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Description
Division of labor, whereby different group members perform different functions, is a fundamental attribute of sociality. It appears across social systems, from simple cooperative groups to complex eusocial colonies. A core challenge in sociobiology is to explain how patterns of collective organization are generated. Theoretical models propose that division of labor self-organizes, or emerges, from interactions among group members and the environment; division of labor is also predicted to scale positively with group size. I empirically investigated the emergence and scaling of division of labor in evolutionarily incipient groups of sweat bees and in eusocial colonies of harvester ants. To test whether division of labor is an emergent property of group living during early social evolution, I created de novo communal groups of the normally solitary sweat bee Lasioglossum (Ctenonomia) NDA-1. A division of labor repeatedly arose between nest excavation and guarding tasks; results were consistent with hypothesized effects of spatial organization and intrinsic behavioral variability. Moreover, an experimental increase in group size spontaneously promoted higher task specialization and division of labor. Next, I examined the influence of colony size on division of labor in larger, more integrated colonies of the harvester ant Pogonomyrmex californicus. Division of labor scaled positively with colony size in two contexts: during early colony ontogeny, as colonies grew from tens to hundreds of workers, and among same-aged colonies that varied naturally in size. However, manipulation of colony size did not elicit a short-term response, suggesting that the scaling of division of labor in P. californicus colonies is a product of functional integration and underlying developmental processes, rather than a purely emergent epiphenomenon. This research provides novel insights into the organization of work in insect societies, and raises broader questions about the role of size in sociobiology.
ContributorsHolbrook, Carter Tate (Author) / Fewell, Jennifer H (Thesis advisor) / Gadau, Jürgen (Committee member) / Harrison, Jon F. (Committee member) / Hölldobler, Berthold (Committee member) / Johnson, Robert A. (Committee member) / Arizona State University (Publisher)
Created2011
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. 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
Description
One hypothesis for the small size of insects relative to vertebrates, and the existence of giant fossil insects, is that atmospheric oxygen levels have constrained body sizes because oxygen delivery would be unable to match the needs of metabolically active tissues in larger insects. This study tested whether oxygen delivery becomes more challenging for larger insects by measuring the oxygen-sensitivity of flight metabolic rates and behavior during hovering for 11 different species of dragonflies that range in mass by an order of magnitude. Animals were flown in 7 different oxygen concentrations ranging from 30% to 2.5% to assess the sensitivity of their behavior and flight metabolic rates to oxygen. I also assessed the oxygen-sensitivity of flight in low-density air (nitrogen replaced with helium), to increase the metabolic demands of hovering flight. Lowered atmosphere densities did induce higher metabolic rates. Flight behaviors but not flight metabolic rates were highly oxygen-sensitive. A significant interaction between oxygen and mass was found for total flight time, with larger dragonflies varying flight time more in response to atmospheric oxygen. This study provides some support for the hypothesis that larger insects are more challenged in oxygen delivery, as predicted by the oxygen limitation hypothesis for insect gigantism in the Paleozoic.
ContributorsHenry, Joanna Randyl (Author) / Harrison, Jon F. (Thesis advisor) / Kaiser, Alexander (Committee member) / Rutowski, Ronald L (Committee member) / Arizona State University (Publisher)
Created2011
Description
The repression of reproductive competition and the enforcement of altruism are key components to the success of animal societies. Eusocial insects are defined by having a reproductive division of labor, in which reproduction is relegated to one or few individuals while the rest of the group members maintain the colony and help raise offspring. However, workers have retained the ability to reproduce in most insect societies. In the social Hymenoptera, due to haplodiploidy, workers can lay unfertilized male destined eggs without mating. Potential conflict between workers and queens can arise over male production, and policing behaviors performed by nestmate workers and queens are a means of repressing worker reproduction. This work describes the means and results of the regulation of worker reproduction in the ant species Aphaenogaster cockerelli. Through manipulative laboratory studies on mature colonies, the lack of egg policing and the presence of physical policing by both workers and queens of this species are described. Through chemical analysis and artificial chemical treatments, the role of cuticular hydrocarbons as indicators of fertility status and the informational basis of policing in this species is demonstrated. An additional queen-specific chemical signal in the Dufour's gland is discovered to be used to direct nestmate aggression towards reproductive competitors. Finally, the level of actual worker-derived males in field colonies is measured. Together, these studies demonstrate the effectiveness of policing behaviors on the suppression of worker reproduction in a social insect species, and provide an example of how punishment and the threat of punishment is a powerful force in maintaining cooperative societies.
