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Ontogeny of Venom Use and Venom Composition in the Western Widow Spider Latrodectus Hesperus David Roger Nelsen

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Ontogeny of Venom Use and Venom Composition in the Western Widow Spider Latrodectus Hesperus David Roger Nelsen Loma Linda University TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works Loma Linda University Electronic Theses, Dissertations & Projects 6-2013 Ontogeny of Venom Use and Venom Composition in the Western Widow Spider Latrodectus Hesperus David Roger Nelsen Follow this and additional works at: http://scholarsrepository.llu.edu/etd Part of the Behavior and Ethology Commons, Biology Commons, Developmental Biology Commons, Public Health Commons, and the Social and Behavioral Sciences Commons Recommended Citation Nelsen, David Roger, "Ontogeny of Venom Use and Venom Composition in the Western Widow Spider Latrodectus Hesperus" (2013). Loma Linda University Electronic Theses, Dissertations & Projects. 292. http://scholarsrepository.llu.edu/etd/292 This Dissertation is brought to you for free and open access by TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works. It has been accepted for inclusion in Loma Linda University Electronic Theses, Dissertations & Projects by an authorized administrator of TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works. For more information, please contact [email protected]. LOMA LINDA UNIVERSITY School of Public Health in conjunction with the Faculty of Graduate Studies ____________________ Ontogeny of Venom Use and Venom Composition in the Western Widow Spider Latrodectus hesperus By David Roger Nelsen ____________________ A Dissertation submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy in Biology ____________________ June 2013 UMI Number: 3566117 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 3566117 Published by ProQuest LLC (2013). 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, MI 48106 - 1346 © 2013 David Roger Nelsen All Rights Reserved Each person whose signature appears below certifies that this dissertation in his/her opinion is adequate, in scope and quality, as a dissertation for the degree Doctor of Philosophy. , Chairperson William K. Hayes, Professor of Biology ________________________________________________________________________ Leonard R. Brand, Professor of Biology and Paleontology ________________________________________________________________________ Penelope Duerksen-Hughes, Associate Dean of Basic Science & Translational Research, School of Medicine ________________________________________________________________________ Kevin Nick, Associate Professor of Geology ________________________________________________________________________ Zia Nisani, Professor of Biology, Antelope Valley College ________________________________________________________________________ Ernest R. Schwab, Associate Dean of Academic Affairs, School of Allied Health Professions iii ACKNOWLEDGEMENTS Earning my Ph.D. has been my dream since I was a small child. Despite my hard work, I would not have been able to accomplish this goal without the support of many people. These past five years have been full of challenges, disappointments, and breakthroughs, but most of all they have been fun. Through this journey my wife, Marie Nelsen, has been by my side as a stalwart supporter. Marie has been instrumental in helping to collect spiders, and has given me strength to complete this dissertation. My parents, Fred and Ana Nelsen, have fostered my love of biology. As a child, they would take me out for hikes and trips to the library so that I could get more books about animals, and they watched many hours of nature documentaries with me. More recently they have encouraged me to pursue my dream while reminding me to stay grounded. My mother and father-in-law, Bobby and Kathy Morris, also deserve recognition for their encouragement and support. Many fellow students deserve recognition as well. Thanks to Allen Cooper for his patience answering many stats questions and helping design studies. Thanks to Gerad Fox for also answering many questions and always having the PDF’s I needed. Thanks to Eric Gren for aiding me with data collection. Thanks to Michael Batech for his support: his vast knowledge of statistics and critical eye during editing. Thanks to the other students, past and present, who have been involved in my academic career and personal life. I have had the pleasure of studying under and being advised by many outstanding professors and researchers. I would like to thank Dr. Leonard Brand for his support, mentoring, and always being willing to listen and advise. Thanks to Dr. Zia Nisani for his iv advice briefly as a fellow student and continuing as a committee member and mentor. His enthusiasm and research have been inspirational. Dr. Kevin Nick has taught me so much, including many technical skills and perspective concerning the importance of my own research. Thanks to Dr. Penelope Duerksen-Hughes and Dr. Ernie Schwab for sharing their experience and expertise, and their willingness to advise me when I became stuck on a particular research problem. Dr. Bill Hayes has been a wonderful mentor. Thank you for taking me on as a student, for seeing the potential in me, and for taking my raw enthusiasm and channeling it into something useful. He has taught me what it means to be a researcher, how to design and execute a study, how to analyze the data, and how to write the results. Dr. Hayes has been a model professor, passing on as much of his knowledge as he can to his students and seeing their success as his own. Thanks must also be given to the other Earth and Biological Sciences professors and staff. There are too many individuals that must be thanked, but know that your support was integral to the completion my dissertation. But special thanks must go to Wayne Kelln; he has been an invaluable resource to me and the Department of Earth and Biological Sciences. There is still so much that I must learn if I am to continue, and succeed, as a researcher and professor. I am glad that my journey does not end here and that I have the privilege of having the family, colleagues, and mentors that I do. I am eternally grateful for what you have already done for me and thank you for your continued support. I must continue to learn, and ask that you all continue to guide me. I sincerely thank you. v DEDICATIONS I would like to dedicate my dissertation to my wife and parents. Marie, you are amazing. Thank you for loving me. & To my parents, Fred and Ana Nelsen, without your guidance I would not have made it this far. Thank you. vi CONTENTS Approval Page .................................................................................................................... iii Acknowledgements ............................................................................................................ iv Dedication .......................................................................................................................... vi Table of Contents ............................................................................................................. viii List of Tables ..................................................................................................................... xi List of Figures .................................................................................................................. xiii Abbreviations .....................................................................................................................xv Abstract ............................................................................................................................ xvi Chapter 1. Introduction ..............................................................................................................1 Specific Objectives ............................................................................................3 References ..........................................................................................................6 2. Poisons, Toxungens, and Venoms: Redefining and Classifying Toxic Biological Secretions and the Organisms that Employ Them .................................8 Abstract ..............................................................................................................9 Introduction ......................................................................................................10 Toxins ..............................................................................................................11 Existing definitions of venom ..........................................................................13 Hierarchy and Exclusiveness .....................................................................16 Source of Secretion ....................................................................................17 Mode of Transmission, Including a Delivery Structure or Delivery System ........................................................................................................18 Biological Role(s) ......................................................................................20 Active Application .....................................................................................22
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