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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
This research focuses on the intricate dynamical systems of eusocial insects, particularly ants, and honey bees, known for their highly organized colonies and cooperative behaviors. Research on eusocial insects contributes to understanding of animal and social behavior and promises to help agriculture and have huge economic impacts. Collaborating closely with

This research focuses on the intricate dynamical systems of eusocial insects, particularly ants, and honey bees, known for their highly organized colonies and cooperative behaviors. Research on eusocial insects contributes to understanding of animal and social behavior and promises to help agriculture and have huge economic impacts. Collaborating closely with ecologists, I construct diverse mathematical models tailored to different environmental contexts. These models encompass individual stochastic (Agent-based model), Ordinary Differential Equation (ODE), non-autonomous, and Delay Differential Equation (DDE) models, rigorously validated with experimental data and statistical methods. Employing dynamical theory, bifurcation analysis, and numerical simulations, I gain deeper insights into the adaptive behaviors exhibited by these insects at both colony and individual levels. Our investigation addresses pivotal questions: 1) What mechanisms underlie spatial heterogeneity within social insect colonies, influencing the spread of information and pathogens through their intricate social networks?2) How can I develop accurate mathematical models incorporating age structures, particularly for species like honeybees, utilizing delayed differential equations? 3) What is the influence of seasonality on honeybee population dynamics in the presence of parasites, as explored through non-autonomous equations? 4) How do pesticides impact honeybee population dynamics, considering delayed equations and seasonality? Key findings highlight:1) The spatial distribution within colonies significantly shapes contact dynamics, thereby influencing the dissemination of information and the allocation of tasks. 2) Accurate modeling of honeybee populations necessitates the incorporation of age structure, as well as careful consideration of seasonal variations. 3) Seasonal fluctuations in egg-laying rates exert varying effects on the survival of honeybee colonies. 4) Pesticides wield a substantial influence on adult bee mortality rates and the consumption ratios of pollen. This research not only unveils the intricate interplay between intrinsic and environmental factors affecting social insects but also provides broader insights into social behavior and the potential ramifications of climate change.
ContributorsChen, Jun (Author) / Kang, Yun (Thesis advisor) / DeGrandi-Hoffman, Gloria (Committee member) / Fewell, Jeniffer (Committee member) / Harrison, Jon (Committee member) / Towers, Sherry (Committee member) / Arizona State University (Publisher)
Created2023
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Description
The Northwest (NW) Atlantic porbeagle Lamna nasus is overfished and captured as bycatch in fisheries within the region. A comprehensive understanding of the population’s life history (e.g., reproduction) and habitat use, and the impact of capture with different gear types (e.g., post-release mortality) is needed to ensure effective fisheries management

The Northwest (NW) Atlantic porbeagle Lamna nasus is overfished and captured as bycatch in fisheries within the region. A comprehensive understanding of the population’s life history (e.g., reproduction) and habitat use, and the impact of capture with different gear types (e.g., post-release mortality) is needed to ensure effective fisheries management plans, develop bycatch mitigation strategies, and support stock recovery. This research used satellite tagging technologies to address gaps in knowledge needed to support management and conservation decisions for the NW Atlantic porbeagle. I provided the first estimate of post-release survival and recovery periods for immature porbeagles captured with rod-and-reel. Although survival was high (100%), juvenile porbeagles exhibited a recovery period in surface waters that may make them vulnerable to further fishing interactions. Next, I described the vertical habitat use of young porbeagles to recommend possible fishing modifications to reduce risk of capture. Young porbeagles spent more time in surface waters during summer compared to fall and during the night compared to day, suggesting that risk of capture may be reduced by setting gear deeper during summer and at night when this life stage’s behavior is reduced to the upper water column. Then, I provided an analysis of the seasonal and life stage-based habitat use of porbeagles. Space use was concentrated in continental shelf waters around Cape Cod, Massachusetts regardless of season and life stage. Given the relatively small and static high occupancy area overlaps with a high concentration of fishing activity, this region could be considered for spatial management of the NW Atlantic porbeagle. Finally, I used ultrasonography and satellite tagging to describe the three-dimensional habitat use of gravid porbeagles for the first time. Gravid porbeagles demonstrated seasonal differences in horizontal and vertical habitat use but spent most of the pupping season in waters southeast of Cape Cod or on Georges Bank, suggesting this region may be serving as a pupping ground for at least a portion of this population. Conservation efforts should focus on these important habitats to protect the next generation of porbeagles.
ContributorsAnderson, Brooke Nicole (Author) / Ferry, Lara (Thesis advisor) / Bowlby, Heather (Committee member) / Hammerschlag, Neil (Committee member) / Kang, Yun (Committee member) / Saul, Steven (Committee member) / Sulikowski, James (Committee member) / Arizona State University (Publisher)
Created2024
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Description
The most advanced social insects, the eusocial insects, form often large societies in which there is reproductive division of labor, queens and workers, have overlapping generations, and cooperative brood care where daughter workers remain in the nest with their queen mother and care for their siblings. The eusocial insects