ContributorsSmith, Adrian A. (Author) / Liebig, Juergen (Thesis advisor) / Hoelldobler, Bert (Thesis advisor) / Gadau, Juergen (Committee member) / Johnson, Robert A. (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2011
Description
At present, almost 70% of the electric energy in the United States is produced utilizing fossil fuels. Combustion of fossil fuels contributes CO2 to the atmosphere, potentially exacerbating the impact on global warming. To make the electric power system (EPS) more sustainable for the future, there has been an emphasis on scaling up generation of electric energy from wind and solar resources. These resources are renewable in nature and have pollution free operation. Various states in the US have set up different goals for achieving certain amount of electrical energy to be produced from renewable resources. The Southwestern region of the United States receives significant solar radiation throughout the year. High solar radiation makes concentrated solar power and solar PV the most suitable means of renewable energy production in this region. However, the majority of the projects that are presently being developed are either residential or utility owned solar PV plants. This research explores the impact of significant PV penetration on the steady state voltage profile of the electric power transmission system. This study also identifies the impact of PV penetration on the dynamic response of the transmission system such as rotor angle stability, frequency response and voltage response after a contingency. The light load case of spring 2010 and the peak load case of summer 2018 have been considered for analyzing the impact of PV. If the impact is found to be detrimental to the normal operation of the EPS, mitigation measures have been devised and presented in the thesis. Commercially available software tools/packages such as PSLF, PSS/E, DSA Tools have been used to analyze the power network and validate the results.
ContributorsPrakash, Nitin (Author) / Heydt, Gerald T. (Thesis advisor) / Vittal, Vijay (Thesis advisor) / Ayyanar, Raja (Committee member) / Arizona State University (Publisher)
Created2013
Description
In social insect colonies, as with individual animals, the rates of biological processes scale with body size. The remarkable explanatory power of metabolic allometry in ecology and evolutionary biology derives from the great diversity of life exhibiting a nonlinear scaling pattern in which metabolic rates are not proportional to mass, but rather exhibit a hypometric relationship with body size. While one theory suggests that the supply of energy is a major physiological constraint, an alternative theory is that the demand for energy is regulated by behavior. The central hypothesis of this dissertation research is that increases in colony size reduce the proportion of individuals actively engaged in colony labor with consequences for energetic scaling at the whole-colony level of biological organization. A combination of methods from comparative physiology and animal behavior were developed to investigate scaling relationships in laboratory-reared colonies of the seed-harvester ant, Pogonomyrmex californicus. To determine metabolic rates, flow-through respirometry made it possible to directly measure the carbon dioxide production and oxygen consumption of whole colonies. By recording video of colony behavior, for which ants were individually paint-marked for identification, it was possible to reconstruct the communication networks through which information is transmitted throughout the colony. Whole colonies of P. californicus were found to exhibit a robust hypometric allometry in which mass-specific metabolic rates decrease with increasing colony size. The distribution of walking speeds also scaled with colony size so that larger colonies were composed of relatively more inactive ants than smaller colonies. If colonies were broken into random collections of workers, metabolic rates scaled isometrically, but when entire colonies were reduced in size while retaining functionality (queens, juveniles, workers), they continued to exhibit a metabolic hypometry. The communication networks in P. californicus colonies contain a high frequency of feed-forward interaction patterns consistent with those of complex regulatory systems. Furthermore, the scaling of these communication pathways with size is a plausible mechanism for the regulation of whole-colony metabolic scaling. The continued development of a network theory approach to integrating behavior and metabolism will reveal insights into the evolution of collective animal behavior, ecological dynamics, and social cohesion.
ContributorsWaters, James S., 1983- (Author) / Harrison, Jon F. (Thesis advisor) / Quinlan, Michael C. (Committee member) / Pratt, Stephen C. (Committee member) / Fewell, Jennifer H. (Committee member) / Gadau, Juergen (Committee member) / Arizona State University (Publisher)
Created2012
Description
Many species follow networked roads. When roads are blocked, the obstruction must be circumnavigated, or traffic rerouted. We obstructed trails of the leaf-cutting ant Atta colombica and compared individual- and group-level circumnavigation as well as trail reuse following obstruction removal. Groups that circumnavigated the obstruction fastest were also the first to return to the original trail once the obstruction was removed. Also, nestward ants returned to using the original trail more quickly than outbound ants. Traffic rate was not related to speed of obstacle solving. The magnitude of reflective flow (reversing direction) explained much of the variation in obstacle-solving time, both comparing nestward versus outbound ants and variation across obstacles. Two other factors explaining variation in obstacle circumnavigation times were percentage of nestward ants carrying leaves and whether ants searched in the appropriate direction for the trail beyond the obstruction, possibly due to variation in the availability of navigation cues or motivation. Reflective flow allows highly-networked leafcutter trails to respond to blockages by using alternative cleared routes, with strength of navigation cues and motivation linked to foraging costs and benefits likely determining the effort expended to “solve” the obstacle versus give up.
