Graduate Theses & Dissertations

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effects of heat dissipation capacity on avian physiology and behaviour
In endotherms, physiological functioning is optimized within a narrow range of tissue temperatures, meaning that the capacity to dissipate body heat is an important parameter for thermoregulation and organismal performance. Yet, experimental research has found mixed support for the importance of heat dissipation capacity as a constraint on reproductive performance. To investigate the effects of heat dissipation capacity on organismal performance, I experimentally manipulated heat dissipation capacity in free-living tree swallows, Tachycineta bicolor, by trimming feathers overlying the brood patch, and monitored parental provisioning performance, body temperature, and offspring growth. I found that individuals with an enhanced capacity to dissipate body heat (i.e., trimmed treatment) provisioned their offspring more frequently, and reared larger offspring that fledged more consistently. Although control birds typically reduced their nestling provisioning rate at the highest ambient temperatures to avoid overheating, at times they became hyperthermic. Additionally, I examined inter-individual variation in body temperature within each treatment, and discovered that body temperature is variable among all individuals. This variability is also consistent over time (i.e., is repeatable), irrespective of treatment. Further, I found that individuals consistently differed in how they adjusted their body temperature across ambient temperature, demonstrating that body temperature is a flexible and repeatable physiological trait. Finally, I used a bacterial endotoxin (lipopolysaccharide) to examine the regulation of body temperature of captive zebra finches (Taeniopygia guttata) during an immune challenge. Exposure to lipopolysaccharide induces sickness behaviours, and results in a fever, hypothermia, or a combination of the two, depending on species and dosage. I asked what the relative role of different regions of the body (bill, eye region, and leg) is in heat dissipation/retention during the sickness-induced body temperature response. I found that immune-challenged individuals modulated their subcutaneous temperature primarily through alterations in peripheral blood flow, particularly in the legs and feet, detectable as a drop in surface temperature. These results demonstrate that the importance of regional differences in regulating body temperature in different contexts. Taken together, my thesis demonstrates that heat dissipation capacity can affect performance and reproductive success in birds. Author Keywords: body temperature, heat dissipation, tree swallow, zebra finch
Comparison of Dehydration Techniques for Acute Weight Management in Rowing
Mild sauna dehydration and fluid abstinence were investigated as weight loss strategies for lightweight rowers. Rowers (N=12) performed a power test, an incremental VO2max test, and a visuomotor battery: once euhydrated, once following sauna dehydration (SAU), and once following fluid abstinence and then sauna dehydration (FA). The percent body mass change (%BMC) achieved, %BMC attributable to sauna dehydration, and %BMC attributable to fluid abstinence were used within linear mixed effects models to predict hydration and performance variables. Sauna and overnight dehydration exerted indistinguishable effects on plasma osmolality, urine osmolality and thirst (p > .05). Fluid abstinence but not sauna dehydration was related to lower power production on the power test (b = 12.14W / 1%BMC, FA = 673.46 ± 79.50, SAU = 683.33 ± 72.08, p = .029), a lower total wattage produced on the incremental VO2max test (b = 4261.51W / 1%BMC, FA = 71029.58 ± 16256.56, SAU = 74001.50 ± 14936.56, p = .006), lower wattages at 2 mmol/L (b = 27.84W / 1%BMC, FA = 180.74 ± 40.27, SAU = 190.82 ± 50.79, p < .001) and 4 mmol/L (b = 20.45W / 1%BMC, FA = 221.90 ± 52.62, SAU = 238.89 ± 40.78, p = .002) blood lactate, and slower movement time on a visuomotor task (b = -38.06ms / 1%BMC, p = .004). Mild fluid abstinence but not sauna dehydration reduces rowing performance when two-hour rehydration is allowed. Author Keywords: crew, fluid, hydration, lightweight, sauna, weight
Contemporary adaptive shifts in the physiology and life history of Pumpkinseed (Lepomis gibbosus) introduced into a warm climate
Contemporary evolution has the potential to help limit the biological impact of rapidly changing climates, however it remains unclear whether wild populations can respond quickly enough for such adaptations to be effective. In this thesis, I used the introduction of native North American Pumpkinseed (Lepomis gibbosus) into the milder climate of Europe over 140 years ago, as a 'natural' experiment to test for contemporary evolution to a change in climate in wild populations. In 2008, four outdoor pond colonies were established in central Ontario using adult Pumpkinseed from two native Canadian populations, and two non-native populations from northeastern Spain. By raising native and non-native Pumpkinseed within a common environment, this design minimized the impact of phenotypic plasticity on differential trait expression, and allowed me to interpret differences in the phenotype among pond-reared Pumpkinseed as evidence of genetic differences among populations. I demonstrated that Canadian and Spanish Pumpkinseed have similar thermal physiology except when acclimated to seasonally warm temperatures; trait differences are consistent with Spanish Pumpkinseed