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Sexual and social signals have long been thought to play an important role in speciation and diversity; hence, investigations of intraspecific communication may lead to important insights regarding key processes of evolution. Though we have learned much about the control, function, and evolution of animal communication by studying several very

Sexual and social signals have long been thought to play an important role in speciation and diversity; hence, investigations of intraspecific communication may lead to important insights regarding key processes of evolution. Though we have learned much about the control, function, and evolution of animal communication by studying several very common signal types, investigating rare classes of signals may provide new information about how and why animals communicate. My dissertation research focused on rapid physiological color change, a rare signal-type used by relatively few taxa. To answer longstanding questions about this rare class of signals, I employed novel methods to measure rapid color change signals of male veiled chameleons Chamaeleo calyptratus in real-time as seen by the intended conspecific receivers, as well as the associated behaviors of signalers and receivers. In the context of agonistic male-male interactions, I found that the brightness achieved by individual males and the speed of color change were the best predictors of aggression and fighting ability. Conversely, I found that rapid skin darkening serves as a signal of submission for male chameleons, reducing aggression from winners when displayed by losers. Additionally, my research revealed that the timing of maximum skin brightness and speed of brightening were the best predictors of maximum bite force and circulating testosterone levels, respectively. Together, these results indicated that different aspects of color change can communicate information about contest strategy, physiology, and performance ability. Lastly, when I experimentally manipulated the external appearance of chameleons, I found that "dishonestly" signaling individuals (i.e. those whose behavior did not match their manipulated color) received higher aggression from unpainted opponents. The increased aggression received by dishonest signalers suggests that social costs play an important role in maintaining the honesty of rapid color change signals in veiled chameleons. Though the color change abilities of chameleons have interested humans since the time of Aristotle, little was previously known about the signal content of such changes. Documenting the behavioral contexts and information content of these signals has provided an important first step in understanding the current function, underlying control mechanisms, and evolutionary origins of this rare signal type.
ContributorsLigon, Russell (Author) / McGraw, Kevin J. (Committee member) / DeNardo, Dale F (Committee member) / Karsten, Kristopher B (Committee member) / Rutowski, Ronald L (Committee member) / Deviche, Pierre (Committee member) / Arizona State University (Publisher)
Created2015
Description
Conditions during development can shape the expression of traits at adulthood, a phenomenon called developmental plasticity. In this context, factors such as nutrition or health state during development can affect current and subsequent physiology, body size, brain structure, ornamentation, and behavior. However, many of the links between developmental and adult

Conditions during development can shape the expression of traits at adulthood, a phenomenon called developmental plasticity. In this context, factors such as nutrition or health state during development can affect current and subsequent physiology, body size, brain structure, ornamentation, and behavior. However, many of the links between developmental and adult phenotype are poorly understood. I performed a series of experiments using a common molecular currency - carotenoid pigments - to track somatic and reproductive investments through development and into adulthood. Carotenoids are red, orange, or yellow pigments that: (a) animals must acquire from their diets, (b) can be physiologically beneficial, acting as antioxidants or immunostimulants, and (c) color the sexually attractive features (e.g., feathers, scales) of many animals. I studied how carotenoid nutrition and immune challenges during ontogeny impacted ornamental coloration and immune function of adult male mallard ducks (Anas platyrhynchos). Male mallards use carotenoids to pigment their yellow beak, and males with more beaks that are more yellow are preferred as mates, have increased immune function, and have higher quality sperm. In my dissertation work, I established a natural context for the role that carotenoids and body condition play in the formation of the adult phenotype and examined how early-life experiences, including immune challenges and dietary access to carotenoids, affect adult immune function and ornamental coloration. Evidence from mallard ducklings in the field showed that variation in circulating carotenoid levels at hatch are likely driven by maternal allocation of carotenoids, but that carotenoid physiology shifts during the subsequent few weeks to reflect individual foraging habits. In the lab, adult beak color expression and immune function were more tightly correlated with body condition during growth than body condition during subsequent stages of development or adulthood. Immune challenges during development affected adult immune function and interacted with carotenoid physiology during adulthood, but did not affect adult beak coloration. Dietary access to carotenoids during development, but not adulthood, also affected adult immune function. Taken together, these results highlight the importance of the developmental stage in shaping certain survival-related traits (i.e., immune function), and lead to further questions regarding the development of ornamental traits.
ContributorsButler, Michael (Author) / McGraw, Kevin J. (Thesis advisor) / Chang, Yung (Committee member) / Deviche, Pierre (Committee member) / DeNardo, Dale (Committee member) / Rutowski, Ronald (Committee member) / Arizona State University (Publisher)
Created2012
Description
Glioblastoma (GBM) is the most common primary brain tumor with an incidence of approximately 11,000 Americans. Despite decades of research, average survival for GBM patients is a modest 15 months. Increasing the extent of GBM resection increases patient survival. However, extending neurosurgical margins also threatens the removal of eloquent brain.

