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THE ALASKA MARMOT and ARCTIC GROUND SQUIRREL by Trixie

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THE ALASKA MARMOT and ARCTIC GROUND SQUIRREL by Trixie Expression And Mechanisms Of Hibernation In The Artic: The Alaska Marmot And Arctic Ground Squirrel Item Type Thesis Authors Lee, Trixie Nicole Download date 02/10/2021 15:52:21 Link to Item http://hdl.handle.net/11122/9128 EXPRESSION AND MECHANISMS OF HIBERNATION IN THE ARCTIC: THE ALASKA MARMOT AND ARCTIC GROUND SQUIRREL By Trixie Nicole Lee RECOMMENDED: f a Dri Barbara Taylor Dr. Brian Barnes Dr. C. Loren Buck, Co-Chair of Advisory Committee Dr. Diane O’Brien, Co-Chair of Advisory Committee Dr. Clg^fcT'ftiulder, Chair, Biology and Wildlife Dept. APPROVED: Dr. Paul Layer, Dean, al Science and Mathematics Dr. cSwrence Duffy, Dean, Graduate School f. Date / EXPRESSION AND MECHANISMS OF HIBERNATION IN THE ARCTIC: THE ALASKA MARMOT AND ARCTIC GROUND SQUIRREL A DISSERTATION Presented to the Faculty of the University of Alaska Fairbanks in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY By Trixie Nicole Lee, B.S. Fairbanks, Alaska May 2012 UMI Number: 3528858 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. UMI 3528858 Published by ProQuest LLC 2012. Copyright in the Dissertation held by the Author. Microform Edition © ProQuest LLC. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 A bstr a c t The Arctic is home to animals that have taken adaptations to overwintering to extremes. In this dissertation, I have investigated one of these adaptations, hibernation, in two species from the Arctic, the Alaska marmot(Marmota broweri) and the arctic ground squirrel (Urocitellus parryii). The expression of hibernation under natural conditions in these species was compared by collecting body temperature records of free- living individuals. The Alaska marmot, a highly social species, demonstrated extreme synchrony in body temperature patterns among a family group, indicating a strong reliance on social thermoregulation. In contrast, the arctic ground squirrel was confirmed to be a solitary hibemator that reduces body temperature below freezing during torpor. Both species must produce heat when soil temperatures are significantly below freezing for most of the winter. At these subfreezing ambient temperatures, the arctic ground squirrel has shown an increasing reliance on nonlipid fuel during torpor, driving a loss of lean mass during hibernation of ~20%. I calibrated deuterium dilution to repeatedly estimate body composition in this species, which dramatically changes adiposity through its annual cycle, and used this technique to quantify lean mass loss throughout hibernation in a study of tissue metabolism. I also developed and applied the natural abundance of nitrogen and carbon stable isotopes as tools for monitoring differential tissue metabolism and differentiating mixed metabolic fuel use in the arctic ground squirrel. These data clarified the mechanism of change in nitrogen stable isotopes and indicated that hibernating ground squirrels rebuild organ tissues while breaking down muscle tissue to meet energetic demands. Furthermore, I corroborated a shift in metabolic fuel use toward nonlipid sources during torpor at low ambient temperatures by using the carbon isotope ratio in exhaled breath in combination with respiratory quotient. This dissertation combines studies of hibernation patterns in free-living animals with experimental data on the tissues and fuels being catabolized at very low temperatures to broaden our understanding of how small mammals successfully hibernate in severe winter conditions. It also presents the development and use of stable isotope ratios as physiological tools in hibernating species. T a bl e o f C o n t en t s Page Signature Page ........................................................................................................................i Title Page...............................................................................................................................ii Abstract .................................................................................................................................iii Table of Contents..................................................................................................................iv List of Figures.................................................................................................................... viii List of Tables.........................................................................................................................x Acknowledgments................................................................................................................ xi General Introduction..............................................................................................................1 Literature Cited .................................................................................................................4 Chapter 1 Body temperature patterns during hibernation in a free-living Alaska marmot (Marmota broweri) ................................................................................................................7 Abstract .............................................................................................................................7 Introduction.......................................................................................................................8 Materials and Methods......................................................................................................9 Results............................................................................................................................. 12 Patterns of Body Temperature....................................................................................12 Cooling and Rewarming Rates...................................................................................12 Soil Temperatures......................................................................................................13 Body Mass..................................................................................................................13 Discussion.......................................................................................................................13 Acknowledgements.........................................................................................................17 References....................................................................................................................... 17 Chapter 2 Sociality influences hibernation patterns of two arctic species, the Alaska marmot and arctic ground squirrel ...................................................................................... 25 Abstract ...........................................................................................................................25 Introduction.....................................................................................................................26 Materials and Methods ................................................................................................... 27 V Page Animals.......................................................................................................................27 Surgical Procedures ....................................................................................................28 Metrics........................................................................................................................ 28 Soil Temperature........................................................................................................ 29 Statistical Analysis..................................................................................................... 29 Results............................................................................................................................. 30 Synchrony...................................................................................................................30 Cooling Rates............................................................................................................. 31 Rewarming Rates....................................................................................................... 31 Characteristics of Heterothermy................................................................................ 31 Discussion....................................................................................................................... 32 Acknowledgments ...........................................................................................................36 Literature Cited ...............................................................................................................37 Chapter 3 Estimating lean mass over a wide range of body composition: a calibration of deuterium dilution in the arctic ground squirrel .................................................................48 Abstract ...........................................................................................................................48 Introduction.....................................................................................................................49
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