experiences such as predation, con-specific competition, and environmental toxins. I show that foraging experience changes individuals' response thresholds to sucrose, a main component of adults’ diets, depending on whether foraging conditions are benign or aversive. Bodily injury is demonstrated to reduce individuals' appetitive responses to new, potentially food-predictive odors. Aversive conditions also impact an individual's social food sharing behavior; mouth-to-mouse trophallaxis with particular groupmates is modulated by aversive foraging conditions both for foragers who directly experienced these conditions and non-foragers who were influenced via social contact with foragers. Although the mechanisms underlying these behavioral changes have yet to be resolved, my results implicate biogenic amine signaling pathways as a potential component. Serotonin and octopamine concentrations are shown to undergo long-term change due to distinct foraging experiences. My work serves to highlight the malleability of a social individual's food-related behavior, suggesting that environmental conditions shape how individuals respond to food and share information with group-mates. This thesis contributes to a deeper understanding of inter-individual variation in animal behavior.
ContributorsFinkelstein, Abigail (Author) / Amdam, Gro V (Thesis advisor) / Conrad, Cheryl (Committee member) / Smith, Brian (Committee member) / Neisewander, Janet (Committee member) / Bimonte-Nelson, Heather A. (Committee member) / Arizona State University (Publisher)
Created2017
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Description
The ability to detect and appropriately respond to chemical stimuli is important for many organisms, ranging from bacteria to multicellular animals. Responses to these stimuli can be plastic over multiple time scales. In the short-term, the synaptic strengths of neurons embedded in neural circuits can be modified and result in

The ability to detect and appropriately respond to chemical stimuli is important for many organisms, ranging from bacteria to multicellular animals. Responses to these stimuli can be plastic over multiple time scales. In the short-term, the synaptic strengths of neurons embedded in neural circuits can be modified and result in various forms of learning. In the long-term, the overall developmental trajectory of the olfactory network can be altered and synaptic strengths can be modified on a broad scale as a direct result of long-term (chronic) stimulus experience. Over evolutionary time the olfactory system can impose selection pressures that affect the odorants used in communication networks. On short time scales, I measured the effects of repeated alarm pheromone exposure on the colony-level defense behaviors in a social bee. I found that the responses to the alarm pheromone were plastic. This suggests that there may be mechanisms that affect individual plasticity to pheromones and regulate how these individuals act in groups to coordinate nest defense. On longer time scales, I measured the behavioral and neural affects of bees given a single chronic odor experience versus bees that had a natural, more diverse olfactory experience. The central brains of bees with a deprived odor experience responded more similarly to odorants in imaging studies, and did not develop a fully mature olfactory network. Additionally, these immature networks showed behavioral deficits when recalling odor mixture components. Over evolutionary time, signals need to engage the attention of and be easily recognized by bees. I measured responses of bees to a floral mixture and its constituent monomolecular components. I found that natural floral mixtures engage the orientation of bees’ antennae more strongly than single-component odorants and also provide more consistent central brain responses between stimulations. Together, these studies highlight the importance of olfactory experience on different scales and how the nervous system might impose pressures to select the stimuli used as signals in communication networks.
ContributorsJernigan, Christopher (Author) / Smith, Brian H. (Thesis advisor) / Newbern, Jason (Committee member) / Harrisoin, Jon (Committee member) / Rutowski, Ronald (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Amongst the most studied of the social insects, the honey bee has a prominent place due to its economic importance and influence on human societies. Honey bee colonies can have over 50,000 individuals, whose activities are coordinated by chemical signals called pheromones. Because these pheromones are secreted from various exocrine