Glioblastoma (GBM) is the most common primary brain tumor with an incidence of approximately 11,000 Americans. Despite decades of research, average survival for GBM patients is a modest 15 months. Increasing the extent of GBM resection increases patient survival. However, extending neurosurgical margins also threatens the removal of eloquent brain. For this reason, the infiltrative nature of GBM is an obstacle to its complete resection. We hypothesize that targeting genes and proteins that regulate GBM motility, and developing techniques that safely enhance extent of surgical resection, will improve GBM patient survival by decreasing infiltration into eloquent brain regions and enhancing tumor cytoreduction during surgery. Chapter 2 of this dissertation describes a gene and protein we identified; aquaporin-1 (aqp1) that enhances infiltration of GBM. In chapter 3, we describe a method for enhancing the diagnostic yield of GBM patient biopsies which will assist in identifying future molecular targets for GBM therapies. In chapter 4 we develop an intraoperative optical imaging technique that will assist identifying GBM and its infiltrative margins during surgical resection. The topic of this dissertation aims to target glioblastoma infiltration from molecular and cellular biology and neurosurgical disciplines. In the introduction we; 1. Provide a background of GBM and current therapies. 2. Discuss a protein we found that decreases GBM survival. 3. Describe an imaging modality we utilized for improving the quality of accrued patient GBM samples. 4. We provide an overview of intraoperative contrast agents available for neurosurgical resection of GBM, and discuss a new agent we studied for intraoperative visualization of GBM.
ContributorsGeorges, Joseph F (Author) / Feuerstein, Burt G (Thesis advisor) / Smith, Brian H. (Thesis advisor) / Van Keuren-Jensen, Kendall (Committee member) / Deviche, Pierre (Committee member) / Bennett, Kevin (Committee member) / Arizona State University (Publisher)
Created2014
Description
Parental care provides many benefits to offspring. One widely realized benefit is enhanced regulation of offspring's thermal environment. The developmental thermal environment during development can be optimized behaviorally through nest site selection and brooding, and it can be further enhanced by physiological heat production. In fact, enhancement of the developmental

Parental care provides many benefits to offspring. One widely realized benefit is enhanced regulation of offspring's thermal environment. The developmental thermal environment during development can be optimized behaviorally through nest site selection and brooding, and it can be further enhanced by physiological heat production. In fact, enhancement of the developmental thermal environment has been proposed as the initial driving force for the evolution of endothermy in bird and mammals. I used pythons (Squamata: Pythonidae) to expand existing knowledge of behavioral and physiological parental tactics used to regulate offspring thermal environment. I first demonstrated that brooding behavior in the Children's python (Antaresia childreni) is largely driven by internal mechanisms, similar to solitary birds, suggesting that the early evolution of the parent-offspring association was probably hormonally driven. Two species of python are known to be facultatively thermogenic (i.e., are endothermic during reproduction). I expand current knowledge of thermogenesis in Burmese pythons (Python molurus) by demonstrating that females use their own body temperature to modulate thermogenesis. Although pythons are commonly cited as thermogenic, the actual extent of thermogenesis within the family Pythonidae is unknown. Thus, I assessed the thermogenic capability of five previously unstudied species of python to aid in understanding phylogenetic, morphological, and distributional influences on thermogenesis in pythons. Results suggest that facultative thermogenesis is likely rare among pythons. To understand why it is rare, I used an artificial model to demonstrate that energetic costs to the female likely outweigh thermal benefits to the clutch in species that do not inhabit cooler latitudes or lack large energy reserves. In combination with other studies, these results show that facultative thermogenesis during brooding in pythons likely requires particular ecological and physiological factors for its evolution.
ContributorsBrashears, Jake (Author) / DeNardo, Dale (Thesis advisor) / Harrison, Jon (Committee member) / Deviche, Pierre (Committee member) / McGraw, Kevin (Committee member) / Smith, Andrew (Committee member) / Arizona State University (Publisher)
Created2012
Description
For animals that experience annual cycles of gonad development, the seasonal timing (phenology) of gonad growth is a major adaptation to local environmental conditions. To optimally time seasonal gonad growth, animals use environmental cues that forecast future conditions. The availability of food is one such environmental cue. Although the importance

For animals that experience annual cycles of gonad development, the seasonal timing (phenology) of gonad growth is a major adaptation to local environmental conditions. To optimally time seasonal gonad growth, animals use environmental cues that forecast future conditions. The availability of food is one such environmental cue. Although the importance of food availability has been appreciated for decades, the physiological mechanisms underlying the modulation of seasonal gonad growth by this environmental factor remain poorly understood.

