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The Development and Improvement of Instructions THE ROLE OF ANDROGENS IN MALE PREGNANCY AND FEMALE COMPETITIVE BEHAVIOR IN A SEX ROLE REVERSED PIPEFISH A Dissertation by SUNNY KAY SCOBELL Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY December 2011 Major Subject: Biology The Role of Androgens in Male Pregnancy and Female Competitive Behavior in a Sex Role Reversed Pipefish Copyright 2011 Sunny Kay Scobell THE ROLE OF ANDROGENS IN MALE PREGNANCY AND FEMALE COMPETITIVE BEHAVIOR IN A SEX ROLE REVERSED PIPEFISH A Dissertation by SUNNY KAY SCOBELL Submitted to the Office of Graduate Studies of Texas A&M University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Approved by: Co-Chairs of Committee, Adam G. Jones Gil G. Rosenthal Committee Members, Duncan S. MacKenzie John T. Jaques Head of Department, U. J. McMahan December 2011 Major Subject: Biology iii ABSTRACT The Role of Androgens in Male Pregnancy and Female Competitive Behavior in a Sex Role Reversed Pipefish. (December 2011) Sunny Kay Scobell, B.A., Augustana College; M.S., University of Oklahoma Co-Chairs of Advisory Committee: Dr. Adam G. Jones Dr. Gil G. Rosenthal The sex-role reversal and male pregnancy found in syngnathids are highly unusual traits in vertebrates. Reproductive hormones likely influence development and regulation of these traits. However, very few studies have examined the underlying hormonal mechanisms that mediate female competitive behavior and male pregnancy. New methodologies and better husbandry practices have made such studies more feasible in recent years. Research on a relatively small number of species has suggested that androgens are likely regulators of spermatogenesis and the development of the male brood pouch prior to pregnancy. Androgens are also potential candidates for mediating sex-role reversed behavior in female syngnathids. The goal of this dissertation was to examine the role of androgens in the male reproductive cycle and female intrasexual competitive behavior in the sex-role reversed Gulf pipefish, Syngnathus scovelli. From review of the literature, I developed a model for the hormonal regulation of the male reproductive cycle in seahorses. I predicted that androgens would be low during the early stages of pregnancy and increase during the end of pregnancy as males go iv through another cycle of spermatogenesis in preparation for the next mating event. My study of 11-ketotestosterone and testis mass across the reproductive cycle in male S. scovelli supported this model. I also conducted several studies on the role of androgens in female competitive behavior. I determined that treatment with 11-ketotestosterone the evening prior to an intrasexual interaction resulted in an increase in competitive behavior in large over small test females. Conversely, treatment with 11-ketotestosterone one hour prior to an intrasexual interaction resulted in a decrease in competitive behavior in large over small females when stimulus female behavior was controlled. A comparative study of competitive and courtship behavior in S. scovelli and the closely related S. floridae suggested that sexual selection has affected competitive and courtship behavior in both males and females of these species. The diversity of reproductive patterns exhibited by syngnathids suggests that they will provide a unique opportunity to assess how hormonal regulation of reproductive behavior and function has evolved within this lineage. v DEDICATION To my mother, father, and sister You are my constant source of strength, support, and inspiration vi ACKNOWLEDGEMENTS I am forever indebted to my co-advisors, Adam Jones and Gil Rosenthal, for bringing me to Texas A&M, for guiding me through this process, for working with me through the rough patches, and for opening up a whole new world of science to me. You have expanded my outlook on biology and challenged me to think bigger than I ever had before. I cannot thank you enough for all that you have done for me. I also want to thank my committee members, Duncan MacKenzie and Scott Jaques; this project would not have been possible without your help. Duncan, thank you so much for translating the Boisseau thesis and co-authoring the review, for help with the assays, for our many conversations on physiology, and for encouraging me to pursue comparative endocrinology. Scott, thank you for being so generous with your time, your lab space, assay supplies, and your help. Working in your lab made these difficult assays seem easy. I want to thank all of the past and present members of the Jones Lab: Kenyon Mobley, Charlyn Partridge, Clay Small, Kim Paczolt, Nick Ratterman, Emily Rose, and Sarah Flanagan; the Rosenthal Lab: Michi Tobler, Macteld Verzijden, Holly Kindsvater, Brad Johnson, Zach Cress, Zach