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71-20,037 BAUST, John George, 1942- INFLUENCES of LOW TEMPERATURE on COLD HARDINESS and NEURAL ADAPTATION in the ALASKAN BEETLE

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71-20,037 BAUST, John George, 1942- INFLUENCES of LOW TEMPERATURE on COLD HARDINESS and NEURAL ADAPTATION in the ALASKAN BEETLE Influences Of Low Temperature On Cold Hardiness And Neural Adaptation In The Alaskan Beetle, Pterostichus Brevicornis (Carabidae) Item Type Thesis Authors Baust, John George Download date 28/09/2021 03:15:03 Link to Item http://hdl.handle.net/11122/9190 71-20,037 BAUST, John George, 1942- INFLUENCES OF LOW TEMPERATURE ON COLD HARDINESS AND NEURAL ADAPTATION IN THE ALASKAN BEETLE PTEROSTICHUS BREVICORNIS (CARABIDAE). University of Alaska, Ph.D., 1970 Physiology University Microfilms, A XEROX Company, Ann Arbor, Michigan THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. INFLUENCES OF LOW TEMPERATURE ON COLD HARDINESS AND NEURAL ADAPTATION IN THE ALASKAN BEETLE PTEROSTICHUS BREVICORNIS (CARABIDAE) A DISSERTATION Presented to the Faculty of the University of Alaska in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY by John G'. Baust, B.A. College, Alaska May 1970 Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. INFLUENCES OF LOW TEMPERATURE ON COLD HARDINESS AND NEURAL ADAPTATION IN THE ALASKAN BEETLE PTEROSTICHUS BREVICORNIS (CARABIDAE) APPROVED: Chairman i J2 _ APPROVED: Date L % ? 0 Vice President for Research and Advanced Study Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ABSTRACT A species of adult carabid beetle, Pterostichus brevicornis, has been found to overwinter in the Fairbanks locale within de­ cayed stumps. Empirical observations indicated that body freezing was tolerated. Indications of continued vital processes through­ out the overwintering period were also noted. In ligh t of these observations the existance of a cryoprotective compound was sus­ pected and determined to be glycerol. The hemolymph glycerol of adult Pterostichus brevicornis was determined u tilizin g quantitative paper chromatography. Seasonal variations were revealed in the concentration of this substance. Mean glycerol levels in excess of 22 gm% have been measured during winter with values decreasing to less than 1 gm% during spring and summer. Acclimation studies gave similar results. In the artificially warmed winter beetles glycerol content decreased to less than 1 qm% within 36 hours at 20°C. The stimulus to the iniation of glycerol synthesis was found to be f ir s t exposure to 0°C following summer. The rate of accumulation varied nearly directly with sub-zero (°C) ex­ posure temperature. Glycerol concentrations were closely correlated with the changing whole body supercooling and hemolymph freezing points in both naturally acclimatized and laboratory acclimated specimens. Both supercooling and freezing points were depressed 0.9°C per 4 gm% increased in gly­ cerol. Hemolymph glucose and trehalose levels were also correlated i i i Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. with fluctuations in glycerol in an attempt to define the probable source of glycerol. Glucose did not vary seasonally whereas tre­ halose changed sign ificantly. Temperature fluctuations within the hibernacula were considered. Also, seasonal variations in locomotor response to temperature were determined in a thermal gradient chamber and correlated with changes in cold tolerance. Mean temperature preferences were found to vary from a summer high of +13.3°C to a winter low of -5.5°C. Sub-freezing exposures were avoided unless glycerol was present. Attempts were made to integrate the above observations with neural function. Electrical recordings from the region of the modified tro­ chanter on the hind thoracic legs revealed a thermosensitive region. Semi-microelectrode recordings have evidenced motor fiber populations displaying narrow bands of differential temperature se n sitiv ity . A ctivity in some fibers has been recorded at temperatures as low as -11.7°C. These fibers vary their tonic discharge patterns with varying states of acclimation and acclimatization, thereby allowing continued activity at sub-zero temperatures. Localization of discharge to motor efferents was made utilizing neurophysiological, surgical and neurohumoral techniques. Fluctuations in the discharge frequencies are thought to reflect changes in nerve tone. iv Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGEMENTS It is with sincere gratitude that I thank my advisor, Dr. L. Keith Miller, for his continued interest and encouragement through­ out this project. Without his guidance, this study would not have been