Urbanization is characterized by profound environmental changes, and urban animals must adjust to an environment vastly different from that of their non-urban conspecifics. Evidence suggests that birds adjust to urban areas by advancing the timing of seasonal breeding and gonad development, compared to their non-urban conspecifics. A leading hypothesis to account for this phenomenon is that food availability is elevated in urban areas, which improves the energetic status of urban birds and enables them to initiate gonad development earlier than their non-urban conspecifics. However, this hypothesis remains largely untested.

My dissertation dovetailed comparative studies and experimental approaches conducted in field and captive settings to examine the physiological mechanisms by which food availability modulates gonad growth and to investigate whether elevated food availability in urban areas advances the phenology of gonad growth in urban birds. My captive study demonstrated that energetic status modulates reproductive hormone secretion, but not gonad growth. By contrast, free-ranging urban and non-urban birds did not differ in energetic status or plasma levels of reproductive hormones either in years in which urban birds had advanced phenology of gonad growth or in a year that had no habitat-related disparity in seasonal gonad growth. Therefore, my dissertation provides no support for the hypothesis that urban birds begin seasonal gonad growth because they are in better energetic status and increase the secretion of reproductive hormones earlier than non-urban birds. My studies do suggest, however, that the phenology of key food items and the endocrine responsiveness of the reproductive system may contribute to habitat-related disparities in the phenology of gonad growth.
ContributorsDavies, Scott (Author) / Deviche, Pierre (Thesis advisor) / Sweazea, Karen (Committee member) / McGraw, Kevin (Committee member) / Orchinik, Miles (Committee member) / Warren, Paige (Committee member) / Arizona State University (Publisher)
Created2014
Description
Colorful ornaments in animals often serve as sexually selected signals of quality. While pigment-based colors are well-studied in these regards, structural colors that result from the interaction of light with photonic nanostructures are comparatively understudied in terms of their consequences in social contexts, their costs of production, and even the

Colorful ornaments in animals often serve as sexually selected signals of quality. While pigment-based colors are well-studied in these regards, structural colors that result from the interaction of light with photonic nanostructures are comparatively understudied in terms of their consequences in social contexts, their costs of production, and even the best way to measure them. Iridescent colors are some of the most brilliant and conspicuous colors in nature, and I studied the measurement, condition-dependence, and signaling role of iridescence in Anna's hummingbirds (Calypte anna). While most animal colors are easily quantified using well-established spectrophotometric techniques, the unique characteristics of iridescent colors present challenges to measurement and opportunities to quantify novel color metrics. I designed and tested an apparatus for careful control and measurement of viewing geometry and highly repeatable measurements. These measurements could be used to accurately characterize individual variation in iridescent Anna's hummingbirds to examine their condition-dependence and signaling role. Next, I examined the literature published to date for evidence of condition-dependence of structural colors in birds. Using meta-analyses, I found that structural colors of all three types - white, ultra-violet/blue, and iridescence - are significantly condition-dependent, meaning that they can convey information about quality to conspecifics. I then investigated whether iridescent colors were condition-dependent in Anna's hummingbirds both in a field correlational study and in an experimental study. Throughout the year, I found that iridescent feathers in both male and female Anna's hummingbirds become less brilliant as they age. Color was not correlated with body condition in any age/sex group. However, iridescent coloration in male Anna's hummingbirds was significantly affected by experimental protein in the diet during feather growth, indicating that iridescent color may signal diet quality. Finally, I examined how iridescent colors were used to mediate social competitions in male and female Anna's hummingbirds. Surprisingly, males that were less colorful won significantly more contests than more colorful males, and colorful males received more aggression. Less colorful males may be attempting to drive away colorful neighbors that may be preferred mates. Female iridescent ornament size and color was highly variable, but did not influence contest outcomes or aggression.
ContributorsMeadows, Melissa (Author) / McGraw, Kevin J. (Thesis advisor) / Rutowski, Ronald L (Committee member) / Sabo, John L (Committee member) / Alcock, John (Committee member) / Deviche, Pierre (Committee member) / Arizona State University (Publisher)
Created2012
Description
The elaborate signals of animals are often costly to produce and maintain, thus communicating reliable information about the quality of an individual to potential mates or competitors. The properties of the sensory systems that receive signals can drive the evolution of these signals and shape their form and function. However,