ContributorsPrendergast, Catherine T (Author) / Harrison, Jon F. (Thesis director) / Baudier, Kaitlin M. (Committee member) / Economics Program in CLAS (Contributor) / Department of English (Contributor) / Barrett, The Honors College (Contributor)
Created2020-05
Description
The retina is the lining in the back of the eye responsible for vision. When light photons hits the retina, the photoreceptors within the retina respond by sending impulses to the optic nerve, which connects to the brain. If there is injury to the eye or heredity retinal problems, this part can become detached. Detachment leads to loss of nutrients, such as oxygen and glucose, to the cells in the eye and causes cell death. Sometimes the retina is able to be surgically reattached. If the photoreceptor cells have not died and the reattachment is successful, then these cells are able to regenerate their outer segments (OS) which are essential for their functionality and vitality. In this work we will explore how the regrowth of the photoreceptor cells in a healthy eye after retinal detachment can lead to a deeper understanding of how eye cells take up nutrients and regenerate. This work uses a mathematical model for a healthy eye in conjunction with data for photoreceptors' regrowth and decay. The parameters for the healthy eye model are estimated from the data and the ranges of these parameter values are centered +/- 10\% away from these values are used for sensitivity analysis. Using parameter estimation and sensitivity analysis we can better understand how certain processes represented by these parameters change within the model as a result of retinal detachment. Having a deeper understanding for any sort of photoreceptor death and growth can be used by the greater scientific community to help with these currently irreversible conditions that lead to blindness, such as retinal detachment. The analysis in this work shows that maximizing the carrying capacity of the trophic pool and the rate of RDCVF, as well as minimizing nutrient withdrawal of the rods and the cones from the trophic pool results in both the most regrowth and least cell death in retinal detachment.
ContributorsGoldman, Miriam Ayla (Author) / Camacho, Erikia (Thesis director) / Wirkus, Stephen (Committee member) / School of Mathematical and Natural Sciences (Contributor, Contributor) / Barrett, The Honors College (Contributor)
Created2017-05
Description
One hypothesis for why insects are smaller than vertebrates is that the blind-ended tracheal respiratory system challenges oxygen delivery for larger insects. Supporting this hypothesis, several studies have documented that larger insect species have larger gas transport structures than expected by isometric scaling. To further test this hypothesis, we performed the first inter-specific study of the scaling of spiracle size, using ten scarab beetle species, including some of the most massive insects. Using micro-CT, we measured the cross sectional area and depth of all eight spiracles. Areas of large spiracles in the anterior portion of the animal showed hypermetric scaling, varying approximately with mass^0.8. However, because diffusive capacities scaled with lower slopes than metabolic rates, larger beetles had a 10-fold higher required PO2 gradient across the spiracles to sustain oxygen consumption by diffusion. Despite this trend, calculations suggest that large beetles can exchange oxygen by diffusion across the spiracles at rest, but likely no beetles can do so during flight. Advective capacities through the spiracles scale with mass^1.8, suggestive of a switch toward greater use of convection and/or reduced required pressures in larger beetles.
ContributorsWagner, Julian Morgan (Author) / Harrison, Jon F. (Thesis director) / VandenBrooks, John (Committee member) / Miller, Laura (Committee member) / School of Mathematical and Statistical Sciences (Contributor) / School of Life Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2017-05
Description
Deserts are natural laboratories in which to study the acute effects of extreme heat and aridity on animal physiology, as well as the physiological adaptations that these animals develop to survive. For small, endothermic fliers in the desert, heat balance and water balance are challenging due to high surface area to volume ratios and the additional heat load imposed as a result of endothermy. Much of the previous fifty years of thermo- and hydroregulation research has focused on larger, charismatic megafauna; extremophiles; or only part of the heat or water balance story. Here, I calculate the first heat budget for an important desert bee, Centris caesalpiniae. As is common in many mammals, avians, and other flying insects, I find that C. caesalpiniae males use an appendage – in this case the abdomen – as a convective radiator to dispel excess heat produced by the thoracic flight muscles at high air temperatures. The few heat budgets measured for flying endotherms are conducted in the shade so as to eliminate the effects of solar radiation. To further refine the accuracy of my heat budget model, I consider how heat gain from solar radiation affects the thermal balance of flying desert endotherms. To this effect, I find that solar radiation contributes 43 to 54% of the total heat gain of a desert Centris pallida bee. I additionally show that large morph male, small morph male, and female C. pallida, use different thermoregulatory tactics while flying in shaded versus sunny conditions; large males and females rely on the abdominal convector in the sun while small morph males increase convective conductance in the sun, but do not use an abdominal convector. Given that evaporative cooling was not a significant part of the heat budget for neither C. caesalpiniae nor C. pallida, I investigated the effects of water loss rates and critical thermal maxima during flight on duration of activity period. I found that male C. caesalpiniae limited their activity period due to high water loss rates rather than overheating, and that Centris critical water contents ranged from 48 to 54%, limiting flight activity to about 3 hours.
ContributorsJohnson, Meredith Grace (Author) / Harrison, Jon F. (Thesis advisor) / Buchmann, Stephen (Committee member) / DeNardo, Dale (Committee member) / Dillon, Michael (Committee member) / Arizona State University (Publisher)
Created2023