being better adapted to a warmer climate. Populations also had similar overwintering ecology, however some differences, such as higher survival under starvation conditions and greater energetic benefits associated with winter feeding, indicated that Canadian populations are better adapted to harsh winter conditions typical of the native range. Finally, I determined that the relatively fast life history expressed in wild European Pumpkinseed is largely driven by plastic responses to the local environment; however, the higher reproductive investment by European populations has a genetic basis. Most climate change research considers taxa that are expected to be negatively impacted by warming: my research demonstrates that even warm-tolerant taxa that are unlikely to experience strong climatic selective forces can respond to a warming environment through evolutionary changes. The potential for adaptive contemporary evolution in warm-tolerant taxa should be taken into account when predicting future ecosystem effects of climate change, and when planning management strategies for species introduced into novel climates. Author Keywords: climate change, contemporary evolution, fish, non-native species, thermal biology, winter ecology
role of corticosterone in breeding effort and reproductive success in tree swallows
Glucocorticoids (e.g., corticosterone (CORT)) are hypothesized to mediate decisions regarding reproductive investment during breeding, but the directionality of the relationship is not clear. The CORT-fitness hypothesis posits that high levels of CORT arise from challenging environmental conditions in which an individual will conserve resources for future reproduction or self-maintenance, and thus result in lower reproductive success (a negative relationship). In contrast, the CORT-adaptation hypothesis suggests that, during energetically demanding periods, CORT will mediate physiological or behavioural changes that result in increased reproductive investment and success (a positive relationship). Inconsistencies arise due to the various species and life-history stages studied, and the complex interactions between fitness and glucocorticoids. Using an experimental approach, I investigated the relationship between CORT and reproductive success by manipulating baseline CORT levels in female tree swallows Tachycineta bicolor prior to laying using silastic implants. Implants failed to raise CORT levels of females during either incubation or the nestling stage, and maternal treatment had no effect on indices of fitness at either stage. Using a correlative approach, partial support for the CORT-adaptation hypothesis was found: There was a positive relationship between CORT and hatching success. This only occurred when CORT was measured during incubation, when baseline CORT levels may stimulate increased reproductive effort and success. In contrast, during the nestling stage baseline CORT levels were not related to reproductive investment or success. Maternal CORT levels during incubation also did not influence nestling phenotype, although nestling stress CORT levels were higher in individuals that survived to fledging. In conclusion, CORT mediates reproductive effort and success during some breeding stages, but it is still unclear why this is the case and whether this same pattern will prevail in other contexts. Author Keywords: corticosterone, HPA axis, maternal effects, reproductive success, tree swallow
Corticosterone Promotes Development of Cannibalistic Morphology and Inhibits Tissue Regeneration in Axolotls (Ambystoma mexicanum)
Salamanders are capable of tissue regeneration throughout all life-stages, which requires the dedifferentiation of mature cells to regrow lost tissues. Dedifferentiation is promoted by degradation of the extracellular matrix by matrix metalloproteases, as well as lysosomal degradation of intracellular and cell-surface proteins that mark cells as part of a mature lineage. Salamanders are also capable of developing cannibalistic phenotypes, plastic traits that are elicited by environmental stressors that result in elevated circulating glucocorticoid (e.g., corticosterone) levels that underlie many fundamental adaptive changes in morphology. Interestingly, the direct effect of corticosterone on regeneration and the cannibalistic phenotype have yet to be examined. In the present thesis, axolotls (Ambystoma mexicanum) were exposed to exogenous corticosterone and 50% of the distal tail tissue was removed. The effects of high corticosterone levels on matrix metalloprotease (MMP-2, MMP-9) and lysosomal acid phosphatase (LAP) activity were assessed; these are two classes of enzymes which are markers of extracellular matrix and intracellular remodeling during regeneration, respectively. We found that elevated corticosterone levels inhibited tissue regeneration, by prolonging the dedifferentiation phase as indicated by increased LAP and reduced MMP-2 and MMP-9 activity. Elevated corticosterone levels also promoted the cannibalistic morphology and this effect was strongest among smaller individuals. Author Keywords: amphibian, cannibalistic morphology, corticosterone, dedifferentiation, regeneration, stress
Island Syndrome and Stress Physiology of Mice in the Genus Peromyscus