The elaborate signals of animals are often costly to produce and maintain, thus communicating reliable information about the quality of an individual to potential mates or competitors. The properties of the sensory systems that receive signals can drive the evolution of these signals and shape their form and function. However, relatively little is known about the ecological and physiological constraints that may influence the development and maintenance of sensory systems. In the house finch (Carpodacus mexicanus) and many other bird species, carotenoid pigments are used to create colorful sexually selected displays, and their expression is limited by health and dietary access to carotenoids. Carotenoids also accumulate in the avian retina, protecting it from photodamage and tuning color vision. Analogous to plumage carotenoid accumulation, I hypothesized that avian vision is subject to environmental and physiological constraints imposed by the acquisition and allocation of carotenoids. To test this hypothesis, I carried out a series of field and captive studies of the house finch to assess natural variation in and correlates of retinal carotenoid accumulation and to experimentally investigate the effects of dietary carotenoid availability, immune activation, and light exposure on retinal carotenoid accumulation. Moreover, through dietary manipulations of retinal carotenoid accumulation, I tested the impacts of carotenoid accumulation on visually mediated foraging and mate choice behaviors. My results indicate that avian retinal carotenoid accumulation is variable and significantly influenced by dietary carotenoid availability and immune system activity. Behavioral studies suggest that retinal carotenoid accumulation influences visual foraging performance and mediates a trade-off between color discrimination and photoreceptor sensitivity under dim-light conditions. Retinal accumulation did not influence female choice for male carotenoid-based coloration, indicating that a direct link between retinal accumulation and sexual selection for coloration is unlikely. However, retinal carotenoid accumulation in males was positively correlated with their plumage coloration. Thus, carotenoid-mediated visual health and performance or may be part of the information encoded in sexually selected coloration.
ContributorsToomey, Matthew (Author) / McGraw, Kevin J. (Thesis advisor) / Deviche, Pierre (Committee member) / Smith, Brian (Committee member) / Rutowski, Ronald (Committee member) / Verrelli, Brian (Committee member) / Arizona State University (Publisher)
Created2011
Description
The impact of urbanization on wildlife is becoming an important topic in conservation. However little is known concerning the proximate mechanisms involved which enable some species to persist in cities, while others perish. Adapting to novel city environments requires individuals to maintain a functional physiological response to stressful stimuli, while

The impact of urbanization on wildlife is becoming an important topic in conservation. However little is known concerning the proximate mechanisms involved which enable some species to persist in cities, while others perish. Adapting to novel city environments requires individuals to maintain a functional physiological response to stressful stimuli, while concurrently using the necessary resources (food) needed to persist. A primary function of the stress response is the mobilization of intrinsic energy resources, and thus both requirements (energy and stress) are explicably linked. This dissertation investigates the interaction of energetic reserves and the physiological stress response in a native bird species, the Curve-billed Thrasher, within the context of this species' colonization of Phoenix, Arizona. This research uses a combination of comparative studies, statistical modeling, and experimental approaches conducted in field and captive settings to demonstrate how urban and desert populations of these species differ in energetic state and stress physiology. These studies reveal that the current energetic status of an individual bird influences the secretion of glucocorticoids (primary stress hormones) and can alter how energy reserves are used for gluconeogenesis to produce energy during acute stress. In addition, this research also identifies how differing levels of a hypothalamic neuropeptide (vasotocin) may play a role in mediating differences in stress physiology between populations. The quantity of food available and even temporal variability in its abundance may alter how native birds respond to stress. Increased body condition offsets the costs of maintaining the stress response in urban areas.
ContributorsFokidis, Haralambos Bobby (Author) / Deviche, Pierre (Thesis advisor) / Arizona State University (Publisher)
Created2010
Description
Organisms regularly face the challenge of having to accumulate and allocate limited resources toward life-history traits. However, direct quantification of how resources are accumulated and allocated is rare. Carotenoids are among the best systems for investigating resource allocation, because they are diet-derived and multi-functional. Birds have been studied extensively with

