<p>Seasonal heterothermy—an orchestrated set of extreme physiological responses—is directly responsible for the over-winter survival of many mammalian groups living in seasonal environments. Historically, it was thought that the use of seasonal heterothermy (i.e. daily torpor and hibernation) was restricted to cold-adapted species; it is now known that such thermoregulatory strategies are used by more species than previously appreciated, including many tropical species. The dwarf and mouse lemurs (family Cheirogaleidae) are among the few primates known to use seasonal heterothermy to avoid Madagascar’s harsh and unpredictable environments. These primates provide an ideal study system for investigating a common mechanism of mammalian seasonal heterothermy. The overarching theme of this dissertation is to understand both the intrinsic and extrinsic drivers of heterothermy in three species of the family Cheirogaleidae. By using transcriptome sequencing to characterize gene expression in both captive and natural settings, we identify unique patterns of differential gene expression that are correlated with extreme changes in physiology in two species of dwarf lemurs: C. medius under captive conditions at the Duke Lemur Center and C. crossleyi studied under field conditions in Madagascar. Genes that are differentially expressed appear to be critical for maintaining the health of these animals when they undergo prolonged periods of metabolic depression concurrent with the hibernation phenotype. Further, a comparative analysis of previously studied mammalian heterotherms identifies shared genetic mechanisms underlying the hibernation phenotype across the phylogeny of mammals. Lastly, conducting a diet manipulation study with a captive colony of mouse lemurs (Microcebus murinus) at the Duke Lemur Center, we investigated the degree to which dietary effects influence torpor patterns. We find that tropical primate heterotherms may be exempt from the traditional paradigms governing cold-adapted heterothermy, having evolved different dietary strategies to tolerate circadian changes in body temperature.</p> / Dissertation

Recent studies of heterothermy in free-ranging mammals have revealed that solar radiation is an important variable influencing torpor patterns. The interaction between solar radiation (SR) and arousal costs can be thought of as a continuum from passive increases in body temperature (Tb) (rewarming at a slower rate but energetically less costly), to supplementation of endogenous heat production (rewarming more rapidly but with costs similar to that in the absence of solar radiation). To better understand the importance of solar radiation I experimentally manipulated its availability for E. myurus rewarming from torpor under natural conditions of air temperature and photoperiod. Tb was recorded for E. myurus housed in deep shade (20 % SR), partial shade (40% SR) or full sun (100% SR), and torpor frequency, rewarming rates, minimum body temperature, torpor bout duration and heterothermy index compared among treatments. Animals in unshaded cages rewarmed more rapidly than individuals in partially shaded cages. Torpor bouts were less frequent, but overall levels of heterothermy were higher in E. myurus receiving natural solar radiation compared to those in partially shaded treatments. This study, as far as I am aware, is the first demonstrating that solar heat gain, separated from the effects of an increase in ambient temperature (Ta), plays an important role in torpor arousal. Taken with the direct evidence for elephant shrews basking while rewarming, this demonstrates that animals in the unshaded treatment used solar radiation to supplement, rather than replace, endogenous metabolic thermogenesis. It is clear that E. myurus does not necessarily depend on solar radiation for rewarming, but will take advantage of this resource if available, primarily by reducing time taken to rewarm. / Dissertation (MSc)--University of Pretoria, 2014. / lk2014 / Zoology and Entomology / MSc / Unrestricted

Human activities and unpredictable changes to environmental habitats impose a suite of stress and challenges to animal homeostatic function. Stress responses are often controlled by the release of glucocorticoids to mobilize energy, primarily corticosterone in small mammals, to help regulate homeostatic function such as heterothermy and changes to body condition, the latter of which serves as a proxy for energy reserves. Adaptive heterothermy is influenced by daily and seasonal patterns, heat produced from daily activity, and has been shown to increase in small mammals during times of environmental stress to conserve energy budgets. Body condition also changes in response to environmental perturbations, mobilization of energy by corticosterone, seasonal changes, and activity. My study aimed to disentangle the effects of environment and activity on homeostatic responses by pharmacologically manipulating corticosterone in kangaroo rats. Kangaroo rats are ecosystem engineers, heterothermic, and their activity periods are functions of their thermoregulatory patterns as well as environmental conditions thus making the species a great candidate for this form of study. I conducted two in-situ field experiments to assess for the effects of stress responses on thermoregulation, activity, and body condition. In my first experiment, I used body temperature (Tb) as a proxy for activity time and examined how pharmacologically increased corticosterone influenced kangaroo rat heterothermy responses to the moon phases and ambient temperatures. I also examined their fat, lean mass, and water content at the end of the study. Moon phase was a significant predictor of activity period as animals typically waited longer during the waxing moon phase to become active and cooled down below activity earlier in the night during the waning moon phase. As nights shortened, activity decreased despite environmental conditions becoming warmer. Corticosterone also significantly decreased total activity time and thus steadily increased heterothermy across the length of my experiment. These results indicate activity, not environment, are stronger drivers of heterothermy patterns. Total fat content (energy content) at the end of the study was not affected by corticosterone. The lack of change in fat content was presumably because kangaroo rat body condition was measured once at the end of the study and likely not a reflection of changed body condition over time. In my second experiment, I examined body condition across a longer period by measuring lean mass, fat content, and total body water across a 2-month period in the summer by pharmacologically increasing corticosterone to test the relationship between stress and body condition and to dissociate the two from environmental factors. Body condition indices generally increased across the summer, but corticosterone implantation did not significantly affect body any of the indices. The loss of heterothermic control and decrease in activity time across the summer suggests that animals are likely conserving energy budgets leading to preservation of condition. Further, banner-tailed kangaroo rats generally breed in the spring where body condition falls due to stress induced by increased competition and then increases across the summer as late summer monsoons promote the growth of primary resources utilized by animals thereby preserving body condition. This study suggests that body condition is driven more by life-history traits, activity time, and environment rather than stress responses.