Culumber, Rongfeng Cui, Victoria Smith, Mattie Squire, Pablo Delclos, and Giovanna Serena; the MacKenzie Lab: Thomas Miller, Richard Jones, and Allison Wilkes; and the Jaques Lab: Malcolm Delovio. Thank you for all of the great conversations – scientific and otherwise, the logistical support in the vii field, the lab, and the fish room, the constructive criticism, the laughs, the lunches, and most of all, your friendship. Thanks to all of my undergraduate research assistants at Texas A&M University: Megan Thompson, Tessia Lamison, Sarah Gaughan, Vanessa Seyle, Jeff Westwood, Jamie Morris, Andy Marshall, and Anand Srikrishnan. You have made working with a small, high-maintenance fish not only possible but also enjoyable. Thank you to the University of Texas Marine Science Institute, Rick Kline, and Dr. Izhar Khan for allowing me to use your centrifuge for my field-collected blood samples. I also want to thank Dr. Ingo Schlupp from the University of Oklahoma for introducing me to Gil Rosenthal and helping me make the transition to Texas A&M University. I want to give a special thanks to Dr. James Deegear. Your ability to turn a complex situation into an eloquent, tangible statement that made sense to me helped me work through many difficult problems. I would not have made it through this process without you. To my parents, Henry and Beverley Scobell, thank you for all of your love and support. Thank you for teaching me strength, determination, and resilience. Words cannot express how much I appreciate what you have done for me. To my sister, Dr. Summer Scobell DeRuyter, thank you for going through this with me. You always had the right words at the right time, and you made me believe where there is a will, there is a way. viii To my love, Edward Gemar, thank you so much for your logic, laughter, and love. It means so much to me that through it all, you have been by my side, encouraging me to pursue my dream. ix NOMENCLATURE 17,20-P 17,20-dihydroxy-4-pregnen-3-one 17β-HSD 17β-Hydroxysteroid dehydrogenase 11β-HSD 11β-Hydroxysteroid dehydrogenase ACTH Adrenocorticotropic hormone AVP Arginine vasopressin AVT Arginine vasotocin E2 Estradiol EE2 Ethinylestradiol FSH Follicle stimulating hormone GnRH Gonadotropin releasing hormone IT Isotocin KA Ketoandrostenedione KT 11-ketotestosterone LH Luteinizing hormone MIS Maturation inducing steroid OHA 11β-hydroxyandrostenedione OT Oxytocin PRL Prolactin T Testosterone TP Testosterone proprionate x TABLE OF CONTENTS Page ABSTRACT .............................................................................................................. iii DEDICATION .......................................................................................................... v ACKNOWLEDGEMENTS ...................................................................................... vi NOMENCLATURE .................................................................................................. ix TABLE OF CONTENTS .......................................................................................... x LIST OF FIGURES ................................................................................................... xii LIST OF TABLES .................................................................................................... xiv CHAPTER I INTRODUCTION AND LITERATURE REVIEW ........................... 1 Introduction .................................................................................... 1 Literature review ............................................................................ 8 Hormones in Syngnathids .............................................................. 10 Future directions ............................................................................. 31 Dissertation justification ................................................................ 34 II THE ROLE OF 11-KETOTESTOSTERONE IN THE MALE REPRODUCTIVE CYCLE OF Syngnathus scovelli .......................... 35 Methods .......................................................................................... 41 Results ............................................................................................ 44 Discussion ...................................................................................... 46 III HORMONE MANIPULATION OF FEMALE INTRASEXUAL COMPETITIVE AGGRESSION IN THE SEX-ROLE REVERSED GULF PIPEFISH, Syngnathus scovelli ................................................ 54 Methods .......................................................................................... 62 Results ............................................................................................ 68 Discussion .....................................................................................
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