Organisms regularly face the challenge of having to accumulate and allocate limited resources toward life-history traits. However, direct quantification of how resources are accumulated and allocated is rare. Carotenoids are among the best systems for investigating resource allocation, because they are diet-derived and multi-functional. Birds have been studied extensively with regard to carotenoid allocation towards life-history traits, but direct quantification of variation in carotenoid distribution on a whole-organism scale has yet to be done. Additionally, while we know that scavenger receptor B1 (SCARB1) is important for carotenoid absorption in birds, little is known about the factors that predict how SCARB1 is expressed in wild populations. For my dissertation, I first reviewed challenges associated with statistically analyzing tissue distributions of nutrients (nutrient profiles) and tested how tissue carotenoid distributions (carotenoid profiles) varied by sex, season, health state, and coloration in two bird species, house finches (Haemorhous mexicanus) and zebra finches (Taeniopygia guttata). Then, I investigated the relationship between dietary carotenoid availability, relative expression of SCARB1, and extent of carotenoid-based coloration in a comparative study of wood-warblers (Parulidae). In my review of studies analyzing nutrient profiles, I found that multivariate analyses were the most common, but studies rarely reported intercorrelations among nutrient types. In house finches, all tissue carotenoid profiles varied by sex, season, and coloration. For example, males during autumn (molt) had higher concentrations of 3-hydroxyechinenone (the major red carotenoid in sexually attractive male feathers) in most but not all tissues compared to other season and sex combinations. However, the relationship between color and carotenoid profiles depended on the color metric. In zebra finches, only muscle and spleen carotenoid profiles varied between immune-challenged and control birds. In wood-warblers, I found that capacity to absorb carotenoids was positively correlated with the evolution of carotenoid-based coloration but negatively associated with liver carotenoid accumulation. Altogether, my dissertation illustrates (a) the context-dependence of tissue carotenoid profile variation, (b) that carotenoid-based integumentary coloration is a reflection of tissue carotenoid profiles, and (c) that digestive physiology (e.g., carotenoid absorption) is an important consideration in the study of diet and coloration in wild birds.
ContributorsWebb, Emily (Author) / McGraw, Kevin J (Thesis advisor) / Deviche, Pierre (Committee member) / Martins, Emilia (Committee member) / Sweazea, Karen (Committee member) / Arizona State University (Publisher)
Created2021
Description
Biodiversity is required to guarantee proper ecosystem structure and function. However, increasing anthropogenic threats are causing biodiversity loss around the world at an unprecedented rate, in what has been deemed the sixth mass extinction. To counteract this crisis, conservationists seek to improve the methods used in the design and implementation

Biodiversity is required to guarantee proper ecosystem structure and function. However, increasing anthropogenic threats are causing biodiversity loss around the world at an unprecedented rate, in what has been deemed the sixth mass extinction. To counteract this crisis, conservationists seek to improve the methods used in the design and implementation of protected areas, which help mitigate the impacts of human activities on species. Marine mammals are ecosystem engineers and important indicator species of ocean and human wellbeing. They are also disproportionally less known and more threatened than terrestrial mammals. Therefore, surrogates of biodiversity must be used to maximize their representation in conservation planning. Some of the most effective surrogates of biodiversity known have only been tested in terrestrial systems. Here I test complementarity, rarity, and environmental diversity as potential surrogates of marine mammal representation at the global scale, and compare their performance against species richness, which is the most popular surrogate used to date. I also present the first map of marine mammal complementarity, and assess its relationship with environmental variables to determine if environmental factors could also be used as surrogates. Lastly, I determine the global complementarity-based hotspots of marine mammal biodiversity, and compare their distributions against current marine protected area coverage and exposure to global indices of human threats, to elucidate the effectiveness of current conservation efforts. Results show that complementarity, rarity, and environmental diversity are all efficient surrogates, as they outcompete species richness in maximizing marine mammal species representation when solving the minimum-set coverage problem. Results also show that sea surface temperature, density, and bathymetry are the top environmental variables most associated with complementarity of marine mammals. Finally, gap analyses show that marine mammals are overall poorly protected, yet moderately exposed to hotspots of cumulative human impacts. The wide distribution of marine mammals justify global studies like the ones here presented, to determine the best strategy for their protection. Overall, my findings show that less popular surrogates of biodiversity are more effective for marine mammals and should be considered in their management, and that the expansion of protected areas in their most important habitats should be prioritized.
ContributorsAstudillo-Scalia, Yaiyr (Author) / Albuquerque, Fabio (Thesis advisor) / Deviche, Pierre (Thesis advisor) / Polidoro, Beth (Committee member) / Kellar, Nicholas (Committee member) / Arizona State University (Publisher)
Created2019