The most advanced social insects, the eusocial insects, form often large societies in which there is reproductive division of labor, queens and workers, have overlapping generations, and cooperative brood care where daughter workers remain in the nest with their queen mother and care for their siblings. The eusocial insects are composed of representative species of bees and wasps, and all species of ants and termites. Much is known about their organizational structure, but remains to be discovered.

The success of social insects is dependent upon cooperative behavior and adaptive strategies shaped by natural selection that respond to internal or external conditions. The objective of my research was to investigate specific mechanisms that have helped shaped the structure of division of labor observed in social insect colonies, including age polyethism and nutrition, and phenomena known to increase colony survival such as egg cannibalism. I developed various Ordinary Differential Equation (ODE) models in which I applied dynamical, bifurcation, and sensitivity analysis to carefully study and visualize biological outcomes in social organisms to answer questions regarding the conditions under which a colony can survive. First, I investigated how the population and evolutionary dynamics of egg cannibalism and division of labor can promote colony survival. I then introduced a model of social conflict behavior to study the inclusion of different response functions that explore the benefits of cannibalistic behavior and how it contributes to age polyethism, the change in behavior of workers as they age, and its biological relevance. Finally, I introduced a model to investigate the importance of pollen nutritional status in a honeybee colony, how it affects population growth and influences division of labor within the worker caste. My results first reveal that both cannibalism and division of labor are adaptive strategies that increase the size of the worker population, and therefore, the persistence of the colony. I show the importance of food collection, consumption, and processing rates to promote good colony nutrition leading to the coexistence of brood and adult workers. Lastly, I show how taking into account seasonality for pollen collection improves the prediction of long term consequences.
ContributorsRodríguez Messan, Marisabel (Author) / Kang, Yun (Thesis advisor) / Castillo-Chavez, Carlos (Thesis advisor) / Kuang, Yang (Committee member) / Page Jr., Robert E (Committee member) / Gardner, Carl (Committee member) / Arizona State University (Publisher)
Created2018
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Description
There has been important progress in understanding ecological dynamics through the development of the theory of ecological stoichiometry. This fast growing theory provides new constraints and mechanisms that can be formulated into mathematical models. Stoichiometric models incorporate the effects of both food quantity and food quality into a single framework

There has been important progress in understanding ecological dynamics through the development of the theory of ecological stoichiometry. This fast growing theory provides new constraints and mechanisms that can be formulated into mathematical models. Stoichiometric models incorporate the effects of both food quantity and food quality into a single framework that produce rich dynamics. While the effects of nutrient deficiency on consumer growth are well understood, recent discoveries in ecological stoichiometry suggest that consumer dynamics are not only affected by insufficient food nutrient content (low phosphorus (P): carbon (C) ratio) but also by excess food nutrient content (high P:C). This phenomenon, known as the stoichiometric knife edge, in which animal growth is reduced not only by food with low P content but also by food with high P content, needs to be incorporated into mathematical models. Here we present Lotka-Volterra type models to investigate the growth response of Daphnia to algae of varying P:C ratios. Using a nonsmooth system of two ordinary differential equations (ODEs), we formulate the first model to incorporate the phenomenon of the stoichiometric knife edge. We then extend this stoichiometric model by mechanistically deriving and tracking free P in the environment. This resulting full knife edge model is a nonsmooth system of three ODEs. Bifurcation analysis and numerical simulations of the full model, that explicitly tracks phosphorus, leads to quantitatively different predictions than previous models that neglect to track free nutrients. The full model shows that the grazer population is sensitive to excess nutrient concentrations as a dynamical free nutrient pool induces extreme grazer population density changes. These modeling efforts provide insight on the effects of excess nutrient content on grazer dynamics and deepen our understanding of the effects of stoichiometry on the mechanisms governing population dynamics and the interactions between trophic levels.
ContributorsPeace, Angela (Author) / Kuang, Yang (Thesis advisor) / Elser, James J (Committee member) / Baer, Steven (Committee member) / Tang, Wenbo (Committee member) / Kang, Yun (Committee member) / Arizona State University (Publisher)
Created2014
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Description
In 1968, phycologist M.R. Droop published his famous discovery on the functional relationship between growth rate and internal nutrient status of algae in chemostat culture. The simple notion that growth is directly dependent on intracellular nutrient concentration is useful for understanding the dynamics in many ecological systems. The cell quota