Biological differences between island and mainland conspecifics have been well studied, but few studies have addressed differences in stress physiology. Stressors, such as predation and competition for resources, cause the release of glucocorticoids (GCs). Characteristics of island wildlife, called “island syndrome”, are attributed to low levels of predators and competitors. I tested the hypothesis that island syndrome includes differences in GC levels between island and mainland rodents using two approaches; first, using white-footed mice (Peromyscus leucopus) from a near-shore archipelago (Thousand Islands, Ontario) and the nearby mainland; second, using study-skins of deer mice (Peromyscus maniculatus) from two archipelagos offshore of Vancouver Island, British Columbia. White-footed mice in the near-shore archipelago did not show characteristics of island syndrome, or changes in GC levels (feces and hair); however deer mice from both archipelagos in British Columbia were heavier and had lower hair GCs for their size than Vancouver Island mice. Author Keywords: Glucocorticoids, Island rule, Island syndrome, Peromyscus, Stress physiology
Pathogen vs. Predator
Stressors are often an inescapable part of an organism’s life. While the effects of many stressors have been well studied individually, potential interactions between stressors exist that may result in greater than additive negative effects. Stressors may be linked by conflicting demands on energy budgets, interfering with important physiological pathways, or necessitating incompatible adaptive responses. Using Ranavirus (FV3) and larval dragonfly predators (Anax spp.) in a 2x2 factorial experiment on green frog (Lithobates clamitans) tadpoles, I investigate the interactions in behaviour, morphology, and metabolism when both stressors were applied in concert. I demonstrate that activity and feeding are reduced additively by both stressors, and tadpoles increase distance between conspecifics in FV3-exposed tanks, but only in the absence of predators. I also note decreases in mass, and a non-significant marginal increase in metabolic rate of tadpoles exposed to FV3. Interestingly, I provide evidence that FV3 can compromise morphometric responses through antagonistic interactions with perceived predation risk exposure, which may result in significantly elevated mortality even when either stressor is present in sub-lethal quantities. Thus, I conclude that sub-lethal exposure to stressors can nonetheless have substantial impacts on organisms and a more integrative approach to examining the impacts of stressors on individual physiology and fitness is necessary. Author Keywords: Behaviour, Interaction, Morphology, Predation Risk, Ranavirus, Tadpoles
Time-dependent effects of predation risk on stressor reactivity and growth in developing larval anurans (Lithobates pipiens)
The predator vs. prey dynamic is an omnipresent factor in ecological systems that may drive changes in life history patterns in prey animals through behavioural, morphological, and physiological changes. Predation risk can have profound effects on the life history events of an animal, and is influenced by the neuroendocrine stress response. Activation of the hypothalamic-pituitary-adrenal/interrenal axis, and the induction of stress hormones (e.g., corticosterone (CORT)) have been shown to mediate the onset of inducible anti-predator defensive traits including increased tail-depth, and reduced activity. The predator-prey relationship between dragonfly nymphs and tadpoles can be a powerful model system for understanding mechanisms that facilitate changes in the stress response in accordance with altered severity of risk. It has been well demonstrated early in tadpole ontogeny that increased corticosterone (CORT) levels, observed within three weeks of predator exposure, are correlated with increased tail depth morphology. However, the reactivity of the stress response in relation to the growth modulation in developing prey has yet to be fully explored. Accordingly, this thesis assessed the stress and growth response processes in tadpoles that were continuously exposed to perceived predation risk later in ontogeny. Continuous exposure of prey to predation risk for three weeks significantly increased CORT levels, and tail depth. However, tadpoles exposed to six weeks of predation risk acclimated to the presence of the predator, which was observed as a significant reduction of stressor-induced CORT levels. In addition, although increased tail depth has been attributed to predator defense, predator-naïve tadpoles began to display similar tail depth morphology as treated tadpoles at the six week time point. Thus, this thesis suggests that the stress response in lower vertebrate systems (e.g., tadpoles) may operate in a similarly complex manner to that observed in higher vertebrates (e.g., rats), for which severity of risk associated with the stressor aids in defining activity of the stress response. Moreover, the lack of morphological difference between treatments among tadpoles exposed later in ontogeny suggests that the mechanisms for inducing defenses are normal morphological traits in the development of the animal. This thesis paves the way for future research to elucidate the relationship between the neuroendocrine stress response and hormonal pathways involved in growth modulation in the presence of environmental pressures. Author Keywords: Acclimation, Corticosterone, Growth Modulation, Predation Risk, R. pipiens, Tadpole
Social thermoregulation and potential for heterothermy