<p>Many endotherms use torpor on a seasonal or daily basis to conserve metabolic resources during difficult conditions, but the capacity to do so has never been recognized as varying intraspecifically. I hypothesized that populations that are exposed to prolonged cold, snow, or scarcity of food resources relative to other conspecifics may express torpor despite the fact that other conspecifics may not use, or even be capable of, torpor. I studied thermoregulation of black-tailed prairie dogs (<I>Cynomys ludovicianus</I>) and Ords kangaroo rats (<I>Dipodomys ordii</I>) at the extreme northernmost periphery of each species range to determine whether there is evidence for geographic variation in torpor patterns.</p><p>Contrary to previous studies of black-tailed prairie dogs near the centre of the species range, I found that northern prairie dogs hibernate during winter, spending up to 95 days per year in torpor. Synchrony of body temperature patterns of some individuals suggests that northern prairie dogs hibernate communally. Similarly, in contrast to previous studies of kangaroo rats in more southern localities, I found that northern Ords kangaroo rats use daily torpor during winter, entering torpor on up to 70 days per year. Kangaroo rats that use deep torpor exhibit comparable survival and pre-winter body mass, but poorer spring body condition, than kangaroo rats that do not use deep torpor. I reported the details of my procedures for studying thermoregulation of small mammals in the wild, to encourage comparable studies that would provide additional insights on intraspecific variation in torpor patterns. My findings are the first to demonstrate that the capacity for torpor varies predictably on a geographic basis.</p>

<p>Many endotherms use torpor on a seasonal or daily basis to conserve metabolic resources during difficult conditions, but the capacity to do so has never been recognized as varying intraspecifically. I hypothesized that populations that are exposed to prolonged cold, snow, or scarcity of food resources relative to other conspecifics may express torpor despite the fact that other conspecifics may not use, or even be capable of, torpor. I studied thermoregulation of black-tailed prairie dogs (<I>Cynomys ludovicianus</I>) and Ords kangaroo rats (<I>Dipodomys ordii</I>) at the extreme northernmost periphery of each species range to determine whether there is evidence for geographic variation in torpor patterns.</p><p>Contrary to previous studies of black-tailed prairie dogs near the centre of the species range, I found that northern prairie dogs hibernate during winter, spending up to 95 days per year in torpor. Synchrony of body temperature patterns of some individuals suggests that northern prairie dogs hibernate communally. Similarly, in contrast to previous studies of kangaroo rats in more southern localities, I found that northern Ords kangaroo rats use daily torpor during winter, entering torpor on up to 70 days per year. Kangaroo rats that use deep torpor exhibit comparable survival and pre-winter body mass, but poorer spring body condition, than kangaroo rats that do not use deep torpor. I reported the details of my procedures for studying thermoregulation of small mammals in the wild, to encourage comparable studies that would provide additional insights on intraspecific variation in torpor patterns. My findings are the first to demonstrate that the capacity for torpor varies predictably on a geographic basis.</p>

Les insectes sont soumis aux fluctuations thermiques de leur environnement mais disposent d’un panel varié de réponses comportementales, physiologiques et biochimiques pour en minimiser les effets délétères et maintenir leur intégrité physiologique. Ainsi certaines espèces régulent activement leur température interne indépendamment de la température de l’environnement. Si ces insectes peuvent s’affranchir des contraintes thermiques imposées par leur environnement, ceux qui se nourrissent du sang chaud d’hôtes vertébrés endothermes n’ont pas d’autres choix que de se confronter à une situation de stress thermique à chaque prise alimentaire. Le principal objectif de ce travail de thèse est de comprendre comment des insectes hématophages, employant des stratégies alimentaires différentes, gèrent le stress thermique associé au flux massif de chaleur engendré par l’ingestion du repas de sang. Nos résultats montrent que ces insectes ont su s’adapter en développant différentes stratégies de thermorégulation. / Insects are submitted to thermal fluctuations of their environment and have developed a wide ranged panel of behavioral, physiological and biochemical responses, to minimize the subsequent deleterious effects and maintain their physiological integrity. Some species actively regulate their internal temperature independently of the temperature of the environment. If these insects can overcome the constraints imposed by their thermal environment, those that feed on warm-blooded vertebrate hosts have no choice but to confront a situation of thermal stress at each feeding event. The main objective of this work is to understand how bloodsucking insects manage heat stress associated with the massive flow of heat generated by the ingestion of the blood meal. Our results show these insects have developed different strategies of thermoregulation to protect themselves from overheating.