In 1968, phycologist M.R. Droop published his famous discovery on the functional relationship between growth rate and internal nutrient status of algae in chemostat culture. The simple notion that growth is directly dependent on intracellular nutrient concentration is useful for understanding the dynamics in many ecological systems. The cell quota in particular lends itself to ecological stoichiometry, which is a powerful framework for mathematical ecology. Three models are developed based on the cell quota principal in order to demonstrate its applications beyond chemostat culture.

First, a data-driven model is derived for neutral lipid synthesis in green microalgae with respect to nitrogen limitation. This model synthesizes several established frameworks in phycology and ecological stoichiometry. The model demonstrates how the cell quota is a useful abstraction for understanding the metabolic shift to neutral lipid production that is observed in certain oleaginous species.

Next a producer-grazer model is developed based on the cell quota model and nutrient recycling. The model incorporates a novel feedback loop to account for animal toxicity due to accumulation of nitrogen waste. The model exhibits rich, complex dynamics which leave several open mathematical questions.

Lastly, disease dynamics in vivo are in many ways analogous to those of an ecosystem, giving natural extensions of the cell quota concept to disease modeling. Prostate cancer can be modeled within this framework, with androgen the limiting nutrient and the prostate and cancer cells as competing species. Here the cell quota model provides a useful abstraction for the dependence of cellular proliferation and apoptosis on androgen and the androgen receptor. Androgen ablation therapy is often used for patients in biochemical recurrence or late-stage disease progression and is in general initially effective. However, for many patients the cancer eventually develops resistance months to years after treatment begins. Understanding how and predicting when hormone therapy facilitates evolution of resistant phenotypes has immediate implications for treatment. Cell quota models for prostate cancer can be useful tools for this purpose and motivate applications to other diseases.
ContributorsPacker, Aaron (Author) / Kuang, Yang (Thesis advisor) / Nagy, John (Committee member) / Smith, Hal (Committee member) / Kostelich, Eric (Committee member) / Kang, Yun (Committee member) / Arizona State University (Publisher)
Created2014
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Description
Grassland habitat restoration activities are occurring within the semi-arid grasslands of the Agua Fria National Monument located 65 km north of Phoenix, AZ. The goal of these restoration activities is to reduce woody species encroachment, remove lignified plant materials and recycle nutrients within the ecosystem thus improving range conditions for

Grassland habitat restoration activities are occurring within the semi-arid grasslands of the Agua Fria National Monument located 65 km north of Phoenix, AZ. The goal of these restoration activities is to reduce woody species encroachment, remove lignified plant materials and recycle nutrients within the ecosystem thus improving range conditions for both wildlife species and livestock. Broadcast burning, juniper thinning and slash pile burns are the principle tools used to accomplish resource objectives. Line cover, belt transect, densities, heights and biomass of vegetation data were collected to determine the response of the vegetative community to habitat restoration activities. Principal Component Analysis (PCA) was used to reduce data analysis to the more influential factors. Regression analysis was conducted for statistically significant response variables. Quadratic regression analysis found low predictive values. In broadcast burn treatment units, all important factors as identified by PCA had low predictive factors but significantly differed (R2 <0.01, p<0.05) between unburned and the years post treatment. Regression analysis found significant, albeit weak, relationships between time since treatment and independent variables. In pile burn treatment units, data reduction by PCA was not possible in a biologically meaningful way due to the high variability within treatment units. This suggests the effect of juniper encroachment on grassland vegetation persists long after junipers have been cut and burned. This study concluded that broadcast burning of the central Arizona grasslands does significantly alter many components of the vegetative community. Fuels treatments generally initially reduced both perennial woody species and grasses in number and height for two year post fire. However, palatable shrubs, in particular shrubby buckwheat, were not significantly different in broadcast burn treatment areas. The vegetative community characteristics of juniper encroached woodlands of central Arizona are unaffected by the removal and burning of junipers aside from the removal of hiding cover for predators for multiple years. It is recommended that habitat restoration activities continue provided the needs of wildlife are considered, especially pronghorn, with the incorporation of state and transition models specific to each of the respective ecological site descriptions and with the consideration of the effects of fire to pronghorn fawning habitat.
ContributorsSitzmann, Paul Roman (Author) / Miller, William (Thesis advisor) / Alford, Eddie (Committee member) / Green, Douglas (Committee member) / Arizona State University (Publisher)
Created2014
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Description
The flexibility and robustness of social insect colonies, when they cope with challenges as integrated units, raise many questions, such as how hundreds and thousands of individual local responses are coordinated without a central controlling process. Answering such questions requires: 1. Quantifiable collective responses of colonies under specific scenarios; 2.

