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The Ultimate and Proximate Regulation of Developmental Polyphenism in Spadefoot Toads Brian L

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The Ultimate and Proximate Regulation of Developmental Polyphenism in Spadefoot Toads Brian L Florida State University Libraries Electronic Theses, Treatises and Dissertations The Graduate School 2009 The Ultimate and Proximate Regulation of Developmental Polyphenism in Spadefoot Toads Brian L. (Brian Lee) Storz Follow this and additional works at the FSU Digital Library. For more information, please contact [email protected] FLORIDA STATE UNIVERSITY COLLEGE OF ARTS AND SCIENCES THE ULTIMATE AND PROXIMATE REGULATION OF DEVELOPMENTAL POLYPHENISM IN SPADEFOOT TOADS By BRIAN L. STORZ A Dissertation submitted to the Department of Biological Science in partial fulfillment of the requirements for the degree of Doctor of Philosophy Degree Awarded: Summer Semester, 2009 Copyright © 2009 Brian L. Storz All Rights Reserved The members of the committee approve the dissertation of Brian L. Storz defended on June 30, 2008. ________________________________ Timothy S. Moerland Professor Directing Dissertation ________________________________ J. Michael Overton Outside Committee Member ________________________________ P. Bryant Chase Committee Member ________________________________ Thomas A. Houpt Committee Member ________________________________ Wu-Min Deng Committee Member Approved: ______________________________ P. Bryant Chase, Chair, Department of Biological Science ______________________________ Joseph Travis, Dean, College of Arts and Sciences The Graduate School has verified and approved the above-named committee members. ii ACKNOWLEDGEMENTS To Shonna for always being there for me and being the most supportive and wonderful wife in the world. To Logan for being crazy and keeping my life entertaining and utterly exhausting. To Timothy S. Moerland for helping me up, dusting me off, and putting me back on the biology bicycle, for teaching me how to think like a CMB biologist, and most importantly, for teaching me how to balance science with family. To Joe Travis for turning me into a scientist and having faith in me to succeed no matter the circumstances. To my committee members Tom A. Houpt, P. Bryant Chase, Wu-Min Deng, and J. Michael Overton for being positive and optimistic influences in my life and providing plenty of constructive criticism. To my past committee members, especially Scott J. Steppan, Alice A. Winn, and Gavin J. P. Naylor. To my lab mates, past and present, for good discussion, good drinking times, and helpful feedback about research and life. To Anne Thistle for reviewing many, many versions of my manuscripts. To Dan R Buchholz and Jeff Arendt for inspiring conversations about science and life. To the helpful people in all the FSU research facilities, including Xixi Jia in histology, Kim and Tom in imaging, Brandon and Rob in the stockroom, and the helpful people in core facilities. To all the undergraduates whom have worked with me in some capacity over the years, and especially Korey Wilson, Jessica Heinrichs, Arash Yazdani, Brett Mulvey, and Ryan Phillips. To all the staff, scientists, especially Ryan Martin, and volunteers, past and present, at the Southwestern Research Station in Portal, Arizona. To my funding sources, American Heart Association, American Museum of Natural History, Sigma Xi, Robert B. Short Fellowship from Florida State University, Timothy S. Moerland, Joe Travis, and at one point Mastercard. To the countless spadefoot tadpoles and adults that gave their lives to make this research possible. iii TABLE OF CONTENTS List of Tables ......................................................................................... vi List of Figures ......................................................................................... ix Abstract ......................................................................................... xii 1. INTRODUCTION .............................................................................. 1 2. ULTIMATE REGULATION OF SPADEFOOT DEVELOPMENTAL POLYPHENISM, ENVIRONMENTAL AND BEHAVIORAL FACTORS Introduction ........................................................................................ 4 Methods ........................................................................................ 10 Results ........................................................................................ 16 Discussion ....................................................................................... 20 3. TEMPORALLY DISSOCIATED, TRAIT-SPECIFIC MODIFICATIONS UNDERLIE PHENOTYPIC POLYPHENISM IS SPEA MULTIPLICATA TADPOLES, WHICH SUGGESTS MODULARITY Introduction ........................................................................................ 31 Methods ....................................................................................... 34 Results ........................................................................................ 37 iv Discussion ....................................................................................... 41 4. PROXIMATE REGULATION OF CARNIVORE MUSCLE ENLARGEMENT; HISTOLOGICAL COMPARISONS OF SPADEFOOT CARNIVORE AND OMNIVORE PHENOTYPES Introduction ....................................................................................... 46 Methods ...................................................................................... 49 Results ...................................................................................... 56 Discussion ...................................................................................... 73 CONCLUSION ..................................................................................... 79 REFERENCES ...................................................................................... 81 BIOGRAPHICAL SKETCH ................................................................ 89 v LIST OF TABLES 1.1 Compilation of studies investigating larval polyphenism in spadefoot toads........................................................................................................ 8 1.2 Environmental differences among ponds from which carnivores and omnivores were collected from behavior analyses................................. 15 1.3 Number of Spea multiplicata carnivore, omnivore, and unknown phenotypes produced in each group treatment........................................ 18 1.4 Number of Spea bombifrons carnivore, omnivore, and unknown phenotypes produced in each group treatment........................................ 20 1.5 Number of Spea multiplicata and Spea bombifrons carnivore, omnivore, and unknown phenotypes produced in each individual treatment................................................................................................. 21 1.6 a T-test of number of mouth movement cycles between carnivores and omnivores from different ponds. b T-test of swim time during a 2-minute interval between carnivores and omnivores from different ponds.................................................... 23 1.7 T-test of number of mouth movement cycles during a 2-minute interval between carnivores and omnivores from Rock Tank pond at different ages........................................................................................... 25 1.8 a T-test of swim time during a 2-minute interval between carnivores and omnivores from Rock Tank at different ages. b T-test of swim time in the presence of fairy shrimp during a 2- minute interval between carnivores and omnivores from Rock Tank at different ages. .................................................................................. 27 2.1 Number of carnivores and omnivores at each Gosner developmental stage measured for analyses.................................................................... 35 2.2 T-test of log (base 10) transformed trait measurements at each Gosner developmental stage................................................................................ 38 2.3 Linear regressions of log-transformed traits on log snout-to-vent length and results of ANCOVA comparing slopes of traits plotted against snout-to-vent length in the carnivore and omnivore phenotypes of spadefoot toads................................................................ 40 vi 3.1 a Carnivore OH myofiber number ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. b Omnivore OH myofiber number ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. c T-test of OH myofiber number at each day.......................................... 56 3.2 a Carnivore TL myofiber number ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. b Omnivore TL myofiber number ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. c T-test of TL myofiber number at each day........................................... 58 3.3 a Carnivore OH peripheral myofiber diameter ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. b Omnivore OH peripheral myofiber diameter ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. c T-test of OH peripheral myofiber diameter at each day........................ 60 3.4 a Carnivore OH internal myofiber diameter ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. b Omnivore OH internal myofiber diameter ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. c T-test of OH internal myofiber diameter at each day........................... 62 3.5 a Carnivore TL peripheral myofiber diameter ANOVA and table of post-hoc, bonferroni-corrected, pairwise comparison probabilities. b Omnivore TL peripheral myofiber diameter
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