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We examined the evolutionary morphological responses of Drosophila melanogaster that had evolved at constant cold (16°), constant hot (25°C), and fluctuating (16° and 25°C). Flies that were exposed to the constant low mean temperature developed larger thorax, wing, and cell sizes than those exposed to constant high mean temperatures. Males

We examined the evolutionary morphological responses of Drosophila melanogaster that had evolved at constant cold (16°), constant hot (25°C), and fluctuating (16° and 25°C). Flies that were exposed to the constant low mean temperature developed larger thorax, wing, and cell sizes than those exposed to constant high mean temperatures. Males and females both responded similarly to thermal treatments in average wing and cell size. The resulting cell area for a given wing size in thermal fluctuating populations remains unclear and remains a subject for future research.
ContributorsAdrian, Gregory John (Author) / Angilletta, Michael (Thesis director) / Harrison, Jon (Committee member) / Rusch, Travis (Committee member) / Barrett, The Honors College (Contributor) / School of Life Sciences (Contributor)
Created2015-05
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Studies of animal contests often focus solely on a single static measurement of fighting ability, such as the size or the strength of the individual. However, recent studies have highlighted the importance of individual variation in the dynamic behaviors used during a fight, such as, assessment strategies, decision making, and

Studies of animal contests often focus solely on a single static measurement of fighting ability, such as the size or the strength of the individual. However, recent studies have highlighted the importance of individual variation in the dynamic behaviors used during a fight, such as, assessment strategies, decision making, and fine motor control, as being strong predictors of the outcome of aggression. Here, I combined morphological and behavioral data to discover how these features interact during aggressing interactions in male virile crayfish, Faxonius virilis. I predicted that individual variation in behavioral skill for decision making (i.e., number of strikes thrown), would determine the outcome of contest success in addition to morphological measurements (e.g. body size, relative claw size). To evaluate this prediction, I filmed staged territorial interactions between male F. virilis and later analyzed trial behaviors (e.g. strike, pinches, and bout time) and aggressive outcomes. I found very little support for skill to predict win/loss outcome in trials. Instead, I found that larger crayfish engaged in aggression for longer compared to smaller crayfish, but that larger crayfish did not engage in a greater number of claw strikes or pinches when controlling for encounter duration. Future studies should continue to investigate the role of skill, by using finer-scale techniques such as 3D tracking software, which could track advanced measurements (e.g. speed, angle, and movement efficiency). Such studies would provide a more comprehensive understanding of the relative influence of fighting skill technique on territorial contests.

ContributorsNguyen, Phillip Huy (Author) / Angilletta, Michael (Thesis director) / McGraw, Kevin (Committee member) / School of Life Sciences (Contributor) / Barrett, The Honors College (Contributor)
Created2021-05
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Land management practices such as domestic animal grazing can alter plant communities via changes in soil structure and chemistry, species composition, and plant nutrient content. These changes can affect the abundance and quality of plants consumed by insect herbivores with consequent changes in population dynamics. These population changes can translate