The flexibility and robustness of social insect colonies, when they cope with challenges as integrated units, raise many questions, such as how hundreds and thousands of individual local responses are coordinated without a central controlling process. Answering such questions requires: 1. Quantifiable collective responses of colonies under specific scenarios; 2. Decomposability of the collective colony-level response into individual responses; and 3. Mechanisms to integrate the colony- and individual-level responses. In the first part of my dissertation, I explore coordinated collective responses of colonies in during the alarm response to an alarmed nestmate (chapter 2&3). I develop a machine-learning approach to quantitatively estimate the collective and individual alarm response (chapter 2). Using this methodology, I demonstrate that colony alarm responses to the introduction of alarmed nestmates can be decomposed into immediately cascading, followed by variable dampening processes. Each of those processes are found to be modulated by variation in individual alarm responsiveness, as measured by alarm response threshold and persistence of alarm behavior. This variation is modulated in turn by environmental context, in particular with task-related social context (chapter 3). In the second part of my dissertation, I examine the mechanisms responsible for colonial changes in metabolic rate during ontogeny. Prior studies have found that larger ant colonies (as for larger organisms) have lower mass-specific metabolic rates, but the mechanisms remain unclear. In a 3.5-year study on 25 colonies, metabolic rates of colonies and colony components were measured during ontogeny (chapter 4). The scaling of metabolic rate during ontogeny was fit better by segmented regression or quadratic regression models than simple linear regression models, showing that colonies do not follow a universal power-law of metabolism during the ontogenetic development. Furthermore, I showed that the scaling of colonial metabolic rates can be primarily explained by changes in the ratio of brood to adult workers, which nonlinearly affects colonial metabolic rates. At high ratios of brood to workers, colony metabolic rates are low because the metabolic rate of larvae and pupae are much lower than adult workers. However, the high colony metabolic rates were observed in colonies with moderate brood: adult ratios, because higher ratios cause adult workers to be more active and have higher metabolic rates, presumably due to the extra work required to feed more brood.
ContributorsGuo, Xiaohui (Author) / Fewell, Jennifer H (Thesis advisor) / Kang, Yun (Thesis advisor) / Harrison, Jon F (Committee member) / Liebig, Juergen (Committee member) / Pratt, Stephen C (Committee member) / Pavlic, Theodore P (Committee member) / Arizona State University (Publisher)
Created2021
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Description
A key factor in the success of social animals is their organization of work. Mathematical models have been instrumental in unraveling how simple, individual-based rules can generate collective patterns via self-organization. However, existing models offer limited insights into how these patterns are shaped by behavioral differences within groups, in part

A key factor in the success of social animals is their organization of work. Mathematical models have been instrumental in unraveling how simple, individual-based rules can generate collective patterns via self-organization. However, existing models offer limited insights into how these patterns are shaped by behavioral differences within groups, in part because they focus on analyzing specific rules rather than general mechanisms that can explain behavior at the individual-level. My work argues for a more principled approach that focuses on the question of how individuals make decisions in costly environments.

In Chapters 2 and 3, I demonstrate how this approach provides novel insights into factors that shape the flexibility and robustness of task organization in harvester ant colonies (Pogonomyrmex barbatus). My results show that the degree to which colonies can respond to work in fluctuating environments depends on how individuals weigh the costs of activity and update their behavior in response to social information. In Chapter 4, I introduce a mathematical framework to study the emergence of collective organization in heterogenous groups. My approach, which is based on the theory of multi-agent systems, focuses on myopic agents whose behavior emerges out of an independent valuation of alternative choices in a given work environment. The product of this dynamic is an equilibrium organization in which agents perform different tasks (or abstain from work) with an analytically defined set of threshold probabilities. The framework is minimally developed, but can be extended to include other factors known to affect task decisions including individual experience and social facilitation. This research contributes a novel approach to developing (and analyzing) models of task organization that can be applied in a broader range of contexts where animals cooperate.
ContributorsUdiani, Oyita (Author) / Kang, Yun (Thesis advisor) / Fewell, Jennifer H (Thesis advisor) / Janssen, Marcus A (Committee member) / Castillo-Chavez, Carlos (Committee member) / Arizona State University (Publisher)
Created2016