Northern and southern flying squirrels (Glaucomys sabrinus and G. volans, respectively) are experiencing a climate change induced increase in range overlap, resulting in recent hybridization. We investigated the occurrence of heterospecific communal nesting, a potential facilitator of hybridization, and aimed to confirm the presence of torpor, a potential barrier to hybridization, in flying squirrels. In wild-caught captive squirrels, we conducted a paired nest choice experiment and found that heterospecific nesting did occur, but in a lower frequency than conspecific nesting. Ambient temperature did not affect the frequency of grouped nesting. We attempted to induce torpor in flying squirrels in a laboratory through cold exposure while measuring metabolic rate and body temperature. Strong evidence of torpor was not observed, and metabolic rate remained unchanged with season. We conclude that torpor is not a barrier to hybridization in flying squirrels, but resistance to heterospecific nesting may indicate the existence of one. Author Keywords: heterospecific group, hybridization, northern flying squirrel, social thermoregulation, southern flying squirrel, torpor
Home range use, habitat selection, and stress physiology of eastern whip-poor-wills (Antrostomus vociferus) at the northern edge of their range
The distribution of animals is rarely random and is affected by various environmental factors. We examined space-use patterns, habitat selection and stress responses of whip-poor-wills to mining exploration activity.To the best of my knowledge, fine scale patterns such as the habitat composition within known home ranges or territories of eastern whip-poor-wills have not been investigated. Using a population at the northern edge of the distribution in an area surrounding a mining exploration site, we tested whether variations in habitat and anthropogenic disturbances influence the stress physiology of individuals. We found no effect of increased mining activity on the stress physiology of birds but found a significant scale-dependent effect of habitat on their baseline and stress-induced corticosterone levels, and we suggest that these are the result of variations in habitat quality. The importance of other factors associated with those habitat differences (e.g., insect availability, predator abundance, and microhabitat features) warrants further research. Author Keywords: anthropogenic disturbances, Antrostomus vociferus, corticosterone, eastern whip-poor-will, habitat selection, radio-­telemetry
USE OF SALIVARY CORTISOL TO EVALUATE THE INFLUENCE OF RIDES ON THE STRESS PHYSIOLOGY OF DROMEDARY CAMELS (CAMELUS DROMEDARIUS)
Many facilities attempt to alleviate the risk of chronic stress in captivity by providing environmental enrichment shown to minimize behavioural disorders and stress in several species. One potential form of enrichment used in zoos is training animals to perform rides for guests, however, the effect of this activity on the welfare of individual animals has never been examined. I validated the use of saliva for assessing stress in dromedary camels (Camelus dromedarius) an animal commonly used for rides. I then measured variation in salivary cortisol in four male camels during animal rides for guests at the Toronto Zoo. The camels were sampled during the ride season (from June to August) using four treatments: 1) in their pasture, 2) at the ride area not performing rides, 3) performing a low number of rides (n=50/day) and 4) performing a high number of rides (n=150/day). Furthermore, samples were taken before and after the ride season for comparison. There was a significant difference between the post-ride season treatment and the three treatments involving guest presence during the ride season (ride area, low rides, high rides. This indicates that performing rides is not a stressful experience based on the stress metrics I used, and suggests that rides may be a form of enrichment for dromedary camels. Author Keywords: ACTH challenge, animal welfare, camels, environmental enrichment, salivary cortisol, stress
Neonatal Environment Influences Behavioural and Physiological Reactivity to Stressors, and Mammary Gland Development in BALB/c Mice
Using rodent models, it is possible to study the behavioural and physiological outcomes of early life stress and the influences on normal mammary gland development and carcinogenic risk. Results demonstrate that the experience of three weeks of prolonged maternal separation (LMS; 4 hrs/day) increased the susceptibility of adult, but not pubertal, female BALB/c mice to engage in higher levels of depressive-related immobility behaviour and lower levels of active floating (a suggested adaptive coping behaviour) in the acute forced swim test, than offspring that experienced three weeks of brief separation (BMS; 15 min/day) events. Despite the increased immobility behaviour, adult LMS female offspring demonstrated lower basal corticosterone levels relative to BMS females. However, the experience of chronic early-life stress, regardless of the length, results in greater changes between non-stressed and stressed corticosterone levels (i.e. stressor reactivity) in adult females compared to their male counterparts. These changes were associated with decreased glucocorticoid receptor and coactivator-associated arginine methyltransferase 1 protein expression in mammary gland of female LMS mice at young adulthood, highlighting potential mechanisms underlying their heightened risk of mammary tumourigenesis. These data suggest that early life environments can induce behavioural and physiological alterations observed in adulthood, which may have an influence on the likelihood of malignancies developing in the breast. Author Keywords: coping, early life stress, mammary gland development, mother-infant interactions, steroid receptors, stressor reactivity

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