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Proquest Dissertations Overwintering Physiology Of Arctic And Subarctic Insects From Interior Alaska Item Type Thesis Authors Sformo, Todd L. Download date 24/09/2021 17:06:40 Link to Item http://hdl.handle.net/11122/9041 OVERWINTERING PHYSIOLOGY OF ARCTIC AND SUBARCTIC INSECTS FROM INTERIOR ALASKA By Todd Sformo RECOMMENDED: mminwv Dr.JfracfreajcyeMy/Martainrstoinnsc -? I "A Ab - y i- — Dr. Bert Boyer /V / f / / K- Dr. John Duman, Co-Chaifof Advisory Committee Dr. Brian Barnes, Co-Chair of Advisory Committee A V ■ Dr. Rich Boone, Chair, Biology and Wildlife Dept. / APPROVED: Dr. Paul Layer, Dean, Co]JSrge-crf Natural Science and Mathematics ' Dr. Lawrence K. Duffy, Dean o f thetGraduate School Date OVERWINTERING PHYSIOLOGY OF ARCTIC AND SUBARCTIC INSECTS FROM INTERIOR ALASKA 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 Todd L. Sformo, B.A., M .A, M.F.A., M.S. Fairbanks, Alaska December 2009 UMI Number: 3401549 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. UMT Dissertation Publishing UMI 3401549 Copyright 2010 by ProQuest LLC. All rights reserved. This edition of the work is protected against unauthorized copying under Title 17, United States Code. uestA ® ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 Abstract This dissertation focuses on the overwintering of three insects from Interior Alaska: a hemipteran, Elasmostethus interstinctus, and a coleopteran, Cucujus clavipes puniceus, that are freeze avoiding in the strict sense of the phrase, and a dipteran, Exechia nugatoria, that is simultaneously partially freeze avoiding and freeze tolerant. The variability within the freeze avoidance strategy itself is a key theme throughout this dissertation. Two significant contributions to comparative physiology are the confirmation of insect vitrification (glass formation) with its attendant extension of freeze avoidance and survival into a new, extreme low temperature record of -100°C and the simultaneous coupling of freeze avoidance and tolerance within an individual, which may more properly be described as a new overwintering strategy. Vitrification is the process by which ice crystallization is circumvented, resulting in a supercooled amorphous solid. Through a combination of antifreeze proteins that inhibit ice nucleation, dehydration tolerance, presence of high glycerol concentration, and low temperatures, the mobility of the remaining liquid water molecules is reduced, effectively by-passing the crystalline state. The second contribution is the discovery of a new overwintering strategy that combines freeze avoidance and freeze tolerance within an individual. In this case, the abdomen freezes (and the insect survives), while the contiguous head/thorax remains supercooled. These findings lead to the following evolutionary and trans-disciplinary questions. Is vitrification an adaptation? What is the selective advantage of compartmentalizing ice between body sections of an individual insect? Is this new overwintering strategy an example of a species transitioning between either becoming exclusively freeze avoiding or free tolerant? Applying new understanding of mechanisms of insect vitrification and avoidance of devitrification to cryomedicine may extend preservation of human tissues and organs. Similarly, for physical and material scientists, by understanding the patterns of ice formation within insects that tolerate, inhibit, and/or impede ice formation below the homogeneous ice nucleation temperature of water (-40°C), new biomimetic possibilities can be envisioned. V Table of Contents Signature Page............................................................................................................................................. i Title Page......................................................................................................................................................ii Abstract........................................................................................................................................................ iii Table of Contents .......................................................................................................................................v List of Figures............................................................................................................................................xi List of Tables...........................................................................................................................................xiv Dedicated to ............................................................................................................................................. xvi Introduction: Brief History of Research on Insect Overwintering Physiology in Alaska... 1 General Insect Research and Climate Characteristics in Alaska .......................................... 1 Per (Pete) Scholander ...........................................................................................................................4 Keith Miller............................................................................................................................................. 6 Mark Oswood ..........................................................................................................................................8 Brian Barnes and Jack Duman .......................................................................................................... 9 Summary of significant results by insect ......................................................................................11 Chapter 1 Elasmostethus interstinctus: Resistance to Inoculative Freezing .....................11 Chapter 2 Cucujus clavipes puniceus: Deep Supercooling and Vitrification .................. 11 Chapter 3 Cucujus clavipes puniceus: Probability of Freezing............................................13 Chapter 4 Exechia nugatoria: Simultaneous Freeze Tolerance and Avoidance ........... 13 References ....................................................................................................................................14 Chapter 1 Overwintering and inoculative freeze resistance in the freeze-avoiding stink bug Elasmostethus interstinctus (Heteroptera: Acanthosomatidae) in Alaska ......................18 VI Abstract.........................................................................................................................................18 Introduction ................................................................................................................................. 19 Materials and Methods .............................................................................................................21 Anim als....................................................................................................................................21 Microhabitat Temperature..................................................................................................22 Supercooling and Survival ................................................................................................ 23 Water Content ........................................................................................................................24 Thermal Hysteresis Activity ..............................................................................................24 Polyol Determinations .........................................................................................................25 Statistical Analysis ............................................................................................................... 26 Results...........................................................................................................................................26 Animals, Microhabitat, and Survival ..............................................................................26 Supercooling and Water Content .....................................................................................28 Polyol Determination .......................................................................................................... 30 Correlation between water content and SCP ................................................................30 Thermal Hysteresis and Melting Points .........................................................................31 Discussion....................................................................................................................................31 Acknowledgements ...................................................................................................................38 References....................................................................................................................................39 Figure............................................................................................................................................44 Tables............................................................................................................................................45 vii Chapter 2 Deep Supercooling, Vitrification, and Limited Survival to -100°C
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