Land management practices such as domestic animal grazing can alter plant communities via changes in soil structure and chemistry, species composition, and plant nutrient content. These changes can affect the abundance and quality of plants consumed by insect herbivores with consequent changes in population dynamics. These population changes can translate to massive crop damage and pest control costs. My dissertation focused on Oedaleus asiaticus, a dominant Asian locust, and had three main objectives. First, I identified morphological, physiological, and behavioral characteristics of the migratory ("brown") and non-migratory ("green") phenotypes. I found that brown morphs had longer wings, larger thoraxes and higher metabolic rates compared to green morphs, suggesting that developmental plasticity allows greater migratory capacity in the brown morph of this locust. Second, I tested the hypothesis of a causal link between livestock overgrazing and an increase in migratory swarms of O. asiaticus. Current paradigms generally assume that increased plant nitrogen (N) should enhance herbivore performance by relieving protein-limitation, increasing herbivorous insect populations. I showed, in contrast to this scenario, that host plant N-enrichment and high protein artificial diets decreased the size and viability of O. asiaticus. Plant N content was lowest and locust abundance highest in heavily livestock-grazed fields where soils were N-depleted, likely due to enhanced erosion and leaching. These results suggest that heavy livestock grazing promotes outbreaks of this locust by reducing plant protein content. Third, I tested for the influence of dietary imbalance, in conjunction with high population density, on migratory plasticity. While high population density has clearly been shown to induce the migratory morph in several locusts, the effect of diet has been unclear. I found that locusts reared at high population density and fed unfertilized plants (i.e. high quality plants for O. asiaticus) had the greatest migratory capacity, and maintained a high percent of brown locusts. These results did not support the hypothesis that poor-quality resources increased expression of migratory phenotypes. This highlights a need to develop new theoretical frameworks for predicting how environmental factors will regulate migratory plasticity in locusts and perhaps other insects.
ContributorsCease, Arianne (Author) / Harrison, Jon (Thesis advisor) / Elser, James (Thesis advisor) / DeNardo, Dale (Committee member) / Quinlan, Michael (Committee member) / Sabo, John (Committee member) / Arizona State University (Publisher)
Created2012
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Three populations of experimentally evolved Drosophila melanogaster populations made up of high temperature (H, constant 25 ᵒC), low temperature (C, constant 16 ᵒC) and temporal homogeneity (T, environment changes between 16 ᵒC and 25 ᵒC) were prepared and assayed to determine difference in citrate synthase activity. Between the three groups,

Three populations of experimentally evolved Drosophila melanogaster populations made up of high temperature (H, constant 25 ᵒC), low temperature (C, constant 16 ᵒC) and temporal homogeneity (T, environment changes between 16 ᵒC and 25 ᵒC) were prepared and assayed to determine difference in citrate synthase activity. Between the three groups, the results were inconclusive: the resulting reaction rates in units of nmol min-1mgfly-1 were 81.8 + 20.6, 101 + 15.6, and 96.9 + 25.2 for the hot (H), cold (C), and temporally homogeneous (T) groups, respectively. We conclude that the high associated variability was due to a lack of control regarding the collection time of the experimentally evolved Drosophila.
ContributorsBelohlavek, David (Author) / Angilletta, Michael (Thesis director) / Francisco, Wilson (Committee member) / Barrett, The Honors College (Contributor) / Department of Chemistry and Biochemistry (Contributor)
Created2015-05
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I am evaluating the genomic basis of a model of heat tolerance in which organisms succumb to warming when their demand for oxygen exceeds their supply. This model predicts that tolerance of hypoxia should correlate genetically with tolerance of heat. To evaluate this prediction, I tested heat and hypoxia tolerance

I am evaluating the genomic basis of a model of heat tolerance in which organisms succumb to warming when their demand for oxygen exceeds their supply. This model predicts that tolerance of hypoxia should correlate genetically with tolerance of heat. To evaluate this prediction, I tested heat and hypoxia tolerance in several genetic lines of Drosophila melanogaster. I hypothesized that genotypes that can fly better at high temperatures are also able to fly well at hypoxia. Genotypes from the Drosophila Genetic Reference Panel (DGRP) were assessed for flight at hypoxia and normal temperature (12% O2 and 25°C) as well as normoxia and high temperature (21% O2 and 39°C). After testing 66 lines from the DGRP, the oxygen- and capacity-limited thermal tolerance theory is supported; hypoxia-resistant lines are more likely to be heat-resistant. This supports previous research, which suggested an interaction between the tolerance of the two environmental variables. I used this data to perform a genome-wide association study to find specific single-nucleotide polymorphisms associated with heat tolerance and hypoxia tolerance but found no specific genomic markers. Understanding factors that limit an organism’s stress tolerance as well as the regions of the genome that dictate this phenotype should enable us to predict how organisms may respond to the growing threat of climate change.
ContributorsFredette-Roman, Jacob Daniel (Author) / Angilletta, Michael (Thesis director) / VandenBrooks, John (Committee member) / Youngblood, Jacob (Committee member) / School of Life Sciences (Contributor, Contributor) / Barrett, The Honors College (Contributor)
Created2020-05
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Ectotherms rely on external heat to attain target body temperatures which can vary based on the animal’s current physiological activity. Many ectotherms become thermophilic (“heat-loving”) during crucial physiological processes like digestion and reproduction, behaviorally thermoregulating to increase body temperature higher than what they otherwise prefer. However, there is a positive