Amongst the most studied of the social insects, the honey bee has a prominent place due to its economic importance and influence on human societies. Honey bee colonies can have over 50,000 individuals, whose activities are coordinated by chemical signals called pheromones. Because these pheromones are secreted from various exocrine glands, the proper development and function of these glands are vital to colony dynamics. In this thesis, I present a study of the developmental ontogeny of the exocrine glands found in the head of the honey bee. In Chapter 2, I elucidate how the larval salivary gland transitions to an adult salivary gland through apoptosis and cell growth, differentiation and migration. I also explain the development of the hypopharyngeal and the mandibular gland using apoptotic markers and cytoskeletal markers like tubulin and actin. I explain the fundamental developmental plan for the formation of the glands and show that apoptosis plays an important role in the transformation toward an adult gland.
ContributorsNath, Rachna (Author) / Gadau, Juergen (Thesis advisor) / Rawls, Alan (Committee member) / Harrison, Jon (Committee member) / Arizona State University (Publisher)
Created2018
Description

Foraging honey bees are challenged to balance the energetic costs of thermoregulating and load-carriage at the same time when flying in hot environments. Honey bees can reduce metabolic rate and wingbeat frequency in response to heat, but the kinematic strategies they use while carrying loads are unknown. I observed honey

Foraging honey bees are challenged to balance the energetic costs of thermoregulating and load-carriage at the same time when flying in hot environments. Honey bees can reduce metabolic rate and wingbeat frequency in response to heat, but the kinematic strategies they use while carrying loads are unknown. I observed honey bees (Apis mellifera) carrying a range of nectar loads (0 to 80% of their own body weight in nectar) when flying at 25 and 40°C air temperatures, and found that hotter honey bees decreased their wingbeat frequency (from 230 to 195 Hz) and increased their stroke amplitude (from 90 to 98°) to generate increasing aerodynamic power as they carry heavier nectar loads. The bees flying at 40°C air temperature carrying heavier loads did increase their wingbeat frequency compared to the unloaded individuals. Despite the kinematic changes, both the hot and cold honey bees were able to generate sufficient power to carry loads of roughly equal mass. Bees flying at 40°C air temperature produced more power than their cooler counterparts, suggesting a more efficient mechanism of load carriage.

ContributorsWeisman, Ethan (Author) / Harrison, Jon (Thesis director) / Glass, Jordan (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor) / Department of Psychology (Contributor)
Created2023-05
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Description
The alarming decline of insect pollinators is due in part to agrochemical exposure and climate warming. This thesis focuses on understanding how exposure to a commonly used fungicide and high air temperature affect the flight behavior and physiology of the very important commercial pollinator, Apis mellifera. I found that honey

The alarming decline of insect pollinators is due in part to agrochemical exposure and climate warming. This thesis focuses on understanding how exposure to a commonly used fungicide and high air temperature affect the flight behavior and physiology of the very important commercial pollinator, Apis mellifera. I found that honey bees reared on pollen contaminated with field-realistic levels of a fungicide (Pristine®) commonly applied to almond blossoms before pollination had smaller thoraxes, possibly due to inhibition of protein digestion, plausibly reducing flight capability. By flying unloaded bees in low density air to elicit maximal performance, I found that consumption of high doses of fungicide during development inhibited maximal flight performance, but consumption of field-realistic doses did not. To understand climatic-warming effects on honey bees, I flew unloaded foragers at various air densities and temperatures to assess the effects of flight muscle temperature (29 to 44°C) on maximal aerobic metabolism. Flight metabolic rate peaked at a muscle temperature of 39°C and decreased by ~2% per degree below and ~5% per degree above this optimum. Carrying nectar loads increased flight muscle temperatures and flight metabolism of foragers flying at air temperatures of 20 or 30°C. Yet, remarkably, bees flying at 40°C were able to carry loads without heating up or increasing metabolic rate. Bees flying at 40°C increased evaporative cooling and decreased metabolic heat production to thermoregulate. High speed video revealed that bees flying at 40°C air temperature lowered their wing beat frequency while increasing stroke amplitude, increasing flight efficiency. My data also suggests that cooler bees use wing kinematic strategies that increase flight stability and maneuverability while generating excess heat that warms their flight muscle toward optimum. High water loss rates during flight likely limit foraging in dry air temperatures above 46°C, suggesting that CTmax measures of resting honey bees significantly overestimate when high air temperature will negatively impact flight and foraging.
ContributorsGlass, Jordan Robert (Author) / Harrison, Jon F. (Thesis advisor) / Denardo, Dale F. (Committee member) / Dudley, Robert (Committee member) / Fewell, Jennifer H. (Committee member) / Arizona State University (Publisher)
Created2023
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Description
Honeybees require the use of their antennae to perceive different scents and pheromones, communicate with other members of the colony, and even detect wind vibrations, sound waves, and carbon dioxide levels. Limiting and/or removing this sense makes bees much less effective at acquiring information. However, how antennal movements might be

Honeybees require the use of their antennae to perceive different scents and pheromones, communicate with other members of the colony, and even detect wind vibrations, sound waves, and carbon dioxide levels. Limiting and/or removing this sense makes bees much less effective at acquiring information. However, how antennal movements might be important for olfaction has not been studied in detail. The focus of this work was to evaluate how restriction of antennae movements might affect a bee’s ability to detect and perceive odors. Bees were made to learn a certain odor and were then split up into a control group, a treatment group that had their antennae fixed with eicosane, and a sham treatment group that had a dot of eicosane on their heads in such a way that it would not affect antennae movements but still add the same amount of weight. Following a period of acclimation, the bees were tested with the conditioned odor, one that was perceptually similar to it, and to a dissimilar odor. Using proboscis-extension duration and latency as response measures, it became clear that both antenna fixation and sham treatments affected the conditioned behavior. However, these treatment effects did not reach statistical significance. Briefly, both fixation of antennae as well as the sham treatment reduced the discriminability of the conditioned and similar odors. Although more data can be collected to more fully evaluate the significance of the treatments, the behavior of the sham group could indicate that mechanoreceptive hairs on the head play an important role in olfaction. It is also possible that there are other factors at play, possibly induced by the fixed bees’ increased stress levels.
ContributorsHozan, Alvin Robert (Author) / Smith, Brian H (Thesis advisor) / Lei, Hong (Committee member) / Cook, Chelsea (Committee member) / Arizona State University (Publisher)
Created2021
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Description
Pollinator populations globally have declined at concerning rates in recent years, which is problematic given that roughly a third of all food production depends on them. Managed honey bee colony losses in particular have alarmed beekeepers and scientists, especially in the United States. Widespread agrochemical use has been implicated as

Pollinator populations globally have declined at concerning rates in recent years, which is problematic given that roughly a third of all food production depends on them. Managed honey bee colony losses in particular have alarmed beekeepers and scientists, especially in the United States. Widespread agrochemical use has been implicated as one of the major causes of these colony losses. While the lethal effects of agrochemicals often receive the most attention, sublethal effects can occur at lower doses and can substantially weaken colonies over time. Impaired associative learning ability is a sublethal effect of a number of agrochemicals, and is particularly concerning, as it may hinder the abilities of bees to forage for food or find their way back to the colony. Here, I focus on the fungicide Pristine® (active ingredients: 25.2% boscalid, 12.8% pyraclostrobin), which is sprayed on honey bee-pollinated crops during bloom and is known to poison bee mitochondria at ppm levels. First, I show that Pristine® impairs performance on an associative learning assay in the laboratory. Next, I show that Pristine® alters carbohydrate absorption in honey bees, providing a possible mechanism underlying this impaired learning performance. Finally, I demonstrate that Pristine® interacts with high temperatures to induce homing failure in exposed bees. My results raise concerns that this common fungicide may not be safe for pollinators and will be relevant to policymakers as they make decisions surrounding the regulation of fungicide use in agriculture.
ContributorsDesJardins, Nicole (Author) / Harrison, Jon F (Thesis advisor) / Smith, Brian H (Thesis advisor) / DeGrandi-Hoffman, Gloria (Committee member) / DeNardo, Dale (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2023