possible. I greatly appreciate the discussions and guidance offered by Dr. L. Irving. His never ending interest did much to smooth the course of th is study. I am also indebted to Dr. T. Kaufmann fo r her kind words and instruction. Special thanks are extended to both Karen Coady and Harry (Don) Draper. Karen, for her perseverance through experiments running beyond the stroke of m idnight, and to Don, fo r his continued and s k il l f u l collecting of the majority of specimens utilized. I thank Don Borchert for his excellent reproduction of photo­ graphic records and figures presented in this dissertation. Deepest appreciation is extended to Helga Wakefield, Kathy Huddleston and Betty Mather for their extended patience in the physical preparation of this volume. My appreciation is also extended to my fellow students, William Arvey, Wayne Heimer and Louis Nauman, for their active interests and criticisms of this work. I also extend sincerest thanks to Dr. S. Swihart of the State University of New York and Dr. Chandler Me. Brooks of the Downstate College of Medicine for their patience in helping provide me with the background required for the fruitful persuit of research en­ deavors . Finally, I give special thanks to both my wife, Judy, and my parents for their continued confidence and support leading to this conclusion. Support by the NASA under grant NGT 02-001-007 in the form of a pre-doctoral fellowship and the NIH under grant GM 10402 is acknowledged. v Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS PAGE LIST OF TABLES viii LIST OF ILLUSTRATIONS ix INTRODUCTION 1 History of Insect Freezing Survival 1 Objects of Study 16 CHAPTER I GENERAL ECOLOGICAL, ETHOLOGICAL AND PHYSIOLOGICAL 18 ASPECTS OF OVERWINTERING Life History and the Microhabitat 19 Thermal Fluctuations Within the Microhabitat 22 Locomotor Responses to Temperature 29 CHAPTER I I VARIATIONS IN COLD HARDINESS ASRELATED TO GLYCEROL 37 Introduction 38 Methods and Material 39 Glycerol Analysis 39 Supercooling Point Determinations 44 Freezing Point Determinations 44 Carbohydrate Analyses 44 Results 48 Thermal Acclimatization Experiments 48 Thermal Acclimation Experiments 58 Discussion 76 vi Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. (TABLE OF CONTENTS, CONT.) PAGE CHAPTER I I I THE PHYSICO-CHEMICAL ASPECTS OF GLYCEROL PROTECTION 84 MECHANISMS OF CRYOPROTECTION Introduction 85 Methods and Results 88 Discussion 93 CHAPTER IV TEMPERATURE INDUCED NEURAL ADAPTATIONS 99 Introduction 100 Methods and M aterials 101 Results 102 D iscussion 112 GENERAL SUMMARY 114 BIBLIOGRAPHY 116 v ii Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF TABLES PAGE Table 1-1. Cooling rates of decayed stumps per centimeter depth 25 (horizontal) upon acute cooling to -30°C. Time 0 temperatures ranged between 18 and 20°C. Table 2-1. Rg values from chromatograms of various cryoprotective 42 compounds (sugars and polyhydric alcohols). Table 2-2. Seasonal changes in glycerol content, supercooling points 51 and freezing points of the hemolymph of naturally acclim­ atized Pterostichus brevicornis. Table 2-3. Changes in glycerol content, freezing points and super­ 61 cooling points during spring (warm) acclimation. Table 2-4. Rate changes per hour of glycerol content, supercooling 62 points and freezing points during spring acclimation temperatures. Table 2-5. Schedule of fall (low temperature) acclimation experi­ 66 ments. Table 2-6. Changes in hemolymph glycerol, trehalose and freezing and 68 whole body supercooling points during fall (low temp­ erature) acclimation in P_. brevicornis. Table 4-1. Lists the variations between winter and summer ranges of 107 activity of the trochanter motor fibers in P. brevicornis. Table 4-2. Extinction temperatures and thermal ranges of a ctivity of 108 motor fibers in acclimated specimens of P.. brevicornis. v iii Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF ILLUSTRATIONS PAGE Figure 1-1. Temperature changes at various depths (cm.) in a de- 24 cayed stump hibernaculum rapidly cooled. The time 0 line represents initial air temperatures. Figure 1-2. Cooling rates of stump at various depths (cm.) upon 27 exposure to -42°+ 2°C. Each line (#1-9) represents horizontal depth of thermocouples. Figure 1-3. Warming rates of a stump at various depths (cm.) upon 28 exposure to air temperature of 18°C. Each line (#1-9) represents horizontal depth of thermocouples. Figure 1-4. Variations in stump temperature effected by oscillatory 30 warming and cooling. Each line (#1-9) represents hor­ izontal depth (cm.) of thermocouple. Figure 1-5. Diagramatic illustration of the thermal gradient chamber. 32 Figure 1-6. A plot
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