Ectotherms rely on external heat to attain target body temperatures which can vary based on the animal’s current physiological activity. Many ectotherms become thermophilic (“heat-loving”) during crucial physiological processes like digestion and reproduction, behaviorally thermoregulating to increase body temperature higher than what they otherwise prefer. However, there is a positive relationship between body temperature and water loss that dictates increasing body temperature typically elicits an increase in water loss. Animals that inhabit areas where water is at least seasonally limited (e.g., deserts, wet-dry forests) may face a tradeoff between prioritizing behavioral thermophily to optimize physiological processes versus prioritizing water balance and potentially sacrificing some aspect of total performance capability.It is thus far unknown how reduced water availability and subsequent dehydration may influence thermophily in ectotherms. I hypothesized that behaviorally thermoregulating ectotherms exhibit thermophily during critical physiological events, and the extent to which thermophily is expressed is influenced by the animal’s hydric state. Using Children’s pythons (Antaresia childreni), I investigated the effects of dehydration on behavioral thermophily during digestion and reproduction. I found that dehydration caused a suppression in digestion-associated thermophily, where dehydrated snakes returned to pre-feeding body temperature sooner than they did when they were hydrated. In contrast, water deprivation at different reproductive stages had no effect on thermophily despite leading to a significant increase in the female’s plasma osmolality. ii Additionally, the timing of water deprivation during reproduction had differing effects on plasma osmolality and circulating triglyceride, total protein, and corticosterone concentrations. My research provides evidence of the sensitive and complex dynamic between body temperature, water balance, and physiological processes. At a time when many dry ecosystems are becoming hotter and drier, my investigation of dehydration and its influence on thermal dynamics and physiological metrics provides insight into cryptic effects on the vital processes of digestion and reproduction.
ContributorsAzzolini, Jill L. (Author) / Denardo, Dale F. (Thesis advisor) / John-Alder, Henry (Committee member) / Angilletta, Michael (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2023
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Understanding how and why animals choose what to eat is one of the fundamental goals of nutritional and behavioral biology. This question can be scaled to animals that live in social groups, including eusocial insects. One of the factors that plays an important role in foraging decisions is the prevalence

Understanding how and why animals choose what to eat is one of the fundamental goals of nutritional and behavioral biology. This question can be scaled to animals that live in social groups, including eusocial insects. One of the factors that plays an important role in foraging decisions is the prevalence of specific nutrients and their relative balance. This dissertation explores the role of relative nutrient content in the food selection decisions of a species that is eusocial and also agricultural, the desert leafcutter ant Acromyrmex versicolor. A dietary choice assay, in which the relative amount of protein and carbohydrates in the available diets was varied, demonstrated that A. versicolor colonies regulate relative collection of protein and carbohydrates. Tracking the foraging behavior of individual workers revelaed that foragers vary in their relative collection of experimental diets and in their foraging frequency, but that there is no relationship between these key factors of foraging behavior. The high proportion of carbohydrates preferred by lab colonies suggests that they forage to nutritionally support the fungus rather than brood and workers. To test this, the relative amounts of 1) fungus, and 2) brood (larvae) was manipulated and foraging response was measured. Changing the amount of brood had no effect on foraging. Although decreasing the size of fungus gardens did not change relative P:C collection, it produced significant increases in caloric intake, supporting the assertion that the fungus is the main driver of colony nutrient regulation. The nutritional content of naturally harvested forage material collected from field colonies was measured, as was recruitment to experimental diets with varying relative macronutrient content. Field results confirmed a strong colony preference for high carbohydrate diets. They also indicated that this species may, at times, be limited in its ability to collect sufficiently high levels of carbohydrates to meet optimal intake. This dissertation provides important insights about fundamental aspects of leafcutter ant biology and extends our understanding of the role of relative nutrient content in foraging decisions to systems that span multiple trophic levels.
ContributorsSmith, Nathan Edward (Author) / Fewell, Jennifer H (Thesis advisor) / Harrison, Jon F (Committee member) / Pavlic, Ted (Committee member) / Cease, Arianne (Committee member) / Hoelldobler, Bert (Committee member) / Arizona State University (Publisher)
Created2023
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Eusocial insect colonies have often been imagined as “superorganisms” exhibiting tight homeostasis at the colony level. However, colonies lack the tight spatial and organizational integration that many multicellular, unitary organisms exhibit. Precise regulation requires rapid feedback, which is often not possible when nestmates are distributed across space, making decisions asynchronously.

