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The Interrelationships Between Rodent Respiration in a Burrow

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The Interrelationships Between Rodent Respiration in a Burrow A The Interrelationships between Rodent Respiration in a Burrow Environment and the Physical Ventilation of the Burrow System: a Chapter in the Physiological Ecology of Sundevall's Jird ( Meriones crassus ) Thesis submitted in partial fulfillment of the requirements for the degree of “DOCTOR OF PHILOSOPHY” by Inbal Brickner-Braun Submitted to the Senate of Ben-Gurion University of the Negev 3 July, 2013 Beer Sheva The Interrelationships between Rodent Respiration in a Burrow Environment and the Physical Ventilation of the Burrow System: a Chapter in the Physiological Ecology of Sundevall's Jird ( Meriones crassus ) Thesis submitted in partial fulfillment of the requirements for the degree of "DOCTOR OF PHILOSOPHY" by Inbal Brickner-Braun Submitted to the Senate of Ben-Gurion University of the Negev Approved by the advisors ________________ _______________ Approved by the Dean of the Kreitman School of Advanced Graduate Studies _________________________________ 3 July, 2013 Beer Sheva This research was done under the supervision of Professors Berry Pinshow and Pedro Berliner The Mitrani Department of Desert Ecology and the Wyler Department for Dryland Agriculture. The Jacob Blaustein Institute for Desert Research Research-Student's Affidavit when Submitting the Doctoral Thesis for Judgment I, Inbal Brickner-Braun, whose signature appears below, hereby declare that (Please mark the appropriate statements): X I have written this Thesis by myself, except for the help and guidance offered by my Thesis Advisors. X The scientific materials included in this Thesis are products of my own research, culled from the period during which I was a research student. ___ This Thesis incorporates research materials produced in cooperation with others, excluding the technical help commonly received during experimental work. Therefore, I am attaching another affidavit stating the contributions made by myself and the other participants in this research, which has been approved by them and submitted with their approval. Date: 3 July 2013 Student's name: Inbal Brickner-Braun Signature: _________________________ ACKNOWLEDGEMENTS First and foremost I thank my advisers, Berry Pinshow and Pedro Berliner, for their boundless support and sharing of ideas; they taught me how to ask questions and how to search for answers; the values they instilled in me as a researcher will continue to guide me and be a part of me for the rest of my life. Berry, you have been my mentor, my friend and together with Hanna, my family. Pedro, your enjoyment of scientific discussion has been an inspiration to me. My special thanks to Ishai Hoffman who has been involved in my project from the beginning; thank you for your friendship and for all that you have taught me during long, enjoyable hours in the technical lab and in the field. My special thanks also to Daniel Zucker for being a friend and my research partner; together we explored ideas during long hours, day and night in the lab and in the field, over many cups of coffee. I thank Irina Khokhlova for teaching me everything I know about animal maintenance, breeding and handling; thank you for your encouragement and generosity. Big thanks to my friends and lab members over the past six years: to Carmi Korin, for several enjoyable evenings catching fruit bats in Beer Sheva and for patiently teaching me how to take blood samples from bats; to Marshall McCue for teaching me how to figure out how things work in the lab; to Itzick Vatnick who held my hand when I took the first steps in the world of acid base – thank you for your friendship, to Miri Ben-Hamo and Shai Pilosof who provided me with invaluable statistical advice; to Edward Westen for his suggestions and advice; and to Cynthia Downs for exchange of knowledge and skills. I thank Amos Ar for advising about my blood acid-base balance experiments, Marc Goldberg for technical support and for keeping the technical workshop door open, Rami Mosley for sharing his excellent creative ideas and David Klepatch for providing information from the Desert Meteorology Weather Station. Most importantly, I thank my loving parents for always being there and Avi, my husband, for his love, support and friendship – for everything, really…. Contents Abstract ................................................................................................................................... 3 General Introduction ............................................................................................................... 6 Study animal ........................................................................................................................... 8 PART 1: THE RESPIRATORY ENVIRONMENT INSIDE THE BURROW OF SUNDEVALL'S JIRD.................................................................................................................... 9 1.1 Introduction ............................................................................................................................ 9 Research goals ...................................................................................................................... 11 1.2 Methods ................................................................................................................................ 13 Description of artificial burrow, type 1 ................................................................................. 13 Description of artificial burrow, type 2 ................................................................................. 14 Measurements of [CO 2] inside the burrows .......................................................................... 15 Measurements of wind speed ................................................................................................ 15 Measurements of temperature ............................................................................................... 16 Injection of CO 2 into the nest chamber ................................................................................. 16 1.2.1 Experiment 1. Mechanisms for burrow ventilation ............................................. 16 Experimental settings ............................................................................................................ 17 1.2.2 Experiment 2. Wind-induced ventilation ............................................................ 18 1.2.3 Experiment 3. The effect of the presence of an animal on nest chamber [CO 2] . 19 1.2.4 Experiment 4. [CO 2] in the nest chamber while young were being raised ......... 20 1.3 Results .................................................................................................................................. 23 1.3.1 Experiment 1. Mechanisms for burrow ventilation ............................................. 23 1.3.2 Experiment 2. Wind-induced ventilation ............................................................ 27 1.3.3 Experiment 3. The effect of the presence of an animal on nest chamber [CO 2] . 30 1.3.4 Experiment 4. [CO 2] in the nest chamber while young were being raised ......... 32 1.4 Discussion ............................................................................................................................ 40 Mechanisms of burrow ventilation ....................................................................................... 40 Wind-induced ventilation ...................................................................................................... 41 The effect of the presence of an animal on nest chamber [CO 2] .......................................... 42 [CO 2] in the nest chamber while young were being raised ................................................... 44 1 PART 2: PHYSIOLOGICAL RESPONSES OF SUNDEVALL’S JIRDS TO HIGH AMBIENT CO 2 CONCENTRATION ............................................................................................................ 47 2.1 Introduction .......................................................................................................................... 47 A brief introduction to acid-base balance ............................................................................. 47 The oxygen dissociation curve and the Bohr effect .............................................................. 49 Acid-base balance when ambient [CO 2] is high ................................................................... 50 Blood buffering capacity ....................................................................................................... 50 Ventilatory Regulation of Acid-Base Balance ...................................................................... 51 Renal Compensation of Acid-Base Balance ......................................................................... 52 Research goals ...................................................................................................................... 52 2.2 Methods ................................................................................................................................ 54 Animal maintenance ............................................................................................................. 54 The fat sand rat...................................................................................................................... 54 Preparation of gas mixtures .................................................................................................. 55 2.2.1 Experiment 1. CO 2 tolerance ............................................................................... 55 2.2.2
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