Eusocial insect colonies have often been imagined as “superorganisms” exhibiting tight homeostasis at the colony level. However, colonies lack the tight spatial and organizational integration that many multicellular, unitary organisms exhibit. Precise regulation requires rapid feedback, which is often not possible when nestmates are distributed across space, making decisions asynchronously. Thus, one should expect poorer regulation in superorganisms than unitary organisms.Here, I investigate aspects of regulation in collective foraging behaviors that involve both slow and rapid feedback processes. In Chapter 2, I examine a tightly coupled system with near-instantaneous signaling: teams of weaver ants cooperating to transport massive prey items back to their nest. I discover that over an extreme range of scenarios—even up vertical surfaces—the efficiency per transporter remains constant. My results suggest that weaver ant colonies are maximizing their total intake rate by regulating the allocation of transporters among loads. This is an exception that “proves the rule;” the ant teams are recapitulating the physical integration of unitary organisms. Next, I focus on a process with greater informational constraints, with loose temporal and spatial integration. In Chapter 3, I measure the ability of solitarily foraging Ectatomma ruidum colonies to balance their collection of protein and carbohydrates given different nutritional environments. Previous research has found that ant species can precisely collect a near-constant ratio between these two macronutrients, but I discover these studies were using flawed statistical approaches. By developing a quantitative measure of regulatory effect size, I show that colonies of E. ruidum are relatively insensitive to small differences in food source nutritional content, contrary to previously published claims. In Chapter 4, I design an automated, micro-RFID ant tracking system to investigate how the foraging behavior of individuals integrates into colony-level nutrient collection. I discover that spatial fidelity to food resources, not individual specialization on particular nutrient types, best predicts individual forager behavior. These findings contradict previously published experiments that did not use rigorous quantitative measures of specialization and confounded the effects of task type and resource location.
ContributorsBurchill, Andrew Taylor (Author) / Pavlic, Theodore P (Thesis advisor) / Pratt, Stephen C (Thesis advisor) / Hölldobler, Bert (Committee member) / Cease, Arianne (Committee member) / Berman, Spring (Committee member) / Arizona State University (Publisher)
Created2022
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By increasing the mean and variance of environmental temperatures, climate change has caused local extinctions and range shifts of numerous species. However, biologists disagree on which populations and species are most vulnerable to future warming. This debate arises because biologists do not know which physiological processes are most vulnerable to

By increasing the mean and variance of environmental temperatures, climate change has caused local extinctions and range shifts of numerous species. However, biologists disagree on which populations and species are most vulnerable to future warming. This debate arises because biologists do not know which physiological processes are most vulnerable to temperature or how to model these processes in complex environments. Using the South American locust (Schistocerca cancellata) as a model system, my dissertation addressed this debate and explained how climate limits the persistence of locust populations. Locusts of S. cancellata are serious agricultural pests with occasional outbreaks covering up to 4 million km2 over six countries. Because outbreaks are largely driven by climate, understanding how climate limits the persistence of locusts may help predict crop losses in future climates. To achieve this aim, I integrated observational, experimental, and computational approaches. First, I tested a physiological model of heat stress. By measuring the heat tolerance of locusts under different oxygen concentrations, I demonstrated that heat tolerance depends on oxygen supply during the hatchling stage only. Second, I modeled the geographic distribution of locusts using physiological traits. I started by measuring thermal effects on consumption and defecation of field-captured locusts, and I then used these data to model energy gain in current and future climates. My results indicated that incorporating physiological mechanisms can improve the accuracy of models and alter predicted impacts of climate change. Finally, I explored the causes and consequences of intraspecific variation in heat tolerance. After measuring heat tolerance of locusts in different hydration states and developmental stages, I modeled survival in historical microclimates. My models indicated that recent climate change has amplified the risk of overheating for locusts, and this risk depended strongly on shade availability, hydration state, and developmental stage. Therefore, the survival of locusts in future climates will likely depend on their access to shade and water. Overall, my dissertation argues that modeling physiological mechanisms can improve the ability of biologists to predict the impacts of climate change.
ContributorsYoungblood, Jacob (Author) / Angilletta, Michael (Thesis advisor) / Buckley, Lauren (Committee member) / Cease, Arianne (Committee member) / Smith, Brian (Committee member) / Vanden Brooks, John (Committee member) / Arizona State University (Publisher)
Created2022
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The migratory grasshopper (Melanoplus sanguinipes) is one of the most economically important grasshoppers in the western rangelands of the United States (US), capable of causing incredible amounts of damage to crops and rangelands. While M. sanguinipes has been the focus of many research studies, areas like field nutritional physiology and

The migratory grasshopper (Melanoplus sanguinipes) is one of the most economically important grasshoppers in the western rangelands of the United States (US), capable of causing incredible amounts of damage to crops and rangelands. While M. sanguinipes has been the focus of many research studies, areas like field nutritional physiology and ecology, and interactions between nutritional physiology and biopesticide resistance have very little research. This dissertation presents a multifaceted approach through three research-driven chapters that examine the nutritional physiology of M. sanguinipes and how it interacts with an entomopathogenic fungus for grasshopper management, as well as the challenges of using biopesticides for grasshopper management. Using the Geometric Framework for Nutrition (GFN), I established baseline macronutrient intake for M. sanguinipes, both in laboratory and field populations. Through this work, I found that field and lab populations can exhibit different protein (p) to carbohydrate (c) ratios, or Intake Targets (ITs), but that the field populations had ITs that matched the nutrients available in their environment. I also used the GFN to show that infections with the fungal entomopathogen Metarhizium robertsii DWR2009 did not alter ITs in M. sanguinipes. Although, when confined to carbohydrate- or protein-biased diets, infected grasshoppers had a slightly extended lifespan relative to grasshoppers fed balanced protein:carbohydrate diets. Interestingly, in a postmortem for the grasshopper, the fungus was only able to effectively sporulate on grasshoppers fed the 1p:1c diets, suggesting that grasshopper diet can have substantial impacts on the spread of fungal biopesticides throughout a population, in the absence of any inhibitory abiotic factors. Lastly, I examined the major barriers to fungal and microsporidian biopesticide usage in the United States, including low efficacy, thermal and environmental sensitivity, non-target effects, unregistered or restricted use, and economic or accessibility barriers. I also explored potential solutions to these challenges. This dissertation's focus on Melanoplus sanguinipes and Metarhizium roberstii Strain DWR2009, generates new information about how nutritional physiology and immunology intersect to impact M. sanguinipes performance. The methodology in each of the experimental chapters provides a framework for examining other problematic grasshopper species, by determining baseline nutritional physiology, and coupling nutrition with immunology to maximize the effectiveness of biological pesticides.
ContributorsZembrzuski, Deanna (Author) / Cease, Arianne (Thesis advisor) / Harrison, Jon (Committee member) / Angilletta, Michael (Committee member) / Jaronski, Stefan (Committee member) / Arizona State University (